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RE 1995 ISSN 0035 - 418X
7.
Pas
ZOOLOGIE
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REVUE SUISSE DE ZOOLOGIE
REVUE SUISSE DE ZOOLOGIE
TOME 102 — FASCICULE 3
Publication subventionnée par l'Académie suisse des Sciences naturelles
et la Société suisse de Zoologie
VOLKER MAHNERT
Directeur du Muséum d'histoire naturelle de Genève
FRANCOIS BAUD
Conservateur au Muséum d'histoire naturelle de Genève
DANIEL BURCKHARDT
Chargé de recherche au Muséum d'histoire naturelle de Genève
Comité de lecture
Le président de la Société Suisse de Zoologie
Le directeur du Muséum de Genève: Volker MAHNERT — Systématique des
vertébrés — Muséum de Genève
Le président du comité: Ivan LOBL — Systématique des Insectes — Muséum de
Genève
Patrick GUERIN — Physiologie et éthologie des arthropodes — Institut de Zoologie,
Neuchatel
Willy MATTHEY — Ecologie, entomologie — Institut de Zoologie, Neuchatel
Claude MERMOD — Ethologie et écologie des vertébrés — Université de Neuchatel
Paul SCHMID-HEMPEL — Ecoéthologie, biologie des populations — Institut f.
Terrestrische Okologie, ETH Ziirich, Schlieren
Steve STEARNS — Biologie de l'évolution — Institut f. Zoologie, Basel
Beat TSCHANZ — Ethologie des Vertébrés — Zoologisches Institut, Bern
Claude VAUCHER — Systématique des Invertébrés — Muséum de Geneve
La préférence sera donnée aux travaux concernant les domaines suivants: Biogéographie,
systématique, écologie, éthologie, morphologie, et anatomie comparée, physiologie.
Administration
MUSEUM D'HISTOIRE NATURELLE
1211 GENEVE 6
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(en francs suisses)
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REVUE SUISSE DE ZOOLOGIE, 102 (3) : 539-552; septembre 1995
Oriental species of the genus Biroina Richards
(Diptera: Sphaeroceridae)
Laszl6 PAPP
Department of Zoology, Hungarian Natural History Museum,
Baross u. 13, Budapest, H-1088 Hungary.
Oriental species of the genus Biroina Richards (Diptera: Sphaeroceri-
dae). - Biroina burckhardti sp. n. (Thailand), B. orientalis sp. n. (Pakistan,
India, Nepal) and B. topali sp. n. (India) are described. The genus Biroina
is reported from the Oriental Region for the first time. Synapomorphies
(especially those of the male genitalia) for the species are summarized and
the new species are compared with the type-species B. myrmecophila
(Knab & Malloch, 1912).
Key-words: Diptera - Sphaeroceridae - Biroina - Taxonomy - Oriental
region.
INTRODUCTION
Dupa (1925) described Biroella as a monotypic subgenus of Leptocera based
on Limosina myrmecophila Knab & Malloch, from New South Wales. That name is
homonymous, so RICHARDS (1973) gave a new name for Biroella Duda, and described
13 new species from Australia and one from New Zealand. MARSHALL (1989) ele-
vated its rank to genus and listed all the 15 species, although the generic attribution of
some of them is questionable (Marshall, pers. comm.). The species presently placed in
Biroina would need a revision. In the course of our studies on sphaerocerid flies, three
species of Biroina were found also in the Oriental Region (see ROHACEK & PAPP
1988). Below these three species are described with a re-description of the type-
species, B. myrmecophila. It is hoped that the present paper could be regarded as a
preparatory step for a revision of the genus Biroina.
The morphological terminology of ROHACEK & PApp (1988) is followed here.
The types and other material are deposited in two museums: MHNG —
Muséum d'histoire naturelle Genève (Switzerland); HNHM — Hungarian Natural
History Museum, Budapest (Hungary).
Manuscript accepted 07.10.1994.
540 LASZLO PAPP
TAXONOMY
Biroina Richards, 1973
Biroella DupA, 1925: 74; type-species: Limosina myrmecophila Knab & Malloch, 1912 (by
monotypy).
Biroina RICHARDS, 1973: 330, new name for Biroella Duda, 1925, a junior homonym of
Biroella Bolivar, 1903, as a subgenus of Leptocera Olivier, 1813. Type-species:
Limosina myrmecophila Knab & Malloch, 1912.
Biroina Richards: MARSHALL, 1989: 602, as genus.
Based on the findings published below, the genus Biroina is defined as follows
(possible synapomorphies are indicated by an asterisk, characteristics as given by
ROHACEK (1982) and ROHACEK & PApp (1988): (1) pvt small or indistinct but occe and
occi rather long; (2) 2 ors; (3) (3)-4-5 ifr; (4) frons with or without dark M-shaped
mark; (5) 1-2 dc, if 2, anterior always short; (6) 8-12 tows of ac microsetae; (7) 1
katepisternal; (8) scutellum flat, semicircular or even broader; (9) male fore and hind
femora not much thickened; (10)* t> chaetotaxy (dorsal side) characterized by 2 pairs
of proximal bristles (though upper proximal ad more proximal than its pd pair in
some species) and 3 very long distal bristles; ventral side without mid ventral seta;
(11) wing with C not extending beyond apex of R445; (12) R243 strongly sinuate; (13)
discal cell short and broad, usually with extensions of M),> and M3,4, one of them or
both approximate wing margin in some species; (14) alula small and narrow; (15)
preabdomen with large, heavily sclerotized terga, preabdominal sternites rather large
and well sclerotized; (16) male sternite 5 without additional structures; (17)
posteromedial part of sternite 5 less sclerotized with short setae; (18) epandrium
(periandrium) mostly semiglobular; (19)* hypandrium without robust bifurcate
ventral appendage but with a slipper-shaped cranioventral process, its ventral surface
with or without small teeth; (19)* hypandrium with a caudally directed apodeme (Figs
20, 29, cf. Fig. 61 of ROHACEK & Papp 1988, Fig. 32 of PAPP 1991); (20) male subanal
plate distinct; (20a)* epandrium lateroventrally usually with a pair of lamelliform
processes (Figs 22, 26), which are not fused with subanal plate, i.e. its derivation from
subanal plate is questionable; (21) surstylus (gonostylus) bilobed, lateral lobe with
numerous long to extremely long setae, medial lobe various; (22)* phallophore long
digitiform or very long; (23) distiphallus complex with dorsal appendages; (24)
postgonite of various shape but always simple; (25) ejaculatory apodeme well
developed; (26)* female postabdomen fully telescopized and hidden in segment 5
when at rest; (27) female 76 and 77 divided into two, also S6 and S7 weakly
sclerotized though they may be broad (e.g. in orientalis); (28)* female tergite 8
divided into 4 long narrow sclerites; (29) tergite 10 very short; (30) female sternite 8
narrow and weakly sclerotized; (31) sternite 10 small; (32) spectacles-shaped sclerites
weakly sclerotized; (33) accessory glands not studied now; (34) spermathecae pear-
shaped or spherical; (35) female cerci with or without sinuate hairs.
ROHACEK's (1982, 1983) revision of the limosinine genera, though its was not
aimed at a complete revision of the World fauna, gave enough basis also for those
genera not considered there to have them judged for their generic identity. Based on
ORIENTAL SPECIES OF BIROINA 541
the synapomorphies hitherto considered, I think, Biroina deserves a generic rank
indeed.
Remark. The genus Apterobiroina L. Papp, 1979 is not related to the genus
Biroina as originally stated by PAPP (1979) but possibly to Minilimosina Rohacek,
1983 (this will be discussed in a future paper).
Biroina burckhardti sp. n.
Holotype d (MHNG): Thailande, Chang Mai, Doi Suthep, 1400 m, 5.X1.1985, D.
Burckhardt - I. Löbl (No. 11).
Measurements in mm: body length 2.79, wing length 2.05, wing width 0.90.
Body dark greyish brown, mesonotum subshining, pleura with thick grey
microtomentum.
Frons light brown, face, genae and occiput light reddish brown. M-shaped
mark, including orbits (fronto-orbital plates), interfrontalia, ocellar triangle and the
sagittal line of frontal vitta silvery. Frontal lunule small, triangular, gena below eye at
narrowest 0.175 mm and strongly widening posteriorad. Cephalic chaetotaxy: pvt
minute and divergent; 2 ors but anterior one reduced to a short thin hair (1/4 length of
posterior ors); ocellars large; 4 pairs of short and thin ifr. Eyes mangoe-shaped.
Antennae large, antennal length 0.29 mm, pedicel reddish brown, flagellomere
conical, light dirty red. Arista 0.725 mm with short cilia.
Thoracic chaetotaxy: | h, 2 superposed np, 2 very long sa, 1 pa. Two dc pairs
but anterior one less than half length of the posterior one. Femora light, reddish,
apices brown, tibiae dark brown, basitarsus and 2nd tarsomere of fore tarsi brown,
tarsomeres 3-5 white; mid and hind tarsomeres dirty yellow. 1 chaetotaxy: two short
anterals at 23/38 and 28/38; ad-s: very short at 7/38, short at 9/38, long at 15/38, very
long at 31/38 of tibia; pd-s: medium long at 9/38, long at 18/38, very long at 30/38 of
tibia. Wings light browb, veins dark brown. R| extremely short and thick, upcurving
to costa at the level of the tip of alula, R4,5 upcurving to costa on a wide arc, R743
very short and sinuate, approximates costa at its middle, distance in its apical third to
C is little more than the diameter of C. mg2/mg3 0.34 mm to 0.775 mm (wing curved,
i.e. not precisely measurable). M;,2 and M3,4 short, dark brown, former one parallel
to R445. lat) 0.207 mm, tp 0.138 mm (i.e. ratio 1.50). Halteres waxy white.
Preabdomen dull black, tergites with medium long and not thick discal and
marginal bristles. Syntergite 1+2 nearly as long as tergite 3 and 4 combined. Male
sternite 5 short, posteromedially with acute short setae (Fig. 2).
Genitalia: periandrium rather high (Fig. 1), slightly asymmetrical. Subanal
plate is an intricate structure: its main part is a pair of wide caudal plates (Fig. 4)
enlarged ventrally to the base of medial lobe of surstylus. Medial lobe of surstylus
(Figs 4, 5) long, digitiform with minute setae only, lateral lobe with several extremely
long setae. Aedeagal complex with comparatively short digitiform phallophore (Fig.
3). Postgonite (Figs 3, 6) geniculate, its basal part much shorter than its apical part.
Ejaculatory apodeme distinct though small. Hypandrium strongly asymmetrical, its
medial anterior part wide and well sclerotized.
542 LASZLO PAPP
Fics 1-3
Terminalia of Biroina burckhardti sp. n., holotype male; 1. epandrium, hypandrium, cercus and
surstylus laterally, 2. sternite 5 and 6, 3. aedeagal complex laterally. Scale bars: 0.2 mm.
ORIENTAL SPECIES OF BIROINA 543
Fics 4-6
Terminalia of Biroina burckhardti sp. n., holotype male; 4. genitalia in subventral (subcaudal)
view, 5. surstylus laterally, 6. postgonite laterally. Scale bars: 0.1 mm, 0.2 mm (Fig. 4).
Female unknown.
Biroina burckhardti sp. n. may be readily distinguished from the other Oriental
species of the genus (see key); it is not closely related to the other two species.
Etymology. This species is dedicated to Dr. Daniel Burckhardt (MHNG), one
of the collectors of the holotype.
Biroina myrmecophila (Knab & Malloch)
Limosina myrmecophila KNAB & MALLOCH, 1912: 236.
Material studied: 1 4 5 2 (HNHM): Australia Birö 1900. — N.S. Wales, Mt. Victoria
(on the reverse side: "XI/12/") — "Biroella d 2: myrmecophila" Det. Dr. O. Duda.
Measurements in mm: body length 2.50-3.30, wing length 2.17-2.71, wing
width 1.09-1.31.
544 LASZLO PAPP
Fics 7-12
Male terminalia of Biroina myrmecophila (Knab & Malloch); 7. male sternite 5, 8. postero-
medial part of sternite 5 in higher magnification, 9. surstylus in ventral view, 10. main (medial)
part of hypandrium, 11. aedeagal complex with a part of the subanal plate, 12. epandrium,
cercus and surstylus laterally. Scale bars: 0.1 mm (Figs 8, 9), 0.2 mm (Figs 7, 10-12).
ORIENTAL SPECIES OF BIROINA 545
Body dark brown subshining.
Frons, face and genae reddish brown, frons without M-shaped mark. Frontal
lunule subtriangular, gena as broad as width of eye, with very strong genal bristle
below eye. Cephalic chaetotaxy: pvt minute but occe and occi very long; 1 short
anterior and 3 very long pairs of ifr. Eyes small. Antennae reddish yellow, pedicel
with extremely long btistles, flagellomere fuscous apically. Arista somewhat longer
than width of head with very short cilia.
Thorax: mesonotum not granulated but very thinly microtomentose. Two dc,
anterior one much shorter. Legs dark brown, tarsomeres somewhat lighter. f
chaetotaxy: ad-s: short at 12/49, long at 19/49, very long at 40/49; pd-s: short at
12/49, long at 19/49, very long at 40/49, a very long and thick anteral at 37/49. Wings
tesselate with darker diffuse dark spots on lighter basic colour, veins light brown.
Proximal section of C with very long setae. R343 much sinuate (strongly S-shaped, see
Fig. 28 of RICHARDS 1973). R4,5 slightly upcurving. Mj+> and M344 distinct, former
one approximates wing margin. ta-tp as long as or even shorter than hind crossvein.
Halteres brown.
Male preabdomen with broad ans subshining terga, which bear medium long
marginal and lateral setae, also sterna broad. Male sternite 5 posteromedially with
rather long inclinate acute bristles (Figs 7, 8).
Male genitalia: Epandrium (periandrium) not high (Fig. 12). Subanal plate
heavily sclerotized. Male cerci well-sclerotized with long setae (Fig. 12). Surstylus
(gonostylus) with long lateral lobe which bears several long setae (Figs 9, 12), medial
lobe very large and wide with numerous medium long but thick setae. Aedeagal
complex with phallophore (Fig. 11) very long, proclinate and very thin apically. Post-
gonite (Fig. 11) rather broad in its basal half, tapering distally, apical part geniculately
bent, apex blunt. Ejaculatory apodeme small but distinct.
Female preabdomen as in male; postabdomen strongly telescopized. Tergite 8
in 4 parts (Fig. 14), sternite 8 long quadratic, sternite 10 in two parts. Cerci compara-
tively large (broad) with 3 pairs of very long and several shorter setae. Spermathecae
(Fig. 13) pear-shaped, their sclerotized duct long, basally with small round mamillae;
sclerotized duct distally with a small swelling.
Biroina myrmecophila (Knab & Malloch, 1912), the type-species of the genus,
is markedly different from the three Oriental species (see key below). However, the
generic characteristics (synapomorphies) listed above may serve as criteria for rele-
gation to this genus when studying other species known as Biroina. RICHARDS (1973)
although he did not see the holotype, depicted the same species as I do (cf. his Figs
40, 41); our specimens are from a site not far from the type-locality.
Biroina orientalis sp. n.
Holotype d (MHNG): India, Uttar Pradesh, Kumaon, Chaubattia, 12-13.X.1979, I.
Löbl (No. 10).
Paratypes: 4 4,3 2 (MHNG, HNHM): same data as for the holotype; 5 4,4 £ (MHNG,
HNHM, 1 d damaged, left wing lost): ibid., Ramgarh, 2000 m, O. Löbl, 9.X.1979 (No. 6/b.); 1
2 (MHNG): India, Uttar Pradesh, Gharwal, Dhanolti, I. Löbl (No. 19): 1 d, 1 2 (HNHM): India,
546 LASZLO PAPP
Fics 13-17
Female terminalia of Biroina spp. 13-14. B. myrmecophila (Knab & Malloch); 13. sperma-
theca, 14. postabdomen in ventral view. 15-17. B. orientalis sp. n., paratype female; 15. post-
abdomen in lateral view, 16. same, dorsal view, 17. spermathecae. Scale bars: 0.1 mm (Figs 13,
17), 0.2 mm (Figs 14-15, 16).
ORIENTAL SPECIES OF BIROINA 547
Uttar Pradesh, Nainital, Sleepy Hollow, ca. 2080 m — sifted and singled from under stones,
4.XII.1989, leg. L. Papp; 1 d (MHNG): Pakistan, Swat, Miandam, C. Besuchet-I. Löbl,
17.V.1983 (No. 15/b.): 1 d (MHNG): ibid., 10.V.1983 (No. 6c.); 1 gd, 2 2 (MHNG): Népal,
Bagmati, Malemchi, 2800 m, 14.IV.81, I. Löbl-A. Smetana (No. 24); 1 2 (MHNG): Népal, Prov.
Bagmati, Tarke Ghyang, 2750 m, 19.IV.81, I. Löbl (No. 32a.); 1 9 (MHNG): Népal: Kosi,
Induwa Kola, 2100 m, 17.IV.81, I. Löbl-A. Smetana (No. 27); 1 6,2 2 (MHNG, 1 2 damaged,
left wing lost): Népal, Bagmati, Gul Bhanjyang, 2600 m, 6.IV.81, I. Löbl-A. Smetana (No. 9).
Measurements in mm: body length 2.21 (holotype); 1.67-2.58 (paratypes),
wing length 1.85 (holotype), 1.50-1.96 (paratypes), wing width 0.88 (holotype),
0.71-0.96 (paratypes).
Body dark brown, mesonotum dull, abdomen subshining dorsally.
Frons dark reddish brown, orbitalia, vertex, interfrontal stripes and a sagittal
line silvery microtomentose, so M-shaped mark well discernible. Frontal lunule
subtriangular, light brown, gena bare in its dorsal two-thirds, face shining. Cephalic
chaetotaxy: pvt small, occe and occi pairs comparatively long though thin; 2 ors close
to each other; four pairs of ifr bristles; genal bristle less strong than in myrmecophila.
Eyes not much reduced, gena below eye less broad than width of eye. Antennae long,
reddish yellow, pedicel comparatively very long with very long bristles subapically,
flagellomere long and pointed dorsoapically. Arista longer than width of head with
medium long cilia.
Thorax dark brown with thick grey micromentum on mesonotum. Two dc,
anterior one only half as long as posterior; | strong katepisternal. Legs dark brown but
fore tarsomeres 2-5 whitish yellow, mid and hind tarsomeres 2-5 yellow. # chaeto-
taxy: ad-s: short at 1/3, longer at 14/33, strong at 16/33, strong at 29/33 of tibia; av-s:
short at 7/33, long at 17/33, very strong at 26/33 of tibia. Wings light brown without
tesselate pattern, veins brown but not dark. R2,3 strongly S-curved but runs close to
costal vein in its whole length, R4,5 upcurving to C along a wide arc. M;,35 and M3,4
rather short, not approximating wing margin. ta-tp as long as hind crossvein. Knob of
halteres long and brown, stalk waxy yellow.
Male preabdomen dull black, tergites with medium long and not thick discal
and marginal bristles. Sternite 1 reduced to a pair of minute round sclerites. Male
sternite 5 posteromedially with blunt and somewhat flattened short setae (Fig. 19).
Male genitalia: periandrium only a little higher than long. Subanal plate similar
to that of topali but lateroventral process of epandrium (Gig. 22) with less broad base.
Ventral surface of slipper-shaped ventral process of hypandrium without small teeth
(Fig. 20). Surstylus (gonostylus) with lateral lobe with numerous very long setae on
its lateral surface but no long bristles in its cranial (apical) 2/5, medial lobe of
surstylus (Fig. 21) lobate, with longer setae. Aedeagal complex with long digitiform
phallophore. Postgonite (Fig. 23) curved in distal third, its basal part much longer
than its apical part. Ejaculatory apodeme bacilliform.
Female preabdomen similar to that of male; postabdomen fully telescopized.
Tergites 6 and 7 divided into two, tergite 8 into four (Figs 15, 16), but sternite 6 to 8
not divided; tergite and sternite 10 minute, partly hidden under tergite and sternite 8
(Fig. 15). Cerci short and pale yellow with comparatively short setae. Spermathecae
sphaerical, their sclerotized duct short (Fig. 17).
548 LASZLO PAPP
Fics 18—23
Terminalia of Biroina orientalis sp. n., paratype male; 18. sternite 5 with a part of sternite 6, 19.
posteroventral part of sternite 5 in a higher magnification, 20. hypandrium in ventral view, 21.
Surstylus in its widest extension, 22. subanal process in its widest, 23. aedeagal complex
laterally. Scale bars: 0.1 mm (Figs 19, 22), 0.2 mm (Figs 18, 20, 21, 23).
ORIENTAL SPECIES OF BIROINA 549
Biroina orientalis sp. n. is closely related to B. topali sp. n.; differentiating
characteristics are given in the key.
Etymology. Derived from its distribution.
Biroina topali sp. n.
Holotype ¢ (HNHM): India, No. 925, Daitari, Orissa, 600-700 m — erdei avarböl
[from forest litter], 1967.X1.23, leg. [György] Topal.
Measurements in mm: body length 2.96, wing length 2.18, wing width 1.05.
Body mainly dark brown, thinly microtomentose, i.e. not shining.
Frons rather flat and as head, very broad, though somewhat longer than wide.
M-shaped mark distinct, ocellar triangle, vertex and occiput dark brown, orbits dark
only to the base of ors, anterior parts of orbits, interfrontal and sagittal stripes
yellowish silvery dusted, other parts of frons light brownish yellow. Frontal lunule
triangular. Gena reddish yellow with honey shine, only peristomal area dusted with
silvery microtomentum, gena at genal bristle 0.216 mm wide and strongly widening
posteriorad. Face bright ochreous. Cephalic chaetotaxy: pvt minute, occe and occi
rather long; 2 ors close to each other; 5 ifr comparatively long but thin, genal bristle
long, ca. 3/5 of vibrissa in length, vte, vti and oc strong (ocellars lost but must be
strong judged by their bases). Eyes long elliptical, longitudinal axis rather oblique,
length/width 0.40/0.26 mm. Antennae light, reddish yellow, only scape brown, the
latter with a long ventrally directed thick hair, pedicel with 6 very long and thick
setae, flagellomere conical with long silvery cilia. Arista medium long with long
hairs.
Thorax: mesonotum flat with numerous (up to 12) rows of small hairs. Two dc
pairs but anterior pair less than half length of posterior one; only 1 katepisternal
(anterior pair reduced to a small hair, which is less than 1/3 in length of the posterior
one). Scutellum flat and very broad, nearly semicircular, 0.33 mm long, 0.585 mm
wide (scutellars broken off). Legs brown, 3rd to Sth tarsomeres of fore tarsi white
(yellowish white), mid and hind tarsi yellow. # chaetotaxy: anterodorsals: two small
ones at 7/39 and 9/39, one strong at middle, small at 30/39, one very strong and long
at 33/39, 1 medium long dorsally directed and rather anteral one at 30/39;
posterodorsals: strong at 8/39, very strong at 28/39, very long and thick at 32/39; no
mid ventral but a row of thick and short setae along distal 3/4 of tibia; apicoventral
thick but only 0.05 mm long. Wings definitely brown, veins dark brown. R| very
thick and short, R2,3 strongly S-shaped (sinuate), runs close to C from its basal
quarter, R4,5 upcurving to C along a wide arc. Mı,> distinct on a section of 0.38 mm
only, i.e. diminishes halfway from tp to wing margin. M3,4 distinct on a section of
0.095 mm only, duscal cell very broad. f-tp only slightly longer (0.190 mm) than #,
(0.155 mm). Halteres rather small, knob ochreous, stalk waxy yellow.
Preabdomen comparatively long and flat without any conspicuous setae.
Sternite 5 posteromedially with blunt and somewhat flattened short setae (Fig. 25).
550 LASZLO PAPP
LAPS
EN AO
u N) NL
Fics 24-30
Terminalia of Biroina topali sp. n., holotype male; 24. sternite 5 with a part of sternite 6, 25.
posteroventral part of sternite 5 in a higher magnification, 26. ventral part of epandrium with
left subanal process in its widest, 27. surstylus in its widest extension, 28. aedeagal complex
laterally, 29. hypandrium in ventral view, 30. male genitalia laterally. Scale bars: 0.1 mm (Fig.
26), 0.2 mm (Figs 24, 25, 27-29, 30).
ORIENTAL SPECIES OF BIROINA 551
Genitalia: epandrium (periandrium) not high (only a little higher than long),
ventrocaudal part of epandrium with a pair of subanal processes (Fig. 26), which bear
1 long and several shorter setae. Lateroventral process of epandrium (Fig. 26) with
broad base. Ventral surface of slipper-shaped ventral process of hypandrium with
small teeth, serrate backwards (Fig. 29), hypandrial apodeme stronger than in
orientalis. Lateral lobe of surstylus (gonostylus) with very long bristles also in its
apical third, medial lobe of surstylus (Fig. 27) lobate, with setae longer than in
burckhardti. Aedeagal complex with long digitiform phallophore. Postgonite (Fig. 28)
curved in distal third (somewhat more curved than in orientalis), its basal part much
longer than its apical part. Ejaculatory apodeme rather long but thin.
Female unknown.
Biroina topali sp. n. is closely related to B. orientalis but the holotype is bigger
than any of the orientalis specimens measured, it possesses 5 pairs of ifr bristles
(instead of 4) and it differs in some details of the male genitalia.
Etymology. The species is dedicated to Dr. György Topäl (HNHM) in
recognition of the invaluable materials collected by him in Argentina, India, Vietnam,
etc.
KEY TO THE ORIENTAL SPECIES OF Biroina
l Wings tesselate with darker diffuse dark spots on lighter basic colour.
Male cerci well-sclerotized with long setae (Fig. 12) Male phallophore
(Fig. 11) very long, proclinate and very thin apically. Postgonite (Fig.
11) rather broad in its basal half. Female spermathecae pearshaped with
longssclerotizediducr(Rie 13) e [myrmecophila (Knab & Malloch)]
- Wings light brown, veins darker brown. Male cerci weakly sclerotized
with short hairs only. Male phallophore shorter, digitiform (e.g. Fig.
23). Postgonites (Figs 3, 23, 28) less broad. Female spermathecae sphe-
rical with short duct (Fig. 17; females of burckhardti and topali
unknown) eu, Ne ae 2
2 Sternite 5 posteromedially with acute short setae (Fig. 2). Medial lobe
of surstylus (Figs 4, 5) long, digitiform with minute setae only.
Postgonite (Fig. 6) geniculate, its basal part much shorter than its apical
Pate N en N eo burckhardti sp. n.
= Male sternite 5 posteromedially with blunt and somewhat flattened
short setae (Figs 19, 25). Medial lobe of surstylus (Figs 21, 27) lobate,
with longer setae. Postgonite (Figs 23, 28) curved in distal third, its
basalparsmuchllonser than ıtstapiealpae e 3
3 Four pairs of ifr bristles. Lateral lobe of surstylus with numerous very
long setae on its lateral surface but no long bristles in its apical (cranial)
2/5 (Fig. 21). Ventral surface of slipper-shaped ventral process of
hypandrium without small teeth (Fig. 20). Smaller (body length
LEGS SOI An orientalis sp. n.
552 LASZLO PAPP
— Five pairs of ifr bristles. Lateral lobe of surstylus (gonostylus) with
very long bristles also in its apical third (Fig. 27). Ventral surface of
slipper-shaped ventral process of hypandrium serrate, i.e. with small
teeth (Fig. 29). Bigger (body length 2.96 mm) ................ topali sp. n.
ACKNOWLEDGEMENTS
My most sincere thanks are due to Drs Ivan L6bl and Daniel Burckhardt
(MHNG) for the loan of numerous sphaerocerid material from the Oriental Region.
Jindfich Rohäëek, Silesian Museum Opava is gratefully acknowledged for his
comments.
REFERENCES
Dupa, O. 1925. Die aussereuropäischen Arten der Gattung Leptocera Olivier = Limosina
Macquart (Dipteren) mit Berücksichtigung der europäischen Arten. Arch. Naturgesch.
A 90(11): 5-215.
KNAB, F. & J.R. MALLOCH. 1912. New Australian Diptera from ants' nests. Trans. R. Soc. South
Austr. 36: 233-237.
MARSHALL, S.A. 1989. 96. Family Sphaeroceridae. In: Evenhuis, N.L. (ed.): Catalog of the
Diptera of the Australasian and Oceanian Regions/ Bishop Museum Press and E.J.
Brill, Honolulu, 1155 pp. (pp. 601-607).
Papp, L. 1979. On apterous and reduced-winged forms of the families Drosophilidae,
Ephydridae and Sphaeroceridae (Diptera). Acta zool. hung. 25(3-4): 357-374.
Papp, L. 1991. Oriental Limosininae: new species and records (Diptera, Sphaeroceridae). Acta
zool. hung. 37(3-4): 225-251.
RICHARDS, O.W. 1973. The Sphaeroceridae (= Borboridae or Cypselidae); Diptera Cyclorrha-
pha) of the Australian Region. Austr. J. Zool., Suppl. Ser. 22: 297-401.
ROHACEK, J. 1982. A monograph and re-classification of the previous genus Limosina
Macquart (Diptera, Sphaeroceridae) of Europe. Part I. Beitr. Ent. 32: 195-282.
ROHACEK, J. 1983. ditto, Part II and III. Beitr. Ent. 33: 3-195, 203-255.
ROHACEK, J. & L. Papp. 1988. A review of the genus Paralimosina L. Papp (Diptera,
Sphaeroceridae), with descriptions of ten new species. Ann/s hist. nat. Mus. natn. hung.
80: 105-143.
REVUE SUISSE DE ZOOLOGIE, 102 (3) : 553-583; septembre 1995
New and little known taxa of Rhytidochrotinae
(Acrididae, Orthoptera) from Costa Rica
C. Hugh F. ROWELL
Zoologisches Institut, Universitàt Basel,
Rheinsprung 9, CH-4051 Basel, Switzerland.
New and little known taxa of Rhytidochrotinae (Acrididae, Orthop-
tera) from Costa Rica. - Three new monospecific genera are described
from montane areas of Costa Rica. Like the other three known Costa Rican
rhytidochrotine genera (Hylopedetes Rehn, 1929, Scirtopaon Descamps &
Rowell, 1984 and Micropaon Descamps & Rowell, 1984) they belong to
the atympanate and apterous division of the subfamily. All are apparently
confined to small geographical areas. Exerythracris volcanica n.sp. is a
fern eater from the light gaps and edges of montane forest, and resembles
in gross morphology the similarly specialised Hylopedetes. It is the most
northerly representative of the subfamily so far described, extending almost
to the Costa Rican/Nicaraguan border. Brakeracris varablancensis n.sp. is
found in successional areas (landslips, anthropogenically altered habitats)
in montane forest; its diet is unknown, but does not apparently include
ferns; morphologically it is closest to Scirtopaon. Talamancacris palustris
n.sp. is confined to a few relict subalpine swamps and appears to eat
Asteraceae preferentially; it is not obviously closely allied to the other
genera. Additionally, new morphological, ecological and biogeographic
data are given for the three previously described genera.
Key-words: Orthoptera - Acrididae - Rhytidochrotinae - Costa Rica -
Taxonomy.
INTRODUCTION
Last globally reviewed by DESCAMPS & AMEDEGNATO (1972b), the group
Rhytidochrotinae was elevated to subfamily status by AMEDEGNATO (1975). It
currently includes 17 genera and 35 species (DESCAMPS & AMEDEGNATO 1972b;
DESCAMPS & ROWELL 1978; DESCAMPS & ROWELL 1984), to date known only from
the northern half of South America and from the southern part of Central America
Manuscript accepted 06.11.1994.
554 C.H.F. ROWELL
(i.e., from Brasil to Costa Rica). The greatest number of genera and species is found
in Colombia. With few exceptions (Paropaon, Piezops, Lathacris, Hylopedetes
mirandus) the Rhytidochrotines are insects of montane forest (roughly 1000 m to
2500 m in altitude) and are patchily distributed. Some genera are behaviourally
specialised to eat ferns (ROWELL er al. 1984). They are often brilliantly coloured,
especially the males, sometimes in unusual shades of red.
The formal morphological characteristics of the subfamily were listed by
DESCAMPS & AMEDEGNATO (1972b) and AMEDEGNATO (1977). All genera but one
(Driphilacris Descamps & Amédégnato, 1972a) are completely apterous, and about
half of them are also atympanate. The pronotum is characteristically short, exposing
the entire meso- and metanota, and traversed dorsally by three sulci; the first of these
is short with ends curved anteriorly, almost joining a further groove running just
behind margin of pronotum, which is obsolete dorsally (Lathacris Descamps &
Amédégnato, 1972a has only 2 transverse sulci). Males have a "furcula" (a notched
and usually toothed posterior margin of the 10th abdominal tergite). Similar pronota
and furculae are however seen in several Ommatolampine genera, e.g. Tamnacris
Descamps & Amédégnato, 1972a. The male genitalia are characteristic and homo-
genous; the aedeagus is extremely short and completely sheathed, with very reduced
inferior sclerites and very short superior ones. The only part of the male genitalia that
is at all useful in distinguishing genera is the epiphallus (especially the shape and
orientation of the lophi). The ventral ovipositor valves are typically slender distally
and along their dorsal edge and lie largely between the dorsal valves. In the Central
American genera both lower and upper valves are prominently toothed on their outer
edges, though this is denied in previous characterisations of the subfamily. In all the
genera I have examined Collins’ tooth (COLLINS, 1991) is present on anterior margin
of the third antennal segment from distal end.
The first rhytidochrotine genera described from Central America were Piezops
Hebard, 1923 from southern Panama and Hylopedetes Rehn, 1929 from central Costa
Rica. Hylopedetes originally included three species; DESCAMPS & ROWELL (1978) des-
cribed 2 additional species, bringing the total to 5. Scirtopaon and Micropaon, also
from Costa Rica, were added by DESCAMPS & ROWELL (1984). The present article
describes three new genera from the uplands of Costa Rica, and provides additional
morphological drawings and new localities for the previously described genera and
species. A new artificial key to the atympanate genera of the Rhytidochrotinae (all the
known Costa Rican taxa fall into this category) is provided. For the remaining
(tympanate) genera the corresponding section of the key given by DESCAMPS &
AMEDEGNATO (1972b) remains valid.
MATERIALS AND METHODS
In addition to the new taxa described, I had access while preparing this paper
to material of the the following Rhytidochrotine genera: Galidacris; Hylopedetes;
Micropaon; Opaonella; Parapiezops; Paropaon; Piezops; Rhytidochrota; Scirtopaon;
Trichopaon. The other genera are known to me only from the literature.
COSTA RICAN RHYTIDOCHROTINAE 555
Dimensions were measured with a graticule eyepiece at a total magnification
of X25 (Wild MS stereomicroscope) in conjunction with a moving stage fitted with a
digital micrometer (Mitutoyo) reading to 0.01 mm. Measurements were repeatable to
within 0.02 mm. The length of the pronotum was measured in the midline; as the
posterior and anterior margins of the pronotum are mostly notched in midline in the
Rhytidochrotinae, the measurements are from the bottom of these notches. Overall
length is from the most anterior part of the frontal ridge to the posterior end of the
subgenital plate (males) or the extremities of the ovipositor valves (females). The size
of the rostrum is expressed as the ratio of the two distances illustrated in Fig. 1.
FIG. 1
Derivation of the rostrum index used in the text and Table 4. On a drawing of a side view of the
head (here of Hylopedetes mirandus) a line is drawn from P, the junction of the outline of the
compound eye and the upper surface of the fastigium, to Q, the notch of the frontoclypeal
suture. À further line YZ is then drawn normal to PQ, passing through the furthest extremity of
the rostrum, Y. The rostrum index is the ratio YZ / PQ.
Drawings of the male internal genitalia were made with a drawing tube fitted
to a Leitz compound microscope, and of other parts via tracings from projected colour
transparencies photographed with a Wild Photomacroscope.
Abbreviations of museums: ANSP, Academy of Natural Sciences, Phila-
delphia; INBio, Instituto Nacional de Biodiversidad, Santo Domingo de Heredia,
Costa Rica; MNHNP, Muséum National d'Histoire Naturelle, Paris; RC, the author's
collection, University of Basel.
556
C.H.F. ROWELL
KEY TO THE ATYMPANATE GENERA OF THE RHYTIDOCHROTINAE (Males only)
1
>|
Un
6 external spines on hind tibia (S. America).
LI GIS CIT El a SENDER ae Lathacris Descamps & Amédégnato, 1972a
Waextemalispines/onmhind tibia: 2% 5.426. . 2022 ar ee 2
Supraanaliplate Withoutblack Spots CT CCE 3
Supraanalsplateswithiblackspots. EE 2222022. 2 CSP PC RECENSE 8
Interocular space more than 3X wider than frontal ridge (S. America).
D'ORO AI E SEA Wnts, ey i) ES AGI I ashe seas Muyscacris Hebard, 1923
Interocular space less than 3X wider than frontal ridge................... 4
Medial carina of pronotum at least partially present. .................... I
Medialicanmajor pronotum/absent:. ©... it ERRE 6
Medial carina thin but entire, cut by sulci (Fig. 9B).
ese re IA SE An es Scirtopaon Descamps & Rowell, 1984
Medial carina of pronotum present only in front of Ist sulcus and
belimd8rdisulcust(Ri AO Brakeracris n. gen.
Dorsal apical spine of hind knee longer than outer knee lobe (Fig. 6G).
Paired tufts of sparse white hairs dorsolaterally on abdominal and tho-
FACICISC CAM EMSS aes LITI TIZI Talamancacris n. gen.
Dorsal apical spine of hind knee equal to or shorter than outer knee lobe. ... 7
Meso- and metanota smooth, "epaulettes" (see p. 7) obsolete, anterior
pronotal sulcus shallow (Figs. 8A & B); knee spine much shorter than
outeriknceil0be (BID) yes eka eet enone Hylopedetes Rehn, 1929
Meso- and metanota heavily pitted, "epaulettes" present, anterior pro-
notal sulcus deep and well marked (Figs. 2B & C). Dorsal apical spine
of hind knee as long as outer lobe (Fig. 2G). .......... Exerythracris n. gen.
Frontal ridge extends to clypeus. Face flat. Medial carina of pronotum
present, cut by only 2 sulci. Supraanal plate with 2 black spots (S.
AMEN CAI rege MINARE Lathacris Descamps & Amédégnato, 1972a
Frontal ridge absent below medial ocellus. Face concave. Medial carina
Of pronotumeabsent on very indistinct) 7.22 Sean 9
Supranal plate with 2 black spots. Points of furcula long and sharp (S.
PANTING CA) RARES. PIO SATO LI Paropaon Hebard, 1923
Supranal plate with 1 medial black spot. Points of furcula short and
blunt. Posterior margins of meso- and metanota thickened and raised in
DATE see ee Micropaon Descamps & Rowell, 1984
DESCRIPTIONS OF NEW TAXA
Exerythracris n. gen.
Type species: Exerythracris volcanica n. sp.
COSTA RICAN RHYTIDOCHROTINAE 557
Male. (Figs. 2 & 3). Size small-medium (10mm<body length<20 mm). Gene-
ral form cylindrical, slender, long legged. Integument polished; dorsal surfaces of
thorax, occiput, hind femora, and to a lesser extent of abdomen, heavily pitted.
Fastigium steeply inclined forwards, tapering, bearing a pair of low longi-
tudinal ridges proximally, distally shallowly grooved medially, the groove not
reaching the anterior margin of the fastigium (Fig. 2B, D). Frontal ridge narrow, not
wider than interocular space, narrow at top and bottom, slightly wider opposite the
upper margin of the antennal foramen, ending and diverging slightly above the very
small medial ocellus (Fig. 2E). Profile of frons markedly concave, the frontal ridge
forming a rounded rostrum, index 0.33 (Fig. 2B). Subantennal sutures thin and rather
indistinct, the ventral sutures short and projecting laterally; ventral margin of frons at
frontoclypeal suture distinctly concave (Fig. 2E). Preocular ridges of frons present.
Eyes large and globose; interocular space narrow, 0.6 - 0.8X as wide as greatest width
of antennal scape. Antennae long, filiform, slender, round in cross section, 17
segments in flagellum of adult, end of terminal segment obliquely pointed.
Prothorax (Fig. 2B,C) without medial carina. Transverse sulci deep. Dorso-
lateral area between 2nd and 3rd sulci inflated into a prominent bulge, bearing a
matte-surfaced oval depression (this structure is characteristic of many rhytido-
chrotine genera, and is here referred to as an "epaulette" - see also the description of
Loepacris, DESCAMPS & AMEDEGNATO 1972b: 1068 & their Fig. 20). Posterior ventral
angle of lateral lobe of pronotum rounded, posterior margin above the ventral angle
straight. Anterior ventral angle of lateral lobe of pronotum obtusely rounded. Both
anterior and posterior margins of pronotum slightly notched in midline. Prosternal
process short, abruptly conical, vertical or slightly angled forwards. Apterous. Pos-
terior margins of meso- and metanotum somewhat raised, giving a saw-tooth dorsal
profile to thorax in lateral view.
Hind femora very long (4.3X as long as pronotum) and slender; outer face of
femur rounded. Dorsal carina of femur terminating in a short spine, which equals but
does not not exceed the external upper lobe of the knee (Fig. 2G). Seven non-terminal
spines on the dorsal external margin of hind tibia, 8 on the dorsal internal. Hind tarsi
long, 0.34X as long as hind femur and 1.45X as long as pronotum. Third tarsal
segment 1.5X as long as the first and second segments together. Claws long and
widely separated.
Tympanum absent. Posterior margin of 10th abdominal tergite with a central
notch and short lateral sclerotised teeth (Fig. 2F). Supraanal plate triangular, without
sclerotised melanic markings, slightly rounded at apex and with a slight medial
furrow at its base (Fig. 2F). Cerci short, conical, simple. Subgenital plate smoothly
rounded, in dorsal view terminating in a short medial furrow. Internal genitalia (Fig.
3) of the type typical for the subfamily.
Female. Size, medium (20mm<body length<30 mm), 1.3-1.4X as long as the
male. Differs from male as follows:
Integument less polished, more extensively pitted than in male. Interocular
space slightly wider than in male, 0.9X as wide as antennal pedicel. Antennae slightly
flattened dorsoventrally and relatively shorter than in male (3.1X pronotum).
558 C.H.F. ROWELL
FIG. 2
Exerythracris volcanica n.sp. Male. A. Lateral view of entire insect. B. Lateral view of head
and pronotum. C. Dorsal view of pronotum. D. Dorsal view of head. E. Frontal view of face. F.
Dorsal view of furcula, supraanal plate and cerci. G. Lateral view of hind knee.
COSTA RICAN RHYTIDOCHROTINAE 559
Fusiform in shape, posterior margin of pronotum, meso and metathorax, and the
anterior segments of abdomen being relatively wider than in male. Pronotum with the
merest suggestion of a median carina, defined more by local absence of pitting than
by a raised ridge. Hind tarsus and femur relatively shorter than in male, the femur
3.9X as long as pronotum. Third segment of hind tarsus 1.4X length of Ist and 2nd
segments together. Genitalia (Fig. 3H-K) typical for the subfamily. Spermatheca not
examined.
Diagnosis. The dimensional comparison with other taxa is shown in Table 4.
Exerythracris has one of the proportionately longest hind femora (4.3X pronotum)
and both the longest last tarsal segment (approx. 1.5X the Ist & 2nd segments
together) and the longest rostrum (index 0.33) of all the genera examined. The genus
is superficially similar to Hylopedetes Rehn (Fig. 8) and shares with it the very
elongate third tarsal segment. It differs from Hylopedetes in numerous details: lophi
of epiphallus not convergent towards the midline as in all species of Hylopedetes (Fig.
8G); proportionately longer and more slender legs and antennae; more pronounced
rostrum (index 0.28 in H. mirandus); deep anterior pronotal sulcus (rather weak in
Hylopedetes (Fig. 8A)); presence of well developed "epaulettes" between 2nd and 3rd
sulci (barely discernible in Hylopedetes, Fig. 8B); anterior ventral angle of pronotum
obtusely rounded, and not with a ventrally directed projection as in Hylopedetes (Fig.
8A); relatively well developed dorsal spine on hind knee (minute in Hylopedetes (Fig.
8F)); furcula with relatively shorter teeth than in Hylopedetes (Fig. 8E).
Etymology: Exerythros, "very red", referring to the coloration of the type
species; acris, conventionally used for "grasshopper".
Exerythracris volcanica n. sp.
Holotype d: COSTA RICA, Prov. Guanacaste: P.N. Guanacaste: Volcan Cacao, 1300
m, map ref. N323300, E375300, (H.E. Braker), 31 July 1987, specimen no. RC 87020.
Allotype 2°: COSTA RICA, Prov. Guanacaste: P.N. Guanacaste: SW slope of Volcan Cacao,
1040 m, map ref. N323300, E375300, (C.H.F. Rowell, N. Elsner, C. Chavez), 24 July 1991,
specimen no. RC 91184. Both in ANSP. Paratypes: COSTA RICA: 5 dd (specimen no.s
87016, 87017, 87018, 87019, 87021), data as holotype, RC. 1 d (specimen no. 91183), data as
allotype, RC. 1 2 (specimen no. 87022), data as holotype. 1 4 (specimen no. CRIOO1 013209),
same locality, but 11 July, 1988 (D. Janzen & W. Hallwachs), INBio. 3 d d (specimen no.s
CRIOO1 013226, CRIOO1 013227, CRIOO1 013229), same data, but 15 Feb 1989 (GNP
Biodiversity Survey), INBio. 1 2 (specimen no. CRI000 384037), same locality, but 1 Mar
1989 (PNG Inventario de Biodiversidad), INBio. 1 d (specimen no. CRI000 283607), same
locality, but 15 June 1990 (II. curso parataxénomo INBio), INBio. 2 dd (specimen no.s
91124, 91125), Prov. Guanacaste, Volcan Tenorio, nr. summit of road from Tierras Morenas to
Bajo Los Cartagos, 1040 m., map ref. N287250 E426500, 21 July 1991 (C.H.F. Rowell & N.
Elsner), RC. 1 9 (specimen no. 80237), Prov. Alajuela, nr. Monteverde, Pefias Blancas, Vargas'
pasture, edge of Elfin Forest, 1320-1370 m, map ref. N253300 E450500, 16 July 1980 (C.H.F.
Rowell, M. Rowell-Rahier, C. Hyde, H.E. Braker), RC.
Male. The live male is bright red with shining jet-black eyes, legs, antennae
and underside. In pinned specimens (described below) the colours become more
dilute.
560 C.H.F. ROWELL
Antennae, blackish purple. Eyes beige to brown. Labrum and mandibles, dark
purple. Palps green, suffused with black. Rest of head carmine red. Pro-, meso- and
metanota, thoracic episterna and epimera, carmine red. Sulci blackish. First 6
abdominal tergites with a medial carmine spot, decreasing in size on the more
posterior segments. Abdomen and thorax otherwise purplish black. All legs and feet
purplish black. Coxa, trochanter, and basal part of femur of fore and mid legs
yellowish green, especially on ventral surfaces. Spines of hind tibia greenish yellow
tipped with black.
Internal genitalia: epiphallus sharply constricted and ventrally depressed in
midline, bearing large conical sclerotized lophi, relatively widely spaced, the inner
faces of which are slightly divergent in axial view (Fig. 3A). Lateral epiphallic
sclerites present. Cingulum robust, almost completely enveloping the aedeagus (Fig.
3D, F). Aedeagus very short, upturned, sheathed: the inferior sclerites joined to the
rest of the endophallus by a narrow, almost vertical flexure (Fig. 3E). Anterior
apodemes of the endophallus flattened dorsoventrally, horizontal and markedly
divergent (Fig. 3G).
Female. In life the general olive brown coloration is marbled with yellow, the
band across the frons, genae and pronotal lobes is bright cream, abdominal segments
are green laterally. Distal abdominal segments (Sth onwards) are also green dorsally.
Basal segments of all legs are green; the black hind femur has a greenish cast.
Antennae dark brown. In pinned specimens (described below) the colours change.
Antennae purplish black. Eyes brown. Head olive brown, a paler stripe
extending across genae and frons and continuing posteriorly on the lobes of the
pronotum. Labrum and mandibles blackish purple. Palps green. Pronotum generally
olive brown; anterior and posterior margins and anterior ventral part of lateral lobes
olive green. Thorax and abdomen olive brown, tinged green on sides of abdominal
segments 2-5. Ovipositor valves purplish brown.
Fore leg: as in male. Mid leg: coxa, trochanter, basal part of femur light brown;
distal femur black; tibia greenish proximally, purplish distally; tarsus purple. Hind
leg: femur olive brown dorsally, inner and outer faces black; knee, tibia and tarsus
purple; spines of tarsus green tipped with black. Claws black.
Ovipositor valves long, extending 1.26 mm beyond end of supraanal plate
(Fig. 3J). Lower ovipositor valves with a proximal shoulder on outer lateral face (Fig.
31, J), laterally compressed, normally held between the dorsal valves (Fig. 3K). Both
upper and lower ovipositor valves toothed on outer margins, inner faces bearing
numerous long hair sensilla. Subgenital plate with concave posterior margins and a
simple central apex (Fig. 3H).
(The single adult female from Peñas Blancas differs from those described
above only in a) a slightly shorter pronotum and b) in coloration: the pale band on the
face and prothorax is faint and restricted to the frons; antennae light brown, not
purple-black; hind femora and tibia green-brown and green respectively, not black
and purple. In the absence of a corresponding male it cannot be decided whether this
specimen represents another species of the genus or a local colour variant. The latter
is assumed here, but this individual is not included in the data of Table 1).
COSTA RICAN RHYTIDOCHROTINAE 561
FIG. 3
Exerythracris volcanica n.sp. Figs. A-G, male. A-C, epiphallus: A, axial view, B, lateral view,
C, dorsal view. D & F, endo- and ectophallus; D, lateral view, F, dorsal view. E & G,
endophallus; E. lateral view, G, dorsal view. Figs H-K, female. H, posterior part of subgenital
plate, ventral view. I, ovipositor valves, ventral view. J, tip of abdomen, lateral view, ovipositor
valves open. K, as J, but valves shut.
562
Dimensions: See Table 1.
C.H.F. ROWELL
MEAN
Dimensions (mm)
Hind femur (F)
Rostrum-subgen. plate (L)
Pronotum (midline) (P)
Interocular space
Antennal pedicel (width)
Antennal flagellum (width)
Antenna (length)
Hind tarsus, Ist + 2nd segments
Hind tarsus, 3rd segment
Ratios
F/P
LP
Interoc./P
Interocular/pedicel
Tarsus 3/1+2
Tarsus 1+2+3/F
Tarsus 1+2+3/P
Antennal width/length as %
Dimensions (mm)
Hind femur (F)
Rostrum-ov. valves (L)
Pronotum (midline) (P)
Interocular space
Antennal pedicel (width)
Antenna (length)
Hind tarsus, Ist + 2nd segments
Hind tarsus, 3rd segment
Ratios
F/P
IC
Interoc./P
Interocular/pedicel
Tarsus 3/1+2
Tarsus 1+2+3/F
Tarsus 1+2+3/P
TABLE |
Exerythracris volcanica n.sp.: dimensions.
SD: SD}
MAX
N
MEAN
Males
13.76 0.42 0.03 14.61 13.27 14
17.52 0.71 0.04 19.00 16.01 14
3.20 0.10 0.03 3.39 3.02 14
0.37 0.03 0.08 0.41 0.32 14
0.53 0.02 0.04 0.57 0.48 14
0.30 1
12.94 1.18 0.09 15.25 11.40 7
1.87 0.11 0.06 1.99 1.62 13
2.80 0.12 0.04 3.00 2.66 12
4.31 0.11 0.03 4.50 4.15 14
5.48 0.22 0.04 5.83 5.09 14
0.12 0.01 0.10 0.13 0.10 14
0.71 0.07 0.10 0.81 0.60 14
IRSA, 0.08 0.06 1.64 1.39 12
0.34 0.01 0.04 0.36 0.32 12
1.45 0.06 0.04 LS 1.36 12
232% 1
Females
15.21 0.61 0.04 15.95 14.65 4
23.98 0.73 0.03 24.84 23.06 4
3.87 0.23 0.06 4.16 3.66 4
0.33 0.04 0.07 0.58 0.51 4
0.60 0.04 0.07 0.65 0.56 4
12.03 1
2.02 0.09 0.04 DA 1.92 4
2.86 0.29 0.10 2112 2.45 4
3.93 0.07 0.02 4.00 3.83 4
6.21 0.45 0.07 6.66 5.80 4
0.14 0.02 0.11 0.16 0.12 4
0.92 0.12 0.13 1.04 0.78 4
1.41 0.09 0.07 1.48 1.28 4
0.32 0.01 0.05 0.33 0.30 4
1.26 0.04 0.04 1.29 1.19 4
COSTA RICAN RHYTIDOCHROTINAE 563
Larvae. The general coloration of the larvae is dull green and brown with
reddish brown eyes and antennae. A number of larvae have been examined, but none
reared throughout their development. Here I assume 5 larval instars:
I. Unknown.
II. Antennae with 8 flagellar segments, reddish brown. Eyes shining reddish
brown. Legs and dorsal surfaces plain green, speckled reddish brown; ventral surface
brown. Hind femora with 2 brown bands on inner and outer faces, knees brownish.
III. Antennae with 9 flagellar segments. Segments 2-6 bear an annulus, giving
the impression of 14 segments in all. Eyes shining reddish brown. Well-marked
medial dorsal carina on all thoracic and abdominal segments. Coloration as in instar
II, except a) hind femoral bands fainter b) dark brown postocular stripe, more marked
on head than on pronotum c) yellow flecks present on disc of pronotum.
IV. Antennae with 15 flagellar segments. Head bright green. Palps green.
Dorsal surface of thorax and abdomen green flecked with brown. Yellow flecks on
pronotum seen in III instar now enlarged to form a horizontal stripe below the
postocular stripe, extending from pronotal lobes and around genae and frons (female
only). Meso- and metathoracic epimera carmine red (male only). Underside black.
Fore and mid legs black. Hind knees black, tibiae black.
V. Coloration as adult. 16 flagellar segments in antenna.
Etymology: "volcanica" of volcanoes, an allusion to both the distribution on
the volcanoes of northern Costa Rica and the bright red colour.
Distribution, natural history. E. volcanica inhabits light gaps, path margins
and woodland edges in wet montane forest between 1000 and 1200 m on the volcanic
peaks of the Cordillera de Guanacaste and the Cordillera de Tilaran. Within the
former it seems to have a disjunct distribution, being known from Volcan Tenorio
(the Tierras Morenas site) in the South and from Volcan Cacao in the North, but not
from the intervening peaks of Volcan Rincon de la Vieja or Volcan Miravalles. On
Tenorio it is sympatric with Hylopedetes surdus, and in the Cordillera de Tilaran with
both H. surdus and H. gemmeus, both of which belong to the same subfamily and
have similar food preferences. Both Exerythracris and Hylopedetes are principally
associated with and eat ferns, especially the secondary succession species Hypolepis
hostilis (see also Rowell et al. 1983). A male E. volcanica has been seen attempting
copulation with a female H. surdus and actively repelling males of the latter species
while doing so.
Brakeracris n. gen.
Type species: Brakeracris varablancensis n. sp.
Male (Fig. 4 & 5). Size small to medium. Integument polished except on head,
the integument of which is matte. Dorsal surfaces of occiput, pro-, meso- and meta-
nota and Ist abdominal tergum heavily pitted. Legs, frons and remaining abdominal
segments smooth. Integument with numerous white hairs, especially on hind tibiae,
genital area and distal underside of abdomen, but also sparsely on the dorsal surface.
564 C.H.F. ROWELL
Fastigium subhorizontal, triangular, truncate anteriorly, bearing a pair of
prominent longitudinal ridges proximally, distally grooved, the groove reaching the
anterior margin of fastigium (Fig. 4B, D, E). Frontal ridge narrower (0.8X) than
interocular space, straight and almost parallel sided, slightly tapering ventrally, irre-
gularly pitted in midline, ending above medial ocellus. Medial and ventral arms of
subantennal suture short and deeply incised, dorsal arms thin and curved (Fig. 4E).
Frons concave, upper part of frontal ridge forming a blunt rostrum (Fig. 4B), index
0.30. Preocular ridges well marked. Eyes globose. Interocular space narrower (0.7X)
than maximum width of antennal pedicel. Antennae of medium length (3.57X length
of pronotum), slightly flattened dorsoventrally and rather thick (width 0.032X their
length), 17 segments in flagellum of adult.
Thorax (Fig. 4B, C). Medial carina present on pronotum anterior to Ist sulcus
and posterior to 3rd sulcus, but absent between Ist - 2nd and 2nd - 3rd sulci.
"Epaulettes" present between 2nd and 3rd sulci, but less well developed than in
Exerythracris or Talamancacris. Posterior ventral angle of lateral lobe of pronotum
rounded, posterior margin of pronotum slightly concave. Anterior ventral angle of
lateral lobe with a small anteriorly directed projection (Fig. 4B). Posterior margins of
pronotum slightly concave in midline, anterior margin straight or slightly notched.
Prosternal process short, abruptly conical, vertical. Medial carina absent on meso- and
metanotum. Apterous.
Hind femur relatively long, 4.1 - 4.2X as long as pronotum. Dorsal carina of
femur minutely toothed proximally, smooth distally, ending in a spine which is
shorter than lateral lobes of knee (Fig. 4G). Ventral carina smooth, lateral carinulae
absent or very slight, outer face of hind femur rounded. Seven non-terminal spines on
the dorsal external margin of hind tibia, 8 on the dorsal internal. Hind foot long,
0.34X as long as femur; 3rd tarsal segment relatively short, 1.26X as long as Ist and
2nd tarsal segments together.
Tympanum absent. Medial carina absent from Ist abdominal segment, present
on 2-6th segments. Posterior margin of last abdominal tergite with a central notch and
short lateral sclerotised teeth (Fig. 4F). Supraanal plate triangular, without sclerotised
melanic markings, with a short longitudinal medial furrow at its base. Cerci short,
conical, simple. Subgenital plate bluntly pointed in lateral view, terminating in a short
medial furrow in dorsal view. Internal genitalia (Fig. 5) of the usual type for sub-
family.
Female. The female is so far known only from several last instar larvae.
Comparison of these with similarly aged larvae of other rhytidochrotine genera for
which the female is known indicates that the adult female is likely to be of
conventional form for the group, somewhat larger and more fusiform than the male,
and with the characteristic ovipositor valves.
Etymology: Named for Dr. H.E. Braker, for her contribution to the discovery
and original collection of all three new genera described in this paper.
Diagnosis. Brakeracris seems to be related to the Costa Rican Scirtopaon (Fig.
9). It shares with this genus the presence of a weak medial carina on the anterior and
COSTA RICAN RHYTIDOCHROTINAE 565
G
FIG. 4
Brakeracris varablancensis n.sp. Male. A, lateral view of entire insect. B, lateral view of head
and pronotum. C, dorsal view of pronotum. D, dorsal view of head. E, frontal view of face. F,
dorsal view of furcula, supraanal plate and cerci. G, lateral view of hind knee.
566 C.H.F. ROWELL
posterior sections of the pronotum (Fig. 9B) and on the posterior abdominal segments;
the slight anterior projection of the anterior ventral angle of the pronotum (Fig. 9A);
the size and shape of the terminal spine of the hind knee (Fig. 9H). It differs from
Scirtopaon as follows: frontal ridge with subparallel margins, and not with a double
constriction as in Scirtopaon (Fig. 9D); medial and ventral antennal sutures strongly
excavated (medial and dorsal in Scirtopaon (Fig. 9D)); shorter and thicker antennae;
more prominent rostrum (index 0.30, compared with 0.26 in Scirtopaon); in the detail
of the rugosities of the pronotum (Fig. 9A, B); in the less well developed teeth of the
furcula; in the shape of the lophi - in Scirtopaon (Fig. 9F, G) these have a prominent
lateral ridge which is absent in Brakeracris, and are not tilted caudally as in
Brakeracris. B. varablancensis shows none of the extreme values in the morpho-
metric ratios (Table. 4), but has a rather larger rostrum than most other taxa and the
relatively thickest antennae of all but Talamancacris.
Brakeracris varablancensis n. sp.
Holotype & (specimen no. RC 86495a) COSTA RICA: Prov. Heredia, 3 km. E. of S.
Rafael de Vara Blanca, 1800-2100 m., map ref. N239800, E524200, 13 April 1986, (C.H.F.
Rowell & H.E. Braker), ANSP. Paratype: 1 é, same data as holotype (RC). Last instar larvae:
6 2 ®, same data as holotype (2 in ANSP, otherwise RC).
Male. Alive a predominantly greenish-black insect with bright yellow mar-
kings and green legs. Antennae black; eyes, black; palps, green; suborbital band on
frons and genae, yellow; remainder of head, black. Pronotum black with yellow
border; thoracic and abdominal tergites olive green with yellow markings. Coloration
otherwise as for pinned specimen.
Description of pinned specimens. Basal segments of antenna green, distal
segments dark brown. Eyes reddish brown. Ventral half of frons and genae cream.
Remainder of head blackish green. Thorax and abdomen olive green, underside olive
brown. Entire margin of pronotum bordered in deep yellow apart from a short stretch
dorsal to prothoracic coxal insertion. Metathoracic episternum yellow. Posterior
margin of metanotum with 2 yellow marks. Terga of abdominal segments 1-8 each
bear a pair of yellow chevrons in the midline and paired yellow spots laterally. Legs
dark green, lunules of hind knee reddish, 3rd tarsal segments black.
Internal genitalia: lophi of epiphallus conical, sclerotised, minutely denticled,
widely separated and with inner faces parallel or only slightly divergent in axial view
(Fig. 5A), tilted slightly caudally in lateral view (Fig. 5B). Remainder of genitalia
without particular distinguishing features.
Larval coloration (male):
I instar: unknown.
II instar: uniform dull reddish brown. Antennae slightly clubbed, pink at tip.
Inner surface of hind femur with 3 weakly marked darker bands.
II. instar: Thoracic tegument more rugose than in II. Fore and mid legs green.
Hind leg brown, femur with 3 dark bands.
IV. instar. Pattern similar to that of the adult, but expressed in pink on reddish
brown instead of yellow on greenish black. Legs tinged green.
COSTA RICAN RHYTIDOCHROTINAE 567
FIG. 5
Brakeracris varablancensis n.sp. Male. A-C, epiphallus: A, axial view, B, lateral view, C,
dorsal view. D, F, endo- and ectophallus; D, lateral view, F, dorsal view. E, G & H,
endophallus; E, lateral view, G, perspective view from below and to one side, H, dorsal view.
V instar: as adult.
Larval coloration (female):
The early instar female larvae do not differ from the males, the last instar larva
differs from the adult and last instar male as follows: coloration duller, less contrasty.
Lower half of pronotal lateral lobe yellow. Pronotum without the clear yellow border
of the male. Abdominal terga with a single triangular yellowish patch, not with paired
chevrons and spots as in the male.
Dimensions: See Table 2.
Etymology: varablancensis, of Vara Blanca, the type locality.
Distribution, natural history. B. varablancensis is known to date only from
around S. Rafael de Vara Blanca, on the northern scarp of the Cordillera Central, in
the headwaters of the Rio Sarapiqui, altitude 1900 to 2200 m. In March 1986 it was
found in regenerating scrub at the forest edge above the farmhouse (now situated in
the Parque Nacional Braulio-Carrillo) and also on landslip areas bordering two stream
568 C.H.F. ROWELL
TABLE 2
Brakeracris varablancensis n.sp.: dimensions.
MEAN MAX MIN N
Males
Dimensions (mm)
Hind femur (F) 12292 13.11 12.73 2
Rostrum-subgen. plate (L) 15299 16.61 15.36 2
Pronotum (midline) (P) 3209 3.09 3.08 2
Interocular space 0.40 0.40 0.40 2
Antennal pedicel (width) 0.57 0.59 0.54 2
Antennal flagellum (width) 0.35 1
Antenna (length) 11.00 1
Hind tarsus, Ist + 2nd segments 1295 1
Hind tarsus, 3rd segment 2.45 1
Ratios
F/R 4.19 4.26 4.12 2
IE 5.18 5.38 4.99 D
Interoc./P 0.13 0 0.13 2
Interocular/pedicel 0.71 0.74 0.68 2
Tarsus 3/ 1+2 1.26 1
Tarsus 1+2+3/F 0.34 1
Tarsus 1+2+3/P 1.43 1
Antennal width/length as % 3.18% 1
beds traversing dense montane wet forest 1-2 km north of the farmhouse. Most
individuals seen were mid or late larval instars; only two adult males were found, and
no adult females, despite intensive search. There was no obvious association between
3. varablancensis and any particular plant species. In captivity it nibbled at a variety
of plants from the habitat, but ate well from none of those tested, which included
several species of ferns. In the same localities were found the acridids Drymacris
nebulicola (Proctolabinae) on Solanaceous plants and Si/vitettix communis (Gompho-
cerinae) on grasses.
Intensive search of the same areas under fine weather conditions in July 1991
produced only a single 3rd instar larva and no adults. This may indicate marked
seasonality, or might be due to the extensive vegetational change seen, corresponding
to 5 years of secondary succession. This larva ate various Senecio spp. from other
parts of Costa Rica in the following days, suggesting that B. varablancensis, like T.
palustris (below), may prefer Asteraceae.
Talamancacris n. gen.
Type species: Talamancacris palustris n. sp.
Male. (Figs. 6 & 7). Size small-medium. Integument highly polished except on
head, which is matte. Frons, genae, dorsal surfaces of occiput, and of pronotum
COSTA RICAN RHYTIDOCHROTINAE 569
posterior to the last transverse sulcus, lightly pitted; remaining areas smooth and
glossy, as though clear varnished. Integument with numerous white hairs, especially
on the tibiae, proximal hind femora, genital area and meso- and metathoracic sterna,
and also bilateral tufts dorsolaterally near anterior margin of each abdominal segment.
Fastigium inclined, triangular, truncate anteriorly, with two low longitudinal
ridges proximally, shallowly grooved distally, lateral margins straight or slightly
concave (Fig. 6B, D). Frontal ridge at its widest point narrower (0.6X) than inter-
ocular space, subparallel sided, somewhat narrower at dorsal and ventral extremities,
extending to lower rim of antennal foramen, slightly divergent above medial ocellus;
shallowly grooved medially for a short distance dorsal to medial ocellus, lightly pitted
between this point and fastigium (Fig. 6E). Medial ocellus barely visible. Medial and
ventral arms of subantennal suture short and rather deeply incised, dorsal arms thin
and indistinct (Fig. 6E). Profile of frons concave, upper part of frontal ridge forming a
short (index 0.30), vertically ended rostrum (Fig. 6B). Preocular ridges present. Eyes
globose. Interocular space slightly wider (1.1X) than width of antennal pedicel.
Antennae filiform, somewhat dorsoventrally flattened, short (2.8X as long as
pronotum) and thick (0.042X their length), 17 segments in flagellum of adult.
Pronotum (Fig. 6B, C) slightly inflated in midline anterior to Ist sulcus, but no
medial carina present. Sulci deep; well developed "epaulettes" present between the
2nd and 3rd sulci. Posterior ventral angle of lateral lobe of pronotum rounded,
somewhat lobate, projecting ventrally and posteriorly (Fig. 6B). Anterior ventral
angle rounded. Posterior margins of pronotum slightly concave in midline, anterior
margin convex and minutely notched (Fig. 6C). Prosternal process short, abruptly
conical, vertical. Medial carina absent from meso- and metanotum. Apterous.
Hind femur relatively short, 3.57X as long as pronotum. Dorsal carinae of
femur well marked over most of its length, slightly toothed, obsolete immediately
before the knee, terminating in a sharp spine which exceeds in length the outer lateral
lobes of the knee (Fig. 6G). Ventral carina strongly developed, smooth, lateral
carinulae absent or very slight, outer face of hind femur rounded in cross section.
Seven non-terminal spines on external dorsal margin of hind tibia, 8 on internal dorsal
margin. Hind tarsus long in relation to femur (0.32X), but smaller than in the other
Costa Rican genera in relation to the pronotum (1.13X); 3rd tarsal segment of hind leg
slightly (1.28 X) longer than Ist and 2nd tarsal segments together.
Tympanum absent. Medial carina absent from Ist but present on 2-6th
abdominal segments. Posterior margin of last abdominal tergite with a central notch
and short lateral sclerotised teeth (Fig. 6F). Supraanal plate triangular, without
sclerotised melanic markings, with a short longitudinal medial furrow at its base (Fig.
6F). Cerci short, conical, simple. Subgenital plate bluntly pointed in lateral view,
terminating in a short medial furrow behind the tip of the supraanal plate, glabrous in
ventral and apical midline, pilose laterally. Internal genitalia (Fig. 7A-G) of the usual
type for the subfamily. Epiphallus distinctive (Fig. 7A-C).
Female. Size, small to medium. Differs from male in following morphological
characters: larger (1.38x in overall length) and more fusiform, wider in posterior
thorax and anterior abdominal segments (Fig. 7K). Ovipositor valves (Fig. 7H-J) long,
570 C.H.F. ROWELL
FIG. 6
Talamancacris palustris n.sp. Male. A, lateral view of entire insect. B, lateral view of head and
pronotum. C, dorsal view of pronotum. D, dorsal view of head. E, frontal view of face. F,
dorsal view of furcula, supraanal plate and cerci. G, lateral view of hind knee.
COSTA RICAN RHYTIDOCHROTINAE 571
IG
Talamancacris palustris n.sp. Figs. A-G, male. A-C, epiphallus; A, axial view, B, lateral view,
C, dorsal view. D & F, endo- and ectophallus, D, lateral view, F, dorsal view. E & G,
endophallus; E, lateral view, G, dorsal view. Figs H-L, female. H, tip of abdomen, dorsal view.
I, ovipositor valves and posterior end of subgenital plate, ventral view. J, tip of abdomen,
lateral view, ovipositor valves open. K, pronotum, dorsal view. L, pronotum, lateral view.
572 C.H.F. ROWELL
projecting 1.12 mm beyond end of the subgenital plate. Both upper and lower
ovipositor valves heavily toothed on exterior outer margins, smooth on inner margins.
Lower valves narrow, normally held between upper valves, with a shoulder
proximally on lateral ventral edge. Subgenital plate (Fig. 71) with concave posterior
edges, forming a medial point. Spermatheca not examined.
Diagnosis. Talamancacris differs from all other Costa Rican rhytidochrotines
in that a) the terminal spine of the hind knee projects well beyond the lateral lobes b)
the interocular space exceeds the maximum width of the first antennal segment and c)
in the shape of the male epiphallus. It is also much more hirsute than the other genera.
The dimensional comparison with other genera is shown in Table 4. Talamancacris
has the highest number of extreme morphometric ratios of any of the taxa studied: the
shortest overall length relative to the pronotum, the widest interocular space, the
shortest hind tarsus relative to the pronotum, and the relatively shortest and thickest
antennae. Talamancacris gives a first impression of a small, short legged version of
Brakeracris; especially the profile of the head and the form of the frontal ridge and
the antennal sutures are very alike in the two genera. The abdomen, legs and antennae
are however relatively much shorter (relative to pronotum), the antennae are relatively
thicker (0.042x their length, 0.032 in Brakeracris), the pronotum has no medial
carina, its anterior ventral angle lacks projections, and the integument is glossy and
smooth.
Etymology: Talamanca, the range of mountains in central Costa Rica where all
known localities of the genus are located; acris, conventionally used for grasshopper.
Talamancacris palustris n. sp.
Holotype d (specimen no. RC 93371): COSTA RICA, Prov. Cartago, Turbera "La
Chonta", 2.7km SSE of El Empalme, 2'380 m, map ref. N188200, E542300, 20 Sep 1993,
(C.H.F. Rowell & G. Vargas). Allotype 9 (specimen no. RC 93372), same data. Both in
ANSP. Paratype: 1 © (specimen no. RC 79228): COSTA RICA, Prov. S. José, Puya bog, Km
67.5, Rta 2, nr. Salsipuedes, 2550 m, 25 August 1979 (H.E. Braker) (MNHNP). Last instar
larvae: 2 dd, 1 ©, same data, but 16 July 1980 (C.H.F. Rowell & M. Rowell-Rahier); 2 86,
same locality as holotype, but 25 January 1991 (G. Vargas & H.E. Braker); all RC.
Both sexes alive are dark metallic bluish green, with black antennae and eyes,
a shining white facial and pronotal stripe, paired lighter green chevrons on each
abdominal tergum, and reddish hind knees. The striking blue-green colour fades to
brown an hour or two after death and is probably structural in origin.
Description of pinned specimens: Male. Antennae black, brown at tips. Eyes
dark brown. Lower frons and genae yellow. Head otherwise dark blue green with
yellow V-shaped marking on occiput and fastigium. Thorax and abdomen blackish
brown shading to green at anterior and posterior margins of pronotum and around
bases of anterior legs. Ventral half of lateral lobe of prothorax yellow. Pro- and
mesothoracic epimera yellow. Legs dark olive brown. Tibial spines olive green tipped
with brown. Hind knees blackish green. Abdominal segments 2-6 with a pair of
yellow triangular marks tending to fuse in midline. Cerci brown.
COSTA RICAN RHYTIDOCHROTINAE
573
Internal genitalia (Fig. 7A-G) of usual type for the group. Epiphallus
distinctive: lophi (Fig. 7A-C) vertical in axial view, very slender at their tips, with a
sloping lateral ridge, posterior margin of lateral processes slightly incised at the base
of the lophi. Lateral sclerites present.
Female. Antennal basal segments green, distal segments dark brown. Eyes
reddish brown. Ventral half of frons and genae cream. Remainder of head olive brown
with a lighter V-shaped marking on the occiput. Thorax, abdomen and legs olive
green with reddish brown mottle. Underside olive brown. Lower half of prothoracic
lateral lobes pale yellow brown.
Dimensions: see Table 3.
TABLE 3
Talamancacris palustris n.sp.: dimensions.
MEAN MAX MIN N
Males
Dimensions (mm)
Hind femur (F) 11.46 1
Rostrum-subgen. plate (L) 14.47 1
Pronotum (midline) (P) 3.21 1
Interocular space 0.60 1
Antennal pedicel (width) 0.54 1
Antennal flagellum (width) 0.38 I
Antenna (length) 9.00 1
Hind tarsus, 1st + 2nd segments 1.60 1
Hind tarsus, 3rd segment 2.04 1
Ratios
ID 3557] 1
L/P 4.51 1
Interoc./P 0.19 1
Interoc./pedicel 1.11 1
Tarsus 3/ 1+2 1.28 1
Tarsus 1+2+3/F 0.32 1
Tarsus 1+2+3/P 1.13 1
Antennal width/length as % 4.22% 1
Females
Dimensions (mm)
Hind femur (F) 13.12 1321 13.02 2
Rostrum-ov. valves (L) 19.97 DSS) 18.41 2
Pronotum (midline) (P) 3.90 3.93 3.87 DI
Interocular space 0.72 0.74 0.70 2
Antennal pedicel (width) 0.61 0.62 0.60 2
Antenna (length) 9.80 10.39 9.20 2
Hind tarsus, 1st + 2nd segments 2.00 2.03 1.97 2
Hind tarsus, 3rd segment DATO 229 2.09 2
Ratios
EE 3.36 3.41 SSL 2
11% 512 5.48 4.76 2
Interoc./P 0.18 0.19 0.18 2
Interoc./pedicel 1.18 1725 113113) 2
Tarsus 3/ 1+2 1.10 1.16 1.03 2
Tarsus 1+2+3/F 0.32 0.33 0.31 2
Tarsus 1+2+3/P 1.07 1.08 1.06 2
574 C.H.F. ROWELL
Larvae:
I instar: unknown.
II instar: uniform dull reddish brown.
IH - V instars: coloration similar to that of adults, but less contrasty. Some
female larvae have 2 pale blotches on the dorsal edge of the hind femur.
Etymology: "palustris", of swamps, after the habitat of the species.
Distribution, natural history. T. palustris is known only from a few bogs lying
between 2300 and 2600 m altitude on the Atlantic side of the north slope of the
Talamanca range. All localities have a similar and very characteristic vegetation
dominated at ground layer by Sphagnum moss and the rush-like Xyris (Xyriaceae);
the main emergents are the large fern Blechnum and the terrestrial Bromeliad Puya.
This characteristic plant assemblage, otherwise known only from the Andes, is
confined to certain areas of restricted drainage which appear to date from after the end
of the last glaciation (G. Vargas, pers. comm.). The only other acridoid found in or
near these bogs is the Oedipodine Chortophaga viridifasciata (De Geer, 1773).
Phenology. Larvae have been found in January, July, August and September,
but not in April; adults have been found in August and September. A third instar
female larva captured 20th September and maintained in captivity became adult on
Sth November. These data are compatible with the hypothesis that there is one
generation per year and that eggs are laid in the first half of the year, as is common
among Costa Rican acridids, but do not exclude the possibility that breeding takes
place year round.
Foodplants. T. palustris is not obviously associated with any particular plant
species in its habitat. It is found on the vegetation covering the bog surface, and not in
the emergent shrubs, bromeliads or ferns. Consistent with this observation it refuses
in captivity Puya, Blechnum and the Ericaceous shrub Pernettya. It also refuses
Sphagnum and Xyris and a variety of other plants growing with them. The only plant
from the native habitat which Talamancacris has accepted in captivity 1s Hieracium
stanleyi (Asteraceae). In captivity it refuses all monocots and most exotic dicots, but
accepts a variety of cultivated Asteraceae, including Doronicum and Dahlia, espe-
cially the latter. (Dahlia imperialis 1s indigenous to Costa Rica and grows within
some kilometres of the bogs inhabited by T. palustris, but is not present within them).
Some but not all individuals accept in captivity the fern Hypolepis hostilis (the
commonest food of two other Costa Rican rhytidochrotine genera, Hylopedetes and
Exerythracris (ROWELL et al. 1983; this paper)) or Rubus nr. glaucus (eaten in the
wild by the Costa Rican rhytidochrotine Scirtopaon dorsatus (pers. obs.)), but do not
eat either of them in quantity. Neither plant is present in the natural habitat. These
observations suggest a specialisation of Talamancacris on Asteraceae, perhaps nor-
mally H. stanleyi.
PREVIOUSLY DESCRIBED TAXA
The original descriptions of the remaining three Costa Rican genera (Hylo-
pedetes, Scirtopaon and Micropaon) were not well provided with measurements or
COSTA RICAN RHYTIDOCHROTINAE 575
anatomical drawings. The following section is intended primarily to allow comparison
of the newly described genera with those previously described.
Table 4 compare the dimensions of the known atympanate genera. The data for
some are incomplete; in several cases only one of the two sexes is known.
Hylopedetes Rehn, 1929 (Fig. 8)
The genus is well differentiated from the remaining Costa Rican genera by the
weak first pronotal sulcus, the ventrally directed process at the anterior angle of the
pronotum (Fig. 8A), and the distinctive convergent lophi of the epiphallus (Fig. 8G).
Hylopedetes has the smallest relative interocular distance of any of the genera
examined, and (together with Talamancacris) the relatively shortest antennae. The
pattern of the antennal sutures is similar to that of Exerythracris, with which it also
shares a proportionately very long last segment of the hind tarsus. The male and
female genitalia of H. mirandus were figured by DESCAMPS & AMEDEGNATO (1972b:
Figs 2-8); Table 4 presents its dimensions. The 5 species of the genus are similar
morphologically, the most different being H. nigrithorax. A key to the species was
given by DESCAMPS & ROWELL (1978).
H. mirandus Rehn, 1929 (the type species) has previously been reported only
from the type locality, La Emilia, Guäpiles, which is now completely deforested and
suburban. Four new localities are now known. Three lie in the valley of the Rio Toro
Amarillo, 15 and 10 km S. and 7 km W. of Guäpiles, at 800, 540 and 305 m
respectively, a fourth 8km SW of Guäpiles in the valley of the Rio Sucio at 550 m. It
is probable that the species occurs in suitable places (at forest edges and in light gaps
on ferns, especially on Hypolepis hostilis) at least throughout the intervening area,
and perhaps further up the scarp beyond. Guäpiles itself lies at 260 m. on the edge of
the Caribbean plain at the foot of the scarp of the Cordillera Central, and the nature of
the original forest vegetation changed sharply north of and below this point. The type
locality probably represented the lower altitudinal limit of the species, the genus and
(at least in Central America) of the subfamily.
H. nigrithorax Descamps & Rowell, 1978. This is to date the only rhytido-
chrotine known from the southwest of Costa Rica (contrary to the statement by
DESCAMPS & AMEDEGNATO (1972b: 1058), the species of Hylopedetes described by
Rehn are from the Atlantic and not the Pacific slope of the country). Originally
described from a single locality on the Fila Cruces near the town of S. Vito de Coto
Brus, H. nigrithorax is now known from several other nearby localities lying to the
north and south along this same ridge, at altitudes between 1100 and 1700 m. The Fila
Cruces forms the most southwesterly ridge of high ground before the Pacific plain of
SW Costa Rica. It is not known whether the species extends south into Panama; it was
not collected in the '30s by D.W. Bishop from El Volcan in Chirriqui, where he found
several other species that in Costa Rica are sympatric with H. nigrithorax. The
preference of H. nigrithorax at the type locality for certain species of ferns (including
again Hypolepis hostilis) was documented previously (ROWELL et al. 1984); however,
it can sometimes be found on Asteraceae.
C.H.F. ROWELL
576
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FIG. 8
Hylopedetes mirandus Rehn, 1929. Male. A, lateral view of head and pronotum. B, dorsal view
of pronotum. C, dorsal view of head. D, frontal view of face. E, dorsal view of furcula,
supraanal plate and cerci. F, lateral view of hind knee. G-I. epiphallus; G, axial view, H, lateral
view, I, dorsal view. J, Endo- and ectophallus, lateral view. K & L, endophallus; K, lateral
view, L, dorsal view. (The whole animal (both male and female) is figured in the original
description. DESCAMPS & AMEDEGNATO (1972b, Figs. 2-8) give other drawings, including
additionally the female genitalia).
COSTA RICAN RHYTIDOCHROTINAE 579
The remaining three species of the genus are characteristic of the northern and
central mountains of Costa Rica, and have rather wider distributions than nigrithorax
and mirandus.
H. surdus Descamps & Rowell, 1978 has the largest range of any Hylopedetes
sp. and is also the commonest. It is sympatric with H. gemmeus over almost the entire
range of the latter and extends further north, at least as far as Volcan Tenorio, where it
is sympatric with E. volcanica. It appears to be less shade tolerant than gemmeus, but
like it is found on a variety of ferns, including Hypolepis. In the North and West of its
range H. surdus has orange hind knees, whereas to the South and East the hind leg is
entirely green. No other morphological difference can be discerned between these
colour forms. Museum specimens of the northern form of H. surdus can at first be
confused with H. mirandus, which also has pale knees. In life the true colours are
distinctive and the two species cannot be confused.
H. cruentus Rehn, 1929 occurs principally in an arc open to the NE around the
headwaters of the Rio Reventazön, between 1100 and 1700 m. The type locality (near
Navarro) lies on the southern arm of this arc, and it is also found at other localities in
the area between Cartago and Tapantî. Not apparently present on the Cerro de
Carpintera on the Continental Divide between Cartago and San José, it occurs again
on the southern slopes of Volcan Turrialba and is present on the north facing slope of
Cerro Zurqui. H. cruentus will eat the usual ferns in captivity, but in nature shows no
strong association with them. It is usually found in mixed vegetation at woodland
edges, often basking on leaves of Piper spp.
H. gemmeus Rehn, 1929 is the species most likely to be found in thick forest at
low light intensities. It too is usually found on ferns, especially members of the genera
Dryopteris and Pteris. It is now known from the Cordillera de Tilaran in the north and
from there south and east along the north slope of the Cordillera Central at least as far
as the Rio Frio. The type locality (the same as that of H. cruentus), however, lies well
to the south of this, in the Talamanca foothills at Navarro, but the species has not been
seen there since the original collection. This may be a consequence of the general
forest degradation in that area.
Scirtopaon dorsatus Descamps & Rowell, 1984 (Fig. 9)
Readily distinguished by the combination of blunt rostrum, incomplete
pronotal medial carina, and long, thin, filiform antennae, and by the pattern of the
antennal sutures. Originally described from Monteverde, in the Cordillera de Tilaran,
S. dorsatus is now additionally known from the head waters of the Rio Sarapiqui, near
Cinchona in the Cordillera Central. Patchy, never common, and rather cryptic, the
species is easy to overlook, and probably will ultimately prove to have a range similar
to that of H. gemmeus. It is not a fern or grass eater, but is associated with a variety of
montane dicots.
580 C.H.F. ROWELL
Fic. 9
Scirtopaon dorsatus Descamps & Rowell, 1984. Male. A, lateral view of head and pronotum.
B, dorsal view of pronotum. C, dorsal view of head. D, frontal view of face. E, dorsal view of
furcula, supraanal plate and cerci. F, lateral view of hind knee. G-I, epiphallus; G axial view, H,
lateral view, I, dorsal view (The original description additionally figures the whole male, the
male furcula and supraanal plate, the female fastigium and metanotum in dorsal view, and the
female ovipositor valves in lateral view).
COSTA RICAN RHYTIDOCHROTINAE 581
=
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E F G
FIG. 10
Micropaon lucens Descamps & Rowell, 1984. Male. A, lateral view of head and pronotum. B,
dorsal view of pronotum. C, lateral view of hind knee. D, frontal view of face. E-G, epiphallus;
E, axial view, F, lateral view, G, dorsal view. (The original description additionally figures the
whole male, the male furcula and supraanal plate, the female fastigium in dorsal view, and the
female ovipositor valves in lateral view).
Micropaon lucens Descamps & Rowell, 1984 (Fig. 10)
The genus is well differentiated morphologically by the flat fastigium without
grooves or ridges (Fig. 10D), the smooth, ungrooved frontal ridge (Fig. 10D), the
short rounded rostrum, the thickened margin to the posterior margins of the meso- and
metanotum, and the single medial black spot on the male supraanal plate. Micropaon
lucens is a very local species of the Pacific slopes of the Talamanca range, so far
known from only a few localities, all lying within a 3 km square on the hills above S.
Isidro del General. It is found in marshes or where water seeps out of the ground,
characteristically where Polygonum (Polygonaceae) and Galinsoga (Asteraceae)
grow, though it does not appear to eat either of these plants. Its normal diet is
unknown.
582 C.H.F. ROWELL
DISCUSSION
The Rhytidochrotinae is known as a very homogeneous group, and this holds
for the six Costa Rican genera, which all comply to a basic morphological plan and
are distinguished from each by rather small differences, as judged by the standards
within other acridid subfamilies. Especially the uniformity of the male genitalia of the
Rhytidochrotinae is unusual within the Acrididae, though the present work has shown
that the male epiphallus offers some taxonomic possibilities (true also of the South
American taxa examined). In spite of their formal similarities and the isolated
geographic position of the Costa Rican taxa, it is not easy to speculate convincingly
about their interrelationships. Only Scirtopaon and Brakeracris appear to share some
possibly derived characters which may indicate a recent common ancestry. The other
genera appear quite distinct from these two and from each other.
It is perhaps necessary to stress that despite formal morphological similarities,
the Costa Rican rhytidochrotines have clearly different ecological requirements, very
different colour patterns and usually do not overlap in distribution. They present no
difficulties at all in field determination, it is merely the technical separation which is
harder.
The Rhytidochrotinae, with its small number of taxa and interesting present day
distribution (see also DESCAMPS & AMEDEGNATO 1972b) would be an interesting group
for an independent study of phylogeny based on genomic sequence data. To date no
rhytidochrotines are known from Central America north of Costa Rica (Exerythracris,
extending to near the Nicaraguan border, is the most northerly known representative of
the subfamily) and the only known Panamanian genus (Piezops) comes from the
extreme South of that country near the Colombian border. All the remaining Central
American rhytidochrotines are confined to Costa Rica. It is quite possible that some of
the apparent isolation of the Costa Rican rhytidochrotines is an artifact of inadequate
collecting or relatively recent human destruction of the habitat, particularly in Northern
Panama. However, as all the Costa Rican rhytidochrotines are montane forms, it is
equally possible that the lowlands of central and southern Panama are indeed a natural
barrier to dispersal, at least under the present climatic conditions. The last Ice Age must
have diminished this barrier very considerably, and the presence of numerous Andean
elements on the paramos of the Talamanca Range of Costa Rica (see WEBER 1958, and
for other acridological consequences, ROWELL & CARBONELL 1977) indicates that many
organisms were able to disperse between these two habitats in the past, possibly
including the rhytidochrotines. The Puya/Blechnum swamps of the Talamancas are a
striking example, and Talamancacris (unless it might be discovered in the equivalent
Andean habitats) would seem to be a relict taxon with a remarkably small range and
population size and potentially extremely vulnerable to extinction by habitat alteration.
ACKNOWLEDGEMENTS
I am grateful to Dr. C. Amédégnato (Paris), Prof. C.S. Carbonell (Montevideo)
and Dr. A. Solis and Dr. J. Jiménez (Instituto Nacional de Biodiversidad, Costa Rica)
for the loan of material, to the Servicio de Parques Nacionales and other Costa Rican
COSTA RICAN RHYTIDOCHROTINAE 583
landowners for permission to collect on their land, and to Dr. H.E. Braker, Sr. C.
Chavez, Prof. N. Elsner, Dr. L.D. Gömez, Prof. D. Janzen, Dr. M. Rowell-Rahier and
Dr. G. Vargas for assistance in the field and for useful discussions. An anonymous
reviewer provided helpful criticism of the manuscript. Some of the relevant field trips
were supported financially by the National Science Foundation (USA), the National
Geographical Society (USA) and the Freie Akademische Gesellschaft (Basel,
Switzerland).
REFERENCES
AMEDEGNATO, C. 1974. Les genres d'Acridiens neotropicaux, leur classification par familles,
sous-familles, et tribus. Acrida 3: 193-204.
AMEDEGNATO, C. 1977. Etude des Acridoidea Centre et Sud Americains (Catantopinae sensu
lato): Anatomie des genitalia, classification, répartition, phylogenie. Thése, Université
Pierre et Marie Curie, Paris. 385 pp.
COLLINS, G.B. 1992. A specialised area of unknown function on the antennae of British
grasshoppers (Orthoptera: Acrididae). Entomologist 111: 195-200.
DESCAMPS, M. & C. AMEDEGNATO. 1972a. Contribution à la faune des Acridoidea de Colombie
(Missions M. Descamps). III. Diagnoses de Catantopinae (sensu lato). Annis. Soc.
entomol. Fr. (N.S.) 8: 505-559.
DESCAMPS, M., & C. AMEDEGNATO. 1972b. Contribution à la faune des Acridoidea de Colombie
(mission M. Descamps). IV. Le groupe Rhytidochrotae. Bull. Mus. nat. d'Hist. nat.,
Paris, 3° ser. Zool. 65: 1057-1096.
DESCAMPS, M., & C.H.F. ROWELL. 1978. Acridiens des clairiéres de Costa Rica: diagnoses,
signalisations, notes biologiques, polymorphisme (Acridomorpha, Acrididae). Annls
Soc. entomol. Fr. (N.S.) 14: 351-367.
DESCAMPS, M., & C.H.F. ROWELL. 1984. Diagnoses d'Acridoidea des forêts de Costa Rica.
Annls Soc. entomol. Fr. (N.S.) 20: 143-161.
REHN, J.A.G. 1929. Studies in Costa Rican Dermaptera and Orthoptera. I. New genera and
species of Acrididae. Trans. Am. entomol. Soc. 55: 9-77.
ROWELL, C.H.F. & C.S. CARBONELL, C.S. 1977. Baeacris talamancensis (gen. and sp. nov.)
(Acrididae, Melanoplinae), a neotropical montane grasshopper: its implication for the
origin of the Dichroplini and of the Costa Rican paramo. Acrida 6: 55-74.
ROWELL, C.H.F., M. ROWELL-RAHIER, H.E. BRAKER, G. COOPER-DRIVER & L.D. GOMEZ. 1983.
The palatability of ferns and the ecology of two tropical forest grasshoppers. Biotropica
15: 207-216.
WEBER, H. 1958. Die Paramos von Costa Rica und ihre pflanzengeographische Verkettung mit
den Hochanden Siidamerikas. Abh. Akad. Wiss. Lit. Jg. 1958: 116-194.
REVUE SUISSE DE ZOOLOGIE, 102 (3) : 585-608; septembre 1995
Regenwürmer aus Bolivien (Oligochaeta) !
Andräs ZICSI
Bodenzoologische Forschungsgruppe der Ungarischen Akademie der Wissenschaften,
am Lehrstuhl für Tiersystematik und Ökologie der Eötvös-Loränd Universität,
Puskin utca 3, H-1088 Budapest, Ungarn.
Earthworms from Bolivia (Oligochaeta). - Twenty-eight species of
terrestrial Oligochaeta from Bolivia were studied. They are distributed in
21 genera and 6 families. One new genera, Tamayodrilus and 6 new species
Andiorrhinus (Amazonidrilus) bolivianus, Andiorrhinus (Andiorrhinus)
montanus, Tamayodrilus roembkei, Inkadrilus hanagarthi, Martiodrilus
silvestris, Belladrilus (Belladrilus) vaucheri are described.
Key-words: Earthworms - Glossoscolecidae - Acanthodrilidae - Octo-
chaetidae - Ocnerodrilidae - Lumbricidae - Megascolecidae - Taxonomy -
Bolivia.
EINLEITUNG
Da aus Bolivien nur vereinzelte Angaben über das Vorkommen von Regen-
würmern vorliegen (BEDDARD, 1892; COGNETTI, 1902 a, b, 1905; MICHAELSEN, 1902,
1918; RıGHI & ROMBKE, 1987; Csuzpi & Zicsi, 1991; Zıcsı, 1992), ist es von
Interesse, die im Rahmen der Ungarischen Bodenzoolologischen Expedition 1966-
1967 (Teilnehmer: Dr. J. Balogh, Dr. S. Mahunka und Dr. A. Zicsi) vorwiegend in
den Provinzen Beni und La Paz gesammelten terrestrischen Oligochaeten bekannt zu
machen. Für die Überlassung einer kleineren Ausbeute von Regenwürmern aus
Bolivien spreche ich den Herren Dr. J. Römbke (Frankfurt/Main)? und Dr. CI.
Vaucher (Genf) an dieser Stelle meinen besten Dank aus. Für einen Arbeitsplatz im
Naturhistorischen Museum von Genf, wo die Bearbeitung des Materials z.T. erfolgte,
wird der Direktion sowie Herrn Dr. Cl. Vaucher bestens gedankt. Ferner gebührt mein
! Regenwiirmer aus Südamerika 23.
2 Vor Abschluss des Manuskriptes wurde mir von Herrn Dr. J. Römbke weiteres, aus
Bolivien vermitteltes, Material zur Bestimmung überlassen. Ferner übersandte mir Herr Dr.
Römbke auch den Entwurf eines Manuskriptes in dem die bisherigen Funde terrestrischer
Oligochaeten aus Bolivien zusammengefasst werden. Es wird die Bitte geäussert, die in
meinem Manuskript enthaltenden Informationen über bolivianische Regenwürmer in sein
Manuskript einbauen zu können. Der Bitte wird Folge geleistet, für die Zusendung neuen
Materials sei auch an dieser Stelle gedankt.
Manuskript angenommen am 13.10.1994.
586 ANDRAS ZICSI
Dank fiir die Uberlassung von Typenmaterial den Herren Prof. Dr. M. Dzwillo,
Zoologisches Institut und Museum, Hamburg und Dr. A. Rolando, Museo ed Istituto
di Zoologia Sistematica della Universita, Torino.
Die Buchstaben AF und Z beziehen sich auf die Sammlung des Zoo-
systematischen und Ökologischen Instituts der Eötvös Loränd Universität, Budapest
und G auf die Sammlung des Naturhistorischen Museums, Genf.
BESCHREIBUNG DER ARTEN
Glossoscolecidae Michaelsen, 1900
Enantiodrilus Cognetti, 1902
Enantiodrilus borellii Cognetti, 1902
Von dieser hologynen Art (2 Paare Ovarien und Eitrichter im 12. und 13. Seg-
ment), die bisher nur aus Argentinien (COGNETTI, 1902a) und Brasilien (MICHAELSEN,
1902) von je einem Fundort erwähnt wurde, liegen mir über 100 Exemplare aus dem
bolivianischem Guayaramerin (Prov. Beni) vor. Die Tiere sind in verschiedenen
Entwicklungsstadien, die meisten sind jedoch adult und besitzen einen gut ent-
wickelten Gürtel. Wie bekannt, ist es die einzige Art innerhalb der Glossoscolecidae,
die über zwei Paare Ovarien und Eitrichter verfügt. Die Beschreibungen von
COGNETTI (1902 a, b, 1905) und die ergänzenden Angaben von Michaelsen (1918)
sind sehr ausführlich und ermöglichen eine sofortige Identifizierung dieser beson-
deren, phylogenetisch alleinstehenden Art.
Die von mir untersuchten Exemplare stimmen in allen wesentlichen Merk-
malen mit der Originalbeschreibung überein und besitzen auch den von MICHAELSEN
(1918) ausführlich beschriebenen handschuhfingerförmigen Anhang der Chylus-
taschen (Abb. 1). Dieser Anhang wird von COGNETTI (1902 b: Fig.12) weder erwähnt
noch abgebildet.
Einen deutlichen Unterschied weist die Form der Samentaschen bei meinen
Exemplaren auf, die bei den einzelnen Populationen eine verschiedene Ausbildung
zeigt. Bei den Exemplaren aus der Umgebung der Ziegelbrennerei bei Guayaramerin
weichen sie am meisten von denen der Originalbeschreibung (COGNETTI, 1902 b: Fig.
3) und denen vom Fundort Estancia Esperanza bei Guayaramerin ab. Hier sind es nur
einfache Einstülpungen der Innenwand mit einem kleinen ampullenförmigen Anhang
(Abb. 3). Dieser Anhang vergrössert sich bei anderen Tieren der Estancia Esperanza
zu taschenförmigen Ampullen (Abb. 4) oder zu warzenförmigen Wucherungen, die
genau das Aussehen der in der Originalbeschreibung angeführten Samentaschen
besitzen (Abb. 5). Es könnte angenommen werden, dass die auf Abb. 3 angeführte
Form der Samentaschen eine Rückbildung dieser Organe darstellt, da ansonst die
Tiere mit einem deutlichen Gürtel versehen sind und die Ovarien mit Eiern besetzt
sind. Dieser Annahme spricht jedoch die Beobachtung entgegen, dass bei juvenilen
Tieren von der Estancia Esperanza, deren adulte Exemplare die typische Samen-
REGENWURMER AUS BOLIVIEN 587
ABB. 1-6
Enantiodrilus borelli Cognetti, 1902. 1. Chylustasche mit weiblichen und männlichen Ge-
schlechtsorgan. H = Hoden, HT = Hodentrichter, O = Ovarien, OT = Ovarientrichter, CT =
Chylustasche, ACH = Handschuhförmiger Anhang der Chylustasche. 2.Ventralansicht mit dem
Geschlechtsfeld.PF = Pubertätsflecke. 3-6.Verschiedene Entwicklungsstadienn der Samen-
taschen.A = Ampulle.
588 ANDRAS ZICSI
taschenform aufweisen, die Samentaschen das Aussehen der Abb. 3 besitzen. Die
deutlichen Unterschiede der Samentaschenformen sind bloss ein verschiedenes Ent-
wicklungsstadium dieser Organe und reichen, wie dies am Serienmaterial eindeutig
nachgewiesen werden konnte, nicht zur Aufstellung neuer Taxa aus (Abb. 6).
Als weiterer Unterschied meiner Tiere können ausserdem die in der ventralen
Medianlinie verlaufenden unpaarigen Pubertätsflecke vom 10., 11. - 15., 16.und 17.
Segment erwähnt werden, die bei allen Exemplaren der 3 Fundorte erkannt werden
konnten. Ferner konnte bei einigen Exemplaren eine deutliche Samenrinne nach-
gewiesen werden, mit der die beiden Kopulationstaschen verbunden sind.
Fundorte: AF/1189 32 adulte und 42 praead. Ex., G/INVE 17975 3 Ex., Prov. Beni,
Guayaramerin Umgebung der Ziegelbrennerei, 4. XII. 1966, leg. Zicsi. - AF/2835, 2846, 2847
65 adulte, 23 praead. und 18 juv. Ex., G/INVE 17976 3 Ex., Guayaramerin, Estancia
Esperanza, Galeriewald am Rande des Mamaore Flusses 30. XI. 1966, leg. Zicsi. AF/2851 14
Juv. Ex., Prov. Beni, Guayaramerin, Insel Nicolas Suarez, 21. XI. 1966, leg. Zicsi. - AF/2860-
61 33 juv. Ex., Prov. Beni, Insel Nicolas Suarez, 27. XI. 1966, leg. Zicsi. - AF/2882 7 Ex.,
Prov. Beni, Esperitu, Uberschwemmungsgebiet, 10. V. 1985, leg. Rombke. - AF/2883 5 Ex.,
Esperitu, trockene Steppe, 10. V. 1985, leg. Rômbke. - AF/2884 2 Ex., Esperitu, 16. IV. 1983,
leg. Römbke. - AF/ 2886 6 Ex., AF/2887 4 Ex., Prov. Ballivian, Esperitu 170 m, 27. II. 1993,
16. IV. 1985, leg. Hanagarth. - AF/2888 2 Ex., Dept. Beni, Quiqusbey 300 m, 10. III. 1988, leg.
Hanagarth.
Andiorrhinus Cognetti, 1908 emend. Righi, 1993
Neuerdings wurde diese artenreiche Gattung von RIGHI (1993) aufgrund der
Zahl und Lage der Intestinalherzen in 4 Untergattungen geteilt (Amazonidrilus 4
Paare Intestinalherzen im 10.-13. Segment, Andiorrhinus 2 Paare Intestinalherzen im
11.-12. Segment, Turedrilus 2 Paare Intestinalherzen im 10. und 11. Segment und
Meridrilus 2 Paare Intestinalherzen im 12. und 13. Segment).Bei der Typusart von
Andiorrhinus, A.salvadori, wurde von Righi nur angenommen, dass sie tiber zwei
Paare Intestinalherzen verfiigt, da in der Originalbeschreibung die Lage der letzten
Intestinalherzen im 12. Segment angegeben ist, ohne andere Intestinalherzen
anzuführen.
Da es mir gelungen ist das Typenmaterial dieser Art im Museum von Torino
(Inv. Nr. OL. 30) zu iberpriifen, kann die Annahme von Righi an dieser Stelle
bestätigt werden. Von den drei Exemplaren, liess sich bei einem aufgeschnittenen
Tier die Lage der Intestinalherzen im 11. und 12. noch genau erkennen.
Da durch die Unterteilung der Gattungen ein Zurechtfinden erleichtert wird,
schliesse ich mich der Einteilung Righis an.
Andiorrhinus (Amazonidrilus) bolivianus sp. n.
Lange des Holotypus 218 mm, Breite 9 mm, Segmentzahl 230. Lange der
Paratypen, adultes Tier 220 mm, Breite 8,9 mm, Segmentzahl 235, Lange der
juvenilen Tiere 55-120 mm, Breite 3,5 mm, Segmentzahl 169-189.
REGENWURMER AUS BOLIVIEN 589
Farbe grau. Kopf eingezogen, 1.-2. Segment verwachsen. Vordere Segmente
ungeringelt. Borsten gepaart, Borsten ab etwas grösser als cd, Borsten aa etwas
grösser als bc. Borstendistanz hinter dem Gürtel aa: ab: bc: cd: dd wie 6,25: 1,25:
8,75: 1: 22,5. Borsten ab vom 6. Segment, Borsten cd vom 9. Segment zu erkennen.
Dorsoventrale Borsten des 7. und 8. Segmentes zu Geschlechtsborsten umgewandelt.
Ventrale Borsten des 20.-22. Segmentes ebenfalls zu Geschlechtsborsten umge-
wandelt und vor den Nephridialporen stehend. Nephridialporen am ganzen Kôrper in
der Borstenlinie d.
Samentaschenporen in Intersegmentalfurche 7/8 und 8/9 in der Borstenlinie cd,
deutliche Offnungen die von einem Hof umrandet sind.
Gürtel sattelförmig vom 17.-24., 1/2 25. Segment. Pubertätsfeld vom 20.-22.
Segment (Abb. 7). Weibliche Poren auf dem 18. Segment. Männliche Poren in Inter-
segmentalfurche 20/21.
Innere Organisation. Dissepimente 6/7-11/12 stark verdickt, 12/13-14/15
schwach verdickt. Schlunddriisen bis ins 6. Segment reichend. Muskelmagen im 6.
Segment. Kalkdriisen im 7.-9. Segment, mit kurzem Stiel versehene Lamellentaschen
ohne Anhang. Zahl der Lamellen tiber 30. Lateralherzen im 7.-9. Segment. Intes-
tinalherzen im 10.-13. Segment. Hoden und Samentrichter im 10. und 11. Segment,
frei. Samensäcke im 11. und 12. Segment, klein. Ovarien im 13. Segment, Ovarient-
richter auf 13/14, Ovarienleiter die Dissepimente 14/15-17/18 durchbrechend im 18.
ABB. 7-8
Andiorrhinus (Amazonidrilus) bolivianus sp. n. 7.Ventralansicht mit dem Geschlechtsfeld. DP
= Drüsenpapillen. 8.Samentaschen des 8. und 9. Segmentes. SK = Samenkämmerchen, A =
Ampulle.
590 ANDRAS ZICSI
Segment ausmiindend. Samenleiter verlaufen bis ins 20. Segment getrennt und ver-
schwinden dann in der Muskelwand und treten in Intersegmentalfurche 20/21 hervor.
Geschlechtsborsten des 20., 21. und 22. Segmentes in Borstensäcken. Lange der
Geschlechtsborsten 2,20-2,22 mm, Dicke 0,44 mm. Die Borsten sind auf allen Seiten
mit 19 Kerben. Geschlechtsborsten des 7. und 8. Segmentes kiirzer, 16,64 mm lang
und 0,41 mm dick, mit je 11 Kerben. Mitteldarm im 23. Segment beginnend und
gleich mit einer Typhlosolis versehen. Nephridien mit Nephridialblasen.
Samentaschen 2 Paare im 8. und 9. Segment mit muskulôsem Ausführungs-
gang der z.T. in der Muskelwand verborgen liegt. Diesem schliesst sich eine mit
Samenkämmerchen gefüllte Vorkammer an. Aus dieser geht eine herzförmige Am-
pulle hervor. Die Ampulle scheint leer zu sein (Abb. 8).
Die neue Art steht Andiorrhinus (Amazonidrilus) torquemadai Righi, 1984, am
nächsten. Unterscheidet sich jedoch von dieser durch die kiirzere Ausdehnung des
Giirtels, durch die Lage der weiblichen Poren, durch die Form der Samentaschen und
die Zahl der Kerben der Geschlechtsborsten.
Fundort. Holotypus AF/2837 Prov. Beni. Guayaramerin Estancia Esperanza,
30. XI. 1966, leg. Zicsi. Paratypus AF/2838 1+3 juv. Ex., Fundort wie Holotypus.
Andiorrhinus (Amazonidrilus) c.f. holmgreni Michaelsen, 1918
Von dieser vorläufig zu holmgreni gestellten Art liegt mir ein sehr gut ent-
wickeltes, mit Giirtel versehenes Exemplar vor. Wie aus der Literatur ersichtlich, soll A.
(A.) holmgreni Michaelsen 1918, A. (A.) paraguayensis (Rosa, 1895) und A. (A.)
evelineae Righi, 1986, nahe stehen. Leider sind jedoch A. (A.) paraguayensis und A.
(A.) holmgreni aufgrund preadulter Tiere beschrieben worden, bei A. holmgreni fehlt
sogar jegliche Angabe der vermutlichen Giirtelausdehnung. Mein Exemplar unter-
scheidet sich in einigen Kennzeichen von allen drei Arten, doch glaube ich, dass diese
zur Aufstellung einer neuen Art nicht ausreichen. Da holmgreni ebenfalls aus Bolivien
beschrieben wurde, leider jedoch ohne nähere Angaben des Fundortes, stelle ich mein
Exemplar zu dieser Art und gebe eine ausfiihrliche Beschreibung davon. Zum
Vergleich werden auch einige Bestimmungsmerkmale der Art holmgreni erwähnt.
Lange 165 mm, Dicke 10 mm, Segmentzahl 179.
Farbe grau, Kopf eingezogen, 1.-2. Segment verwachsen. Segmente un-
geringelt. Borsten eng gepaart. Ventralborsten am Vorderkörper vom 3.-4. Segment,
cd-Borsten nur hinter dem Gürtel beginnend, erkannt. Bei holmgreni sind keine cd-
Borsten beschrieben worden, bei paraguayensis sind ab und cd Borsten am ganzen
Körper vorhanden, doch sollen diese sehr klein sein.
Borstendistanz hinter dem Gürtel aa: ab: bc: cd: dd: wıe 12,5: 1,5: 20: 1: 60.
Nephridialporen in der Borstenlinie cd, am Vorderkörper im vorderen Drittel der
mutmasslichen Borstenlinie cd.
Samentaschenporen in Intersegmentalfurche 6/7-8/9 in der angenommenen
Borstenlinie cd, vor der der Nephridialporen. Weibliche Poren auf dem 17. Segment,
männliche Poren auf dem 20. Segment. -
REGENWURMER AUS BOLIVIEN 591
Gürtel sattelförmig, stark drüsig vom 16.-26. Segment; Segment 15 und ein
Teil des 27. Segmentes ebenfalls verfarbt, doch nicht driisig angeschwollen. Puber-
tätsstreifen vom 20.-24. Segment. Borsten ab des 17.-23. Segmentes von Papillen
umgeben und in Geschlechtsborsten umgewandelt. Bei holmgreni stehen die ab
Borsten des 18., 19.und 23. Segmentes auf Papillen und sind zu Geschlechtsborsten
umgewandelt. Undeutlich begrenzte Driisenverdickungen in der Region der Samen-
taschen, wie dies bei holmgreni erwähnt wurde, konnten nicht erkannt werden.
Innere Organisation. Dissepimente 6/7-14/15 deutlich verdickt. Pharingialer
Bulbus ohne Schleimdriisen, vorderer Teil des Oesophagus von Schleimdriisen und
mächtigen Peptonephridien umgeben. Peptonephridien bis ins 15. Segment, in jedem
Segment vorhanden. Nephridien im hinteren Teil des Kôrpers mit Endblasen ver-
sehen. Mächtiger Muskelmagen im 6. Segment. Chylustaschen im 7.-9. Segment,
entspringen dorsal und hängen lateral in die Kôrperhôhle. Es sind stiellose Lamellen-
taschen am Ende ohne Verschniirung. Es konnten ungefähr 30 Lamellen gezählt
werden. Lateralherzen im 7.-9. Segment. Intestinalherzen im 10.-13. Segment. Hoden
und Samentrichter im 10. und 11. Segment, sie sind in perioesophageale Testikel-
blasen eingeschlossen. Die Testikelblasen schliessen auch die Herzen des 10. und 11.
Segmentes sowie die Samensäcke des 11. Segmentes ein. Zwei Paare Samensäcke im
11. und 12. Segment. Ovarien im 13. Segment, Ovarienleiter durchbrechen die zu-
sammengedrückten und vom Muskelmagen nach hinten verschobenen Dissepimente
13/14-16/17 und treten im 17. Segment aus. Samenleiter sind in der Muskelwand
eingebettet und verlaufen bis zum 20. Segment, wo sie dann austreten. Geschlecht-
borsten vom 17.-23. Segment. Die Borsten stehen in Borstensäcken, die an der
Innenwand befestigt sind. Länge der Geschlechtsborsten 3-3,5 mm, sie sind mit in 4
regelmässigen Längsreihen angeordneten Narben ornamentiert. Es konnten bis 26-27
Narben gezählt werden. Mitteldarm beginnt im 23. Segment, Typhlosolis vorhanden.
Samentaschen im 7., 8. und 9. Segment, bestehen aus einem kurzen Aus-
führungsgang, der durch eine mässige Verengung in eine Ampulle übergeht. Die
Ampulle ist mit Samenkämmerchen und Samenmassen gefüllt (Abb.9). Samen-
taschendrüsen, die bei holmgreni angegeben sind, konnten nicht erkannt werden.
Pubertätsstreifen vom 20.-24. Segment. Borsten ab des 17.-23. Segmentes von
Papillen umgeben und in Geschlechtsborsten umgewandelt. Bei holmgreni stehen die
ab Borsten des 18., 19., 23. Segmentes auf Papillen und sind zu Geschlechtsborsten
umgewandelt. Undeutlich begrenzte Drüsenverdickungen in der Region der
Samentaschen, wie dies bei holmgreni erwähnt wurde, konnten nicht erkannt werden.
Aufgrund des vorliegenden Materials steht c.f. holmgreni dem A. (A.)
evelineae Righi, 1986, am nächsten, unterscheidet sich jedoch von ihm deutlich in der
Lage des Gürtels, in der Zahl der Geschlechtsborsten tragenden Segmente am Gürtel
und zu einem gewissen Grade auch in der Form der Samentaschen. Es ist jedoch nicht
ausgeschlossen, dass holmgreni, c.f. holmgreni und evelineae mit paraguayensis
identisch sind und so mit dieser zu synonymisieren wären. Leider ist bei keiner der
bisher beschriebenen drei Arten die Lage der weiblichen Poren angegeben, die bei c.f.
holmgreni im 17. Segment erkannt werden konnten.
Fundort AF/2843 1 Ex., Prov. La Paz, Unduavi, 3800 m, 20. XII. 1966, leg. Zicsi.
592 ANDRAS ZICSI
ABB. 9
Andiorrhinus (Amazonidrilus) c.f. holmgreni Michaelsen, 1918. 9.Samentaschen des 7., 8. und
9. Segmentes. SK = Samenkämmerchen.
Andiorrhinus (Andiorrhinus) montanus sp. n.
Von den 6 bisher bekannten Arten der Untergattung Andiorrhinus sind bisher 3
aus Venezuela und 3 aus Brasilien beschrieben worden. Soweit Höhenangaben vorlie-
gen, stammt allein A. (A.) venezuelanus aus einer grösseren Höhe (3000 m). Das mir
aus Bolivien vorliegende Tier wurde von mir ebenfalls in einer Höhe von 3900 m
erbeutet.
Die in der Literatur angeführten Merkmale zur Unterscheidung der Arten sind
sehr bescheiden. Der Gürtel erstreckt sich bei allen Arten vom 16., 17.-25. Segment
und auch die Pubertätsstreifen sind auf dem 1/2 19., 20-22., 23. Segment gelegen.
Weitere Merkmale, wie Geschlechtsborsten tragende Segmente sind im Bereich der
Samentaschen und in der Gürtelregion bei allen Arten angeführt. Ob die Angaben
über Fehlen oder Vorhandensein von Samensäcken richtig sind, ist fraglich, da diese
mit den Testikelblasen verwachsen zu sein scheinen (A. (A.) salvadori Cognetti) und
so nicht eindeutig erkannt werden können. Auch die 3 Paare Samentaschen scheinen
bei allen Arten konstant zu sein, wenn auch bei A. (A.) muku Righi (1989) nicht im 7.,
8. und 9. Segment sondern im 6., 7. und 8. Segment gelegen. Nur in der Form und
Gestalt der Samentaschen lassen sich Unterschiede erkennen. Obwohl die Unter-
schiede meines Exemplares zu denen der bisher beschriebenen Arten gering sind,
betrachte ich mein vollkommen entwickeltes Tier aufgrund seines von den anderen
weit entfernten Fundorts als eine für die Wissenschaft neue Art.
REGENWURMER AUS BOLIVIEN 593
ABB. 10-11
Andiorrhinus (Andiorrhinus) montanus sp. n. 10. Ventralansicht mit Giirtel und Pubertats-
streifen. DP = Driisenpapillen. 11. Samentaschen des 7., 8. und 9. Segmentes. SK = Samen-
kämmerchen.
Länge 140 mm, Dicke am Gürtel 10 mm, hinter dem Gürtel 6 mm, Segment-
zahl 97 (einige Schwanzsegmente fehlen).
Farbe grau, auf der Dorsalseite vielleicht rot gewesen. Kopf rüsselförmig, 1-2.
Segment verwachsen. Borsten ab vom 6. Segment, cd hinter dem Gürtel beginnend,
erkannt. Borsten ab des 7. und 8. Segmentes zu Geschlechtsborsten verwandelt, sind
von kleinen Papillen umgeben. Nephridialporen in der Borstenlinie cd.
Gürtel sattelförmig vom 16.-25. Segment, Pubertätsstreifen vom 19.-23. Seg-
ment. Borsten ab des 18. und 19. stehen auf kleinen Papillen, die des 20.-23.
594 ANDRAS ZICSI
Segmentes sind von drüsiger Struktur umgeben. Sämtliche Borsten der angeführten
Segmente sind zu Geschlechtsborsten verwandelt. Die Giirtelregion ist stark aus-
gebreitet, Intersegmentalfurchen 19/20, 20/21 und 21/22 mit deutlichen Vertiefungen
(Abb. 10.), die dem Geschlechtsfeld ein charakteristisches Aussehen verleihen. Weib-
liche Poren auf dem 17. Segment hinter der Borstenreihe ab, in Linie der Borste b.
Männliche Poren auf Intersegmentalfurche 20/21 (bei A. (A.) rubescens Michaelsen
(1926) konnten die von Michaelsen nicht erkannten weiblichen Poren ebenfalls auf
dem 17. Segment nachgewiesen werden).
Innere Organisation. Dissepimente 6/7-12/13 deutlich ausgebildet, doch nicht
verdickt. Schlund ohne Schlunddriisen, Oesophagus langgestreckt, von zwei
mächtigen Peptonephridien umgeben. Muskelmagen im 6. Segment, weit nach hinten
verzogen und so im 14.-16. Segment liegend. Drei Paare Chylustaschen im 7.-9.
Segment, dorsoventral mit einem kurzen Stiel an das Blutgefass geheftet, ventral
ebenfalls mit dem Blutgefass verbunden, ohne abgeschniirtes Ende. Typische Lamel-
lentaschen mit ungefähr 25 Lamellen ausgebildet. Lateralherzen im 7.-9. Segment,
Intestinalherzen im 11. und 12. Segment. Hoden und Samentrichter im 10. und 11.
Segment, in perioesophageale Testikelblasen eingeschlossen, die dorsal miteinander
verschmolzen sind. Sie schliessen so die Herzen und Samensäcke des 11. Segmentes
ein. Zwei Paare lappenförmige Samensäcke im 11. und 12. Segment. Ovarien länglich
im 13. Segment, Ovarienleiter durchbrechen die zusammengedrängten Dissepimente
13/14-16/17 und münden im 17. Segment aus. Samenleiter verlaufen parallel vom 18.
Segment an der Innenwand entlang und verschwinden in Höhe des 20/21 Segmentes
in der Leibeswand. Mitteldarm im 23.-25. Segment beginnend, Typhlosolis vor-
handen.
Die Geschlechtsborsten des 7. und 8. Segmentes sind nicht von Geschlechts-
borstendrüsen umgeben wie dies bei A. (A.) rubescens Michaelsen, 1926, und A. (A.)
pictus Michaelsen, 1926, der Fall ist. Es sind einfache, kleine Borstensäcke, die nicht
an die Leibeswand angeheftet sind. Die Geschlechtsborsten der Gürtelregion sind in
Borstensäcken und münden in einen länglichen Muskelstrang ein der ental an die
Innenseite der Leibeswand angeheftet ist und lateral verschieden hoch sein kann,
entsprechend den Einbuchtungen auf Intersegmentalfurche 19/20, 20/21 und 21/22.
Es sind also nicht Drüsenzellen, die die Geschlechtsborstensäcke umgeben wie bei
rubescens, sondern Muskelstränge wie von pictus beschrieben. Geschlechtsborsten
des 7. Segmentes 2,2 mm lang, mit 11 Kerben, Geschlechtsborsten der Gürtelregion
5,0 mm lang, mit 20 Kerben. Nephridien im Vorderkörper Peptonephridien, im
hinteren Teil des Körpers mit einer Endblase.
Samentaschen im 7., 8. und 9. Segment mit muskulösem breiten Ausführungs-
gang der in eine Ampulle übergeht. Ampulle mit kleinen Samenkämmerchen dicht
besetzt. Ausführungsgang mit einigen kugelförmigen Kämmerchen versehen (Abb.
IN)
Wie vorher erwähnt, wurde das Typenmaterial von A.(A.) salvadori Cognetti,
1908, überprüft. An dieser Stelle sei erwahnt, dass auch das Typenmaterial von A.
(A.) rubescens Michaelsen, 1926 (Inv. Nr. V.9809 Brasilien, Staat Manaus, Mana-
capuru am Rio Amazonas, VII.-VIH. 1924, leg. W. Ehrhardt) eingesehen wurde. Es
REGENWURMER AUS BOLIVIEN 595
konnte nachgewiesen werden, dass die von RIGHI (1993) als fraglich betrachtete Lage
der Intestinalherzen im 11. und 12. Segment nun erwiesen wurde.
Die neue Art steht A. (A.) rubescens am nächsten, unterscheidet sich von
diesem jedoch durch die inneren Geschlechtsdriisen im 7., 8. und 18.-23. Segment
sowie die Form und Gestalt der Samentaschen.
Fundort. Holotypus AF/2844 Prov. La Paz, Unduavi 3900 m, 20. XII. 1966, leg. Zicsi.
Andiorrhinus (Andiorrhinus) sp. juv.
Fundort AF/2890 1 Ex., Prov. Larecoja, Mapiri 1. VII. 1992, leg. Arce.
Tamayodrilus gen. n.
Typusart: Tamayodrilus roembkei sp. n.
Gattungsdiagnose. Normale Borsten in 8 Längslinien. Männliche Poren
intraclitellial. Vordere Dissepimente schwach verdickt. 6 Paare Chylustaschen im 10.-
15. Segment, verzogene Rispenschlauchtaschen. Geschlechtsapparat holoandrisch und
metagyn. Samensäcke im 11. und 12. Segment, kurz. Samentaschen vorhanden.
Die Gattung Tamayodrilus unterscheidet sich von allen tibrigen Gattungen mit
holoandrischem und metagynem Geschlechtsapparat durch die Zahl und Anordnug
der Chylustaschen.
Tamayodrilus roembkei sp. n.
Holotypus: Lange 200 mm, Dicke 10 mm, Segmentzahl 205. Paratypus: Lange
201 mm, Dicke 9,8 mm, Segmentzahl 201.
Farbe grau, Kopf eingezogen, 1.-2. Segment verwachsen. Borsten ab vom 3.
Segment, Borsten cd vom 6. Segment beginnend, erkannt. Borsten ab etwas grösser
als cd, Borsten aa gleich bc. Borstendistanz hinter dem Giirtel aa: ab: be: cd: dd wie
10: 1,3: 10: 1: 40. Segmente bis zum 8. Segment ungeringelt, vom 9. Segment doppelt
geringelt. Borsten ab vom 13.-19. Segment auf Papillen angeordnet und zu Ge-
schlechtsborsten umgewandelt.
Weibliche Poren auf dem 14. Segment zwischen der Borstenlinie bc. Männ-
liche Poren auf dem 20. Segment. 3 Paare Samentaschenporen auf Intersegmental-
furche 6/7-8/9, in der Borstenlinie cd gelegen.
Gürtel sattelförmig vom 13.-25. Segment, auf dem 26. Segment nur dorsal eine
diinne Verdickung zu erkennen. Pubertatsstreifen vom 1/4 19.-2/3 25. Segment (Abb.
12).
Innere Organisation. Dissepimente 6/7-10/11 mässig verdickt Schlund im 3-4.
Segment ohne Speicheldriisen. Diesem folgt ein gewundener Teil des Oesophagus der
beiderseits mit Speicheldriisen besetzt ist. Grosser Muskelmegen im 6. Segment.
Lateralherzen im 7.-9. Segment, Intestinalherzen im 10. und 11. Segment. 6 Paare
Chylustaschen im 10.-15. Segment, entspringen dorsolateral und sind der Bauchseite
596 ANDRAS ZICSI
ABB. 12-13
Tamayodrilus roembkei sp. n. 12. Ventralansicht mit Giirtel und Pubertätsstreife. DP
Driisenpapillen, PS = Pubertatsstreifen. 13. Samentaschen des 7., 8. und 9. Segmentes. A
Ampulle.
zu gerichtet. Es sind schinkenförmige Gebilde, die am Ende keine Verschnürung
besitzen. Die innere Struktur der Chylustaschen deuten auf etwas verzogene Rispen-
schlauchtaschen hin. Hoden und Samentrichter im 10. und 11. Segment, Samensäcke
im 11. und 12. Segment, klein. Der Mitteldarm beginnt im 17., Typhlosolis im 26.
Segment. Peptonephridien bis ins 15. Segment reichend, von da Nephridien mit
Blindsack und Nephridialblase.
Die Geschlechtsborsten sind innen von mächtigen drüsigen Zellen vom 13.-19.
Segment umgehen, Pubertätsstreifen sind ebenfalls durch eine drüsige Struktur von
innen markiert. Samenleiter verlaufen an der Innenwand deutlich bis zum 20.
Segment.
Samentaschen 3 Paare im 7., 8. und 9. Segment. Es sind längliche Gebilde mit
löffelartiger Ampulle, und sehr langem Stiel. Ampulle mit Samenmasse gefüllt (Abb.
IS):
Die neue Art wird Herrn Dr. J. Römbke (Frankfurt am Main) zu Ehren be-
nannt, der mir das Material zur Bestimmung überlassen hat.
Fundort. Holotypus AF/2885 Dpto. de La Paz, Prov. Franz Tamayo, Serrania Macho
Pelechuco, 4060 m, 28. X. 1982, leg. J. P. Arce. Paratypus AF/2891 | praead. Ex., Fundort wie
Holotypus.
REGENWURMER AUS BOLIVIEN 597
Goiascolex Righi, 1971
Goiascolex vanzolinii Righi, 1984
Obwohl nur giirtellose Tiere vorliegen, besteht kein Zweifel, dass es sich um
eine Art der Gattung Goiascolex handelt. Die unpaarigen männlichen Poren, die drei
Paare Chylustaschen (Kompositenschlauchtaschen), der holoandrische und metagyne
Geschlechtsapparat sowie Kopulationstaschen und das Fehlen der Samentaschen,
ermöglichen ein sicheres Einreihen zur Gattung Goiascolex. Von den bisher be-
schriebenen Arten G. cabrelli Righi, 1971, G. pepus Righi, 1972, G. edgardi Righi,
1986, und G. vanzolinii Righi, 1984, scheint sie der letzten am nähesten zu stehen. Da
die Originalbeschreibung nur aufgrund eines Exemplars erfolgte, gebe ich eine
Beschreibung meiner Tiere.
Lange 85-130 mm, Dicke 6-8 mm, Segmentzahl 148-178.
Kopf eingezogen rüsselförmig. 1.-2. Segment verwachsen. Borsten am ganzen
Körper eng gepaart. Borstenverhältnis am 32. Segment aa: ab: bc: cd: dd wie 15: 2: 20
: 1: 47,5. Borsten ab vom 13. Segment, cd vom 23. Segment beginnend, erkannt.
Borsten des 17. 18., 20., 21. und 29.-32. Segmentes auf kleinen Drüsenpapillen an-
geordnet, zu Geschlechtsborsten verwandelt. Bei G. vanzolinii Geschlechtsborsten
vom 16.-18. und 20.-28. Segment vorhanden. Geschlechtsborsten mit Kerben in
Reihen angeordnet, es konnten bis zu 7 Kerben gezählt werden. Weibliche Poren auf
dem 14. Segment, männliche Poren auf dem 19. Segment, y-förmiger Schlitz (bei
vanzolinii sind die weiblichen Poren nicht erkannt worden).
Innere Organisation. Verdickte Dissepimente 6/7-11/12 stark, 12/13-14/15
schwächer verdickt. Lateralherzen im 7.-9. Segment. Chylustaschen im 7.-9. Segment,
dorsolateral mit kurzem Stiel angeheftet, schinkenförmig ohne Abschnürung am Ende.
Kompositenschlauchtaschen. Hoden und Samentrichter im 10. und 11. Segment, frei.
Samensäcke im 11. und 12. Segment. Ovarien im 13. Segment. Kopulationstaschen im
19. Segment, rundliche Gebilde die auch etwas auf das 18. und 20. Segment übergehen.
Ventral sind sie durch ein verdicktes Drüsenfeld miteinander verbunden. Samenleiter
treten hinter Dissepiment 14/15 hervor und laufen der Innenwand angeschmiegt bis
zum 18. Segment wo sie in die Kopulationstaschen eintreten und dort im halbkreis ver-
laufend bis zur Mitte des 19. Segmentes weiter zu verfolgen sind. Mitteldarm im 15.-
16. Segment beginnend, Typhlosolis und Nephridialblasen vorhanden. Samentaschen
fehlen.
Obwohl meine praeadulten Tiere einige Abweichungen zu G. vanzolinii
aufweisen, werden sie wegen Form und Lage der Kopulationstaschen und deren Ge-
schlechtsborsten-Anordnung dieser Art zugeordnet. Die deutlichen Geschlechts-
borsten des 29.-32. Segmentes wurden von RIGHI (1984b) nicht erwähnt.
Fundorte. AF/2839, AF/2840 2 praead. und 2 juv. Ex., Guayaramerin, Nueva Cuba, 24.
und 28. XI. 1966, leg. Zicsi.
Diachaeta Benham, 1886
Sämtliche auf der Insel Nicolas Suarez bei Guayaramerin gesammelten Tiere
von Diachaeta waren so jung, dass sie nicht zur Art bestimmt werden konnten.
598 ANDRAS ZICSI
Fundort. AF/2849, AF/2862 37 juv. Ex., Nicolas Suarez, 21. und 27. XI. 1966, leg.
Zicsi.
Inkadrilus Michaelsen, 1918
Da es sich um den ersten Fund einer /nkadrilus-Art seit 1900 handelt, sollen
hier die Kenntnisse über diese Gattung zusammengefasst werden.
MICHAELSEN (1918) ordnete der Untergattung /nkadrilus Arten aus der
Sammelgattung Thamnodrilus (=Martiodrilus) mit Fachkapseltaschen und Saum-
leistentaschen zu. Wie bekannt besitzen die Arten der Gattung Martiodrilus 7-8 oder
6-8 Chylustaschen. Bei den meisten dieser Arten handelt es sich im Bau der
Chylustaschen um Kompositenschlauchtaschen, die sich zu Wabentaschen verein-
fachen kônnen (MICHAELSEN, 1918). Später wurden von MICHAELSEN (1935) die der
Untergattung /nkadrilus zugeordneten Arten in zwei selbstständige Gattungen ge-
stellt. Für Rhinodrilus (Thamnodrilus) cameliae Michaelsen, 1913, mit 8 Paar im 7.-
14. Segment liegenden Chylustaschen, Fachkapseltaschen, wurde die Gattung Quim-
baya errichtet und fiir Anteus aberratus Michaelsen, 1900, und Anteus octocystis
Michaelsen, 1900, mit 8 Paar Chylustaschen im 7.-14. Segment, Saumleistentaschen
wurde die Untergattung /nkadrilus zur Gattung erhoben (MICHAELSEN, 1935). O.
cameliae wurde in Kolumbien gesammelt, /. octocystis in Peru. Fiir /. aberratus
wurde von MICHAELSEN (1918, p. 57) die fragliche “Heimatsnote wahrscheinlich
Peru” beigelegt und dies wegen seiner nahen Verwandtschaft mit der peruanischen /.
octocystis. Bei der Revision der Gattung Aymara (Typusart A. voogdi Michaelsen)
fand ich in der Sammlung des Zoologischen Instituts und Museums von Hamburg
(Typenmaterial: Inv. Nr. 12305, Peru, Dept. Huanuco, 10-15 km N.O. von Chogosh,
9020” S.B. 75030’W.L.) auch zwei gut erhaltene Exemplare der /. aberratus. Durch
diesen Fund ist das Vorkommen von /. aberratus in Peru einwandfrei erwiesen. Die
mir zur Beschreibung vorliegende neue Art aus Bolivien (160 4’ 30°’ S.) weist darauf
hin, dass die Arten mit Saumleistentaschen und einer hohen Zahl von Chylustaschen
eine siidliche Verbreitung in den Kordilleren besitzen und die beiden schon bekannten
Arten in Peru den nördlichsten Vorposten darstellen.
Inkadrilus hanagarthi sp. n.
Es liegt ein gut entwickeltes, adultes Exemplar dieser Art vor.
Holotypus: Länge 160 mm, Dicke 9 mm, Segmentzahl 154.
Farbe am Vorderkörper grau-rot. Kopf eingezogen. 1.-2. Segment verwachsen.
Borsten am ganzen Körper eng gepaart. Borstenverhältnis hinter dem Gürtel aa : ab:
bc : cd: dd wie 15: 1,5 : 10: 1 : 27. Borsten ab vom 5., bc vom 6. Segment erkannt.
Borsten ab vom 19.-26. Segment von Drüsenpapillen umgeben und in Geschlechts-
borsten verwandelt. Geschlechtsborsten 1,5 - 1,7 mm lang und mit 10-11 Kerben in
jeder der 4 unregelmässigen Längsreihen. Nephridialporen in der Borstenlinie cd.
Samentaschenporen in Intersegmentalfurche 6/7 - 8/9 etwas unterhalb der Borsten-
linie cd.
REGENWÙRMER AUS BOLIVIEN 599
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Inkadrilus hanagarthi sp. n. 14.Ventralansicht mit Gürtel und Pubertätsstreifen. DP
Drüsenpapillen, PS = Pubertätsstreifen. 15.Samentaschen des 7., 8. und 9. Segmentes. A
Ampulle, SK = Samenkämmerchen.
Gürtel sattelförmig, stark drüsig vom 14. - 1/2 27. Segment. Pubertätsstreifen
vom 21. - 1/4 27. Segment. Weibliche Poren auf Intersegmentalfurche 14/15, auf der
Innenseite des 14. Segmentes, männliche Poren auf dem 21. Segment (Abb. 14.). Die
Poren konnten von Aussen nicht beobachtet werden, von Innen konnte der Eileiter
und Samenleiter verfolgt und die Ausmündungen erkannt werden.
Innere Organisation. Dissepimente 6/7 - 9/10 sehr stark verdickt. 10/11 und
11/12 schwach angedeutet. Schlund bis ins 4. Segment reichend. Mächtige Pepton-
ephridien im 5. Segment. Weitere Peptonephridien bis ins 12. Segment vorkommend,
von da Nephridien mit Blindsack und Nephridialblase versehen. Mächtiger Mus-
kelmagen im 6. Segment. Lateralherzen im 7. - 9. Segment, Intestinalherzen im 10.
und 11. Segment. 9 Paare Chylustaschen im 7. - 15. Segment, die des 7.-9. Segmentes
sehr klein, 10.-11. etwas grösser, vom 12. - 15. Segment gross, von fächerförmiger
Gestalt, Saumleistentaschen die dorsolateral am Darm angeheftet sind. Hoden und
Samentrichter im 10. und 11. Segment in oesophageale Testikelblasen eingeschlos-
sen. Samensäcke im 11. und 12. Segment, klein. Ovarien im 13. Segment, mit gros-
sem Ovariensack aus dem die Eileiter hervorgehen und im 14. Segment ausmünden.
Samenleiter treten im 11. und 12. Segment hervor und verlaufen an der Innenwand bis
600 ANDRAS ZICSI
zum 21. Segment. Mitteldarm im 16. Segment, Typhlosolis im 27. Segment beginn-
end.
Samentaschen drei Paare im 7., 8. und 9. Segment, es sind fadenfôrmige
Gebilde, die am Ende verschiedenartig angeschwollen und mit Samenmasse gefüllt
sind (Abb. 15).
Die neue Art steht /. aberratus und I. octocystis gleicherweise nahe, unter-
scheidet sich von beiden aber durch die 9 Paare Chylustaschen.
Die neue Art wird Herrn Dr. W. Hanagarth (Okologisches Institut, La Paz), der
mir einen Teil des Materials zur Bestimmung zugesandt hat, zu Ehren benannt.
Fundort. Holotypus AF/2892. Dept. La Paz Prov. Manco Capac, Comunidad
Copacabana, Kusijata, 3312 m, 160 4' 30 S 640 4' 48 O, 16. VII. 1991, leg. Fecho.
Martiodrilus Michaelsen, 1936
Aus Bolivien wurde bisher nur eine Martiodrilus-Art (M. bolivianus Righi &
Römbke, 1987) beschrieben. Bei dieser Art reichen die Chylustaschen, wie bei der
jetzt zur Beschreibung vorliegenden Art, bis ins 15. Segment. Also ein Segment
weiter nach hinten, als dies fiir die zahlreichen tibrigen Martiodrilus-Arten der Fall ist.
Im 7. Segment hingegen fehlen die Chylustaschen bei den beiden bolivianischen
Arten. Ein Fehlen der Chylustaschen im 7. Segment ist auch bei einer Gruppe von
Martiodrilus-Arten bekannt (M. ecuadoriensis und M. savanicola), doch besitzen
diese Arten nur 7 Paar Chylustaschen. Vorläufig soll dieser Verschiebung kein
supraspezifischer Wert beigemessen werden, da wir es in beiden Fallen nur mit je
einem Exemplar zu tun haben. Weiteres Material aus Bolivien ist notwendig, um die
phylogenetische Bedeutung der Verschiebung der Chylustaschen im südlichen Teil
der Kordilleren beurteilen zu können.
Martiodrilus silvestris sp. n.
Von der neuen Art liegt nur ein sehr erweichtes Exemplar vor.
Holotypus: Lange 100 mm, Dicke 7 mm, Segmentzahl 117.
Farbe dunkelrot, irisierend. Kopf kolbenförmig, eingezogen. Borsten am
ganzen Kôrper eng gepaart. Borstenverhältnis hinter dem Giirtel aa : ab : be : cd : dd
wie 10: 1 : 12 : 1 : 37. Borsten ab vom 8. - 14. und vom 19. - 20. Segment auf
Borstenpapillen angeordnet, zu Geschlechtsborsten verwandelt. Geschlechtsborsten
mit 20-23 Kerben versehen. Die meisten Borsten sind wegen der Erweichung des
Tieres ausgefallen. Nephridialporen in der Borstenlinie cd.
Weibliche Poren auf dem 14. Segment hinter der Borstenlinie b, männliche
Poren auf dem 20. Segment. Samentaschenporen in Intersegmentalfurche 6/7 - 8/9 in
der Borstenlinie cd.
Gürtel sattelförmig vom 14. - 26. Segment, Pubertätsstreifen vom 1/2 20. - 1/2
22. Segment (Abb. 16).
REGENWURMER AUS BOLIVIEN 601
ABB. 16-17
Martiodrilus silvestris sp. n. 16. Ventralannsicht mit Giirtel und Pubertätsstreifen. DP
Driisenpapillen, PS = Pubertatsstreifen. 17. Samentaschen des 7., 8. und 9. Segmentes. A
Ampulle.
Innere Organisation. Verdickte Dissepimente fehlen im vorderen Teil des
Körpers, dadurch Muskelmagen stark nach hinten verschoben, im 6. Segment liegend.
Chylustaschen im 7/8 - 14/15. Segment. Die im 8. und 9. Segment würstchenförmig,
die übrigen am Basalteil immer breiter werdend. Die Chylustaschen sind dorsolateral
am Darm angeheftet und besitzen einen kurzen Stiel. Es sind Kompositenschlauch-
taschen mit Übergang zu Wabentaschen. Hoden und Samentrichter im 10. und 11.
Segment in oesophageale Testikelblasen eingeschlossen aus denen die Samensäcke
des 11. und 12. Segmentes hervorgehen. Die Samensäcke sind längliche Gebilde, die
des 11. Segmentes nach vorne gerichtet und 3-4 Segmente einneimend, die des 12.
Segmentes nach hinten gerichtet und 6-7 Segmente einnehmend. Intestinalherzen im
10. und 11. Segment. Ovarien im 13. Segment. Vom 8. - 14. Segment und vom 19. -
20. Segment sind die Geschlechtsborsten von mächtigen Drüsenzellen umgeben.
Samenleiter verlaufen doppelt bis zum 20. Segment. Mitteldarm im 17. Segment
beginnend. Nephridien mit Blindsack und Nephridialblase.
Samentaschen im 7., 8. und 9. Segment. Es sind löffelartige Gebilde mit
langem Stiel. Ampulle voll mit Samenmasse (Abb. 17).
Die neue Art steht hinsichtlich der Form der Pubertätsstreifen M. jordani
(Rosa, 1895) am nächsten. Hinsichtlich der männlichen Geschlechtsorgane erinnert
sie an die Arten M. savanicola und M. agricola. Von allen unterscheidet sie sich
jedoch durch die bis ins 15. Segment reichenden Chylustaschenpaare.
602 D. BORCARD
Fundort. Holotypus AF/289, Cota Pata Dpto. de La Paz, Caja de Montana, 2900 m,
Urwald, 26. I. 1985, leg. Hanagarth.
Pontoscolex Schmarda, 1861
Pontoscolex corethrurus (F. Miiller, 1857)
Fundorte. AF/2845, AF/2852 47+6 juv. Ex., Guayaramerin, 18. XI. und 2. XII. 1966,
leg. Zicsi. - AF/2848 72 Ex., Guayaramerin, Insel Nicolas Suarez, 21. und 27. XI. 1966, leg.
Zicsi. - AF/2864 2 Ex., Plantage am Ufer des Mamore Flusses, Guayaramerin, 24. XI. 1966,
leg. Zicsi.
Periscolex Cognetti, 1905
Periscolex sp.
Die beiden winzigen Exemplare waren so erweicht, dass sie nicht bis zur Art
bestimmt werden konnten.
Fundort: AF/2897 2 Ex., Dept. Beni, Prov. Ballivian, Esperitu Rio Yacuma, 18. X.
1980, leg. Hanagarth.
Acanthodrilidae Claus, 1880
Microscolex Rosa, 1887 emend. Pickford, 1937
Microscolex dubius (Fletcher, 1887)
Fundort. AF/2833 103 Ex., Prov. La Paz, La Paz 3600 m, 25. XII. 1966 leg. Zicsi.
Microscolex phosphoreus (Ant. Diigès, 1837)
Fundort. AF/2834 3 Ex., Prov. La Paz, La Paz, 3600 m, 25. XII. 1966, leg. Zicsi.
Yagansia Michaelsen 1899 emend. Zicsi, 1989
Yagansia peruana Cernosvitov, 1939
Es sind bisher 3 Arten mit zwei Divertikeln der Samentaschen beschrieben
worden. Es sind dies Y. peruana Cernosvitov vom Titicaca-See, Y. chiloensis Michael-
sen und Y. parinacotana Zicsi aus Chile. Fiir die Revision von Yagansia lagen mir
keine Exemplare von Y. peruana vor und ich konnte auch das Typenmaterial dieser Art
nicht einsehen (Zıcsı, 1989). Obwohl bei den vorliegenden Exemplaren die muskulôse
Kammer an der Ausführungsstelle der männlichen Poren nicht erkannt werden konnte,
stelle ich die vorliegenden Tiere Y. peruana zu, da sie in der Ausdehnung der
Gürtelorgane, in der Form der Samentaschen und auch in der Form Geschlechts-
borsten mit dieser übereinstimmen.
ORIBATES D'UNE TOURBIERE JURASSIENNE 603
Fundort. AF/721 4., Ex., Prov. La Paz, zwischen Unduavi und Cota 4654 m, 29. XII.
1966, leg. Zicsi. - AF/719-720 7 Ex., Titicaca-See, beim Jachtklub, 3800 m, 28. XII. 1966, leg.
Zicsi - AF/2896 4 Ex. Prov. La Paz, Titicaca-See, 3850 m, 8. VIII. 1993, leg. Beck.
Octochaetidae Michaelsen, 1900
Dichogaster Beddard 1888
Dichogaster saliens (Beddard, 1893)
Fundorte. AF/2855, AF/2871 10 Ex., Prov. Beni, Guayaramerin, Park, 18. XI. und 2.
XII. 1966, leg. Zicsi. - AF/2850 2 Ex., Guayaramerin, Insel Nicolas Suarez, 21. XI. 1966, leg.
Zicsi. -AF/2865 5+9 juv. Ex., Guayaramerin, Plantage am Ufer des Mamore Flusses, 24. XI.
1966, leg. Zicsi.
Dichogaster affinis (Michaelsen, 1890)
Fundort. AF/2870 2 Ex., Prov. Beni, Guayaramerin, Park, 2. XII. 1966, leg. Zicsi.
Dichogaster modigliani (Rosa, 1886)
Fundort: AF/2889 2 Ex., Depto. de La Paz, Prov. Larecoja, Mabiri, 1. VII. 1982, leg.
Arce.
Ocnerodrilidae Beddard, 1891.
Eukerria Michaelsen, 1935
Da ich ausser der peregrin verbreiteten Art E. saltensis (Beddard, 1895) zum
ersten Mal jetzt auch anderen in Siidamerika vorkommenden Arten begegnet bin und
diese aufgrund der Originalbeschreibungen auch identifizieren konnte, befasse ich
mich an dieser Stelle nicht mit den zahlreichen, neuerdings beschriebenen Arten, die
sich nur unwesentlich von den alten Arten unterscheiden. Es soll jedoch betont
werden, dass die Verbreitung dieser im letzten Jahrhundert beschriebenen Arten durch
Verschleppung eine bedeutend grössere Ausdehnung besitzen, als bisher ange-
nommen worden ist. Ein Vergleich der neu beschriebenen Taxa mit den Typen der
alten Arten ist unbedingt erforderlich.
Eukerria eiseniana (Rosa, 1895)
Fundorte. AF/2853 16 Ex., Prov. Beni, Guayaramerin, Garten, 18. XI. 1966, leg. Zicsi.
- AF/2867 1 Ex., Guayaramerin, Park, 2. XII. 1966, leg. Zicsi.
Eukerria garmani (Rosa, 1895)
Fundorte. AF/2854 16 Ex., Prov. Beni, Guayaramerin, Garten, 18. XI. 1966, leg. Zicsi.
- AF/2869 55 Ex., Guayaramerin, Park, 2. XII. 1966, leg. Zicsi.
604 ANDRAS ZICSI
Eukerria asuncionis (Rosa, 1895)
Fundort. AF/2863 30 Ex., Prov. Beni, Guayaramerin, Plantage am Ufer des Mamore
Flusses, 24. XI. 1966, leg. Zicsi.
Eukerria saltensis (Beddard, 1896)
Fundorte. Prov. La Paz. Zwischen Teoponte und Alcoche, 550 m., 19. XII. 1966, leg.
Zicsi. - AF/2881 1 Ex., G/INVE 17977 1 Ex., Prov. Tarija, km 15 Entre Rios nach Tarija, 4. XI.
1993, leg. Vaucher.
Ocnerodrilus Eisen, 1878
Ocnerodrilus occidentalis Eisen, 1878
Die Revisionen der Gattung Ocnerodrilus VON GATES (1973, 1979) und RIGHI
(1994) beriicksichtigend, ordne ich meine Exemplare O. occidentalis Eisen zu.
Fondorte: AF/2836 1 Ex., AF/2856 6 Ex., AF/2866 1 Ex., Prov. Beni,
Guayaramerin und Umgebung, 18. XI- 2. XII. 1966, leg. Zicsi.
Ilyogenia Beddard, 1892
Ilyogenia tuberculatus Eisen, 1900
Fundorte: AF/2857 13 Ex., Prov. Beni, Guayaramerin Nueva Cuba, 26. XI. 1966, leg.
Zicsi. - AF/2868 1 Ex., Guayaramerin, Park, 2. XII. 1966, leg. Zicsi.
Belladrilus Righi, 1984
Belladrilus (Belladrilus) vaucheri sp. n.
Belladrilus wurde von RIGHI (1994a) aufgrund der Chylustaschen-Struktur in 2
Untergattungen geteilt. Belladrilus s. s. besitzt keine Längsfurchen, während bei
Belladrilus (Santomeria) im Querschnitt der Chylustaschen kein einziger Hohlraum
sondern nur Längsfurchen zu erkennen sind. Meine zur Beschreibung vorliegenden
Exemplare besitzen nur einen Hohlraum im Querschnitt der Chylustaschen und
werden so zur Untergattung Belladrilus gestellt.
Lange des Holotypus 178 mm, Dicke 2 mm, Segmentzahl 157, Paratypen:
Lange 55-168 mm, Dicke 1,5-1,8 mm, Segmentzahl 130-152. Farbe weiss, unpig-
mentiert. Kopf prolobisch-epilobisch, 1/3 zu. Borsten entlang des Körpers eng ge-
paart. Borsten ab am Vorderkörper etwas kleiner als cd; aa gleich bc. Borstenver-
hältnis hinter dem Giirtel aa : ab : bc: cd: dd wie 5: 1 : 5,3,: 1 : 18,3.
Weibliche Poren auf dem 14. Segment vor der Borstenlinie b. Giirtel sattel-
förmig vom 13.-22. Segment, nur leicht angeschwollen; es ist anzunehmen, dass die
Tiere einen zurückgebildeten Giirtel besitzen. Pubertätsfeld zwei grosse längliche
Tuberkeln auf dem 17. und 18. Segment bildend, wo auch die Borsten ab fehlen.
Prostataporen auf dem 17. Segment in einer kleinen Vertiefung, durch eine Samen-
REGENWURMER AUS BOLIVIEN 605
rinne mit den männlichen Poren des 18. Segmentes verbunden (Abb. 18). Ein Paar
Samentaschenporen auf Intersegmentalfurche 8/9 in der Borstenlinie cd, es sind
kleine Schlitze zwischen den Borsten c und d.
ABB. 18-19
Belladrilus (Belladrilus) vaucheri sp. n. 18. Ventralansicht mit dem Geschlechtsfeld. PP =
Prostataporen, SR = Samenrinne. 19. Samentasche des 9. Segmentes.
Innere Organisation. Dissepimente 6/7-8/9 stark verdickt, 9/10-11/12 nur an-
gedeutet, nicht verdickt. Schlunddriisen reichen bis ins 7. Segment und tiberdecken
den im 7. Segment liegenden, muskulôsen Muskelmagen. Der Muskelmagen besitzt
auch ein kragenförmiges Gebilde im 6. Segment. Kalkdriisen im 9. Segment, sie sind
nach vorne gerichtete, langliche Gebilde die am vorderen Teil mit einem seitlich
verlaufenden Blutgefäss in Verbindung stehen. Innere Struktur mit einem Hohlraum
ohne Längsfurchen. Hoden und Samentrichter im 10. Segment, Samensäcke im 9. und
11. Segment, klein. Mächtige Intestinalherzen im 10. und 11. Segment, Rückengefäss
im 12. und 13. Segment perlschnurartig verdickt. Ovarien im 13. Segment, grosse mit
Eiern gefiillte herzformige Gebilde, Ovarientrichter in 13/14 Segment, Eileiter im
14.Segment ausmiindend. Samenleiter durchbrechen im 11./12. Segment das Disse-
piment und verlaufen der Innenwand angeschmiegt bis unter die Einmiindung der
Prostata ins 18. Segment, wo sie ohne Bildung einer Kopulationstasche ausmiinden.
Prostata im 17. Segment mit 4-5 Segmente einnehmendem muskulösen Ausführungs-
gang und einem langen gewundenen Driisenteil, der bis ins 40. Segment reichen kann.
Mitteldarm im 13. Segment beginnend, ohne Typhlosolis. Nephridien ohne
Nephridialblase.
Samentaschen im 9. Segment, mächtige Ampulle mit kurzem Ausführungs-
gang (Abb. 19)
Die neue Art steht Belladrilus (B.) jimi am nächsten. Unterscheidet sich jedoch
von diesem durch die Samensäcke im 9. Segment, durch die Form der Samentaschen,
die viel langeren Prostatadrüsen und durch das Vorhandensein von Ovarientrichtern.
Die neue Art wird zu Ehren des Sammlers, Dr. Cl. Vaucher (Naturhistorisches
Museum Genf) benannt.
Fundort: Holotypus G/INVE 17978. Prov. Tarija, km 15 Entre Rios nach Tarija, 4. XI.
1993, leg. Vaucher. Paratypen: AF/2898 4 Ex., G/INVE 17079 2 Ex., Fundort wie Holotypus.
606 ANDRAS ZICSI
Lumbricidae Rafinesque-Schmaltz, 1815
Allolobophora Eisen, 1874
Allolobophora caliginosa trapezoides (Ant. Dugès, 1828)
Fundorte: Z/11411 75 Ex., La Paz, 3600 m, 25. XII. 1966, leg. Zicsi. - Z/11414 1+1
juv. Ex., Z/11422 8 Ex., Prov. La Paz, Titicaca-See, 3800 m, 28. XII. 1966, leg. Zicsi. -
Z/11417 2 Ex., Prov. La Paz, Oberhalb Unduavi 4650 m, 19.XII. 1966, leg. Zicsi. - Z/12008 4
Ex., G/2510 12 Ex., Prov. Tarija, km 15 Entre Rios nach Tarija, 4. XI. 1993, leg. Vaucher. -
Z/12012 2 Ex., Copacabana La Paz, Prov. Munco Capae 7. IV. 1991. leg. Gemio.
Allolobophora rosea (Savigny, 1826)
Fundorte: Z/11412 2 Ex., La Paz, 3600 m, 25. XII. 1966, leg. Zicsi. - Z/11413 23+8
Juv. Ex., Z/11421 13 Ex., Prov. La Paz, Titicaca-See, 3800 m, 28. XII. 1966, leg. Zicsi. -
Z/11416 8 Ex., Prov. La Paz, Oberhalb Unduavi 4654 m, 29. XII. 1966, leg. Zicsi. - Z/12013 2
Ex., Prov. Munco Capae, Copacabana La Paz, 7. IV. 1991, leg. Gemio.
Dendrodrilus Omodeo, 1956
Dendrodrilus rubidus (Savigny 1826)
Fundorte: Z/11415 2 Ex., Prov. La Paz, Titicaca-See, 3800 m, 28. XII. 1966, leg. Zicsi.
- Z/11419 2 Ex., Prov. La Paz, Oberhalb Unduavi 4654 m, 29. XII. 1966, leg. Zicsi. - Z/11412
2 Ex., G/1511 7 Ex., Prov. Tarija, km 15 Entre Rios nach Tarija, 4. XI. 1993, leg. Vaucher.
Dendrobaena Eisen, 1874
Dendrobaena octaedra (Savigny, 1826)
Fundorte: Z/11420 18+8 juv. Ex., Prov. La Paz, Oberhalb Unduavi 4654 m, 29. XI.
1966, leg. Zicsi.
Octolasium Orley, 1885
Octolasium lacteum (Orley, 1881)
Fundort: Z/11418 9 Ex., Prov. La Paz, Oberhalb Unduavi, 4654 m, 29. XII. 1966, leg.
Zicsi.
Megascolecidae Rosa, 1891
Amynthas Kinberg, 1867 emend. Easton, 1982
Amynthas corticis (Kinberg, 1867)
Fundort: AF/2894 7 Ex., Dept. La Paz, Prov. Murillo, Valle de Zongo, Est Cahura, 150
m, 26. XI. 1991, leg. Hanagarth.
REGENWURMER AUS BOLIVIEN 607
LITERATUR
BALOGH, J. & S. MAHUNKA & A. ZIcsi, 1969. The Scientific Results of the Hungarian Soil
Zoological Expeditions to South America 14. A Report on the Collectings of the
Second Expedition. Folia Ent. Hung. (Ser. nov.) 22: 453-474.
BEDDARD, F.E. 1892. On some new species of Earthworms from various parts of the world.
Proc. zool. Soc. Lond. 666-706:45-46.
CERNOSVITOV, L. 1939. The Percy Sladen Trust Expedition to Lake Titicaca in 1937. VI.
Oligochaeta. Trans. Linn. Soc. London 3:39-52.
COGNETTI DE MARTIIS, L. 1902 a. Terricoli boliviani ed argentini. Boll. Mus. Torino 17:1-11.
COGNETTI DE MARTIS, L. 1902 b. Un nuovo genere della Fam. Glossoscolecidae. Atti Acad.
Torino 37: 432-446.
COGNETTI DE MARTIS, L. 1905. Gli Oligocheti della Regione Neotropicale. Mem. R. Acad. Soc.
Torino 56:1-72.
COGNETTI DE Martius, L. 1908. Lombrichi di Costa Rica e del Venezuela. Atti Acad. Torino 43:
913-926.
Csuzpi, Cs. & A. Zıcsı 1991. Uber die Verbreitung neuer und bekannter Dichogaster und Eutri-
gaster Arten aus Mittel- und Südamerika (Oligochaeta; Octochaetidae) Regenwürmer
aus Südamerika 15. Acta Zool. Hung. 37:177-192.
EASTON, E.G. 1982. Australian Pheretimoid Earthworms (Megascolecidae, Oligochaeta): A
synopsis with the description of a new genus and five new species. Aust. J. Zool. 30:
711-735.
GATES, G.E. 1973. Contributions to North American earthworms (Annelida) 7. Contribution to
a revision of the earthworm family Ocnerodrilidae IX. What is Ocnerodrilus
occidentalis? Bull. Tall. Timbers Res. Stat. 14: 13-28.
GATES, G.E. 1979. A new genus of larger Ocnerodrilid earthworms in the American hemi-
sphere. Megadrilogica 3: 162-164.
MICHAELSEN, W. 1900. Die Terricolen-Fauna Columbiens. Arch. Naturg. 66/1/: 231-266.
MICHAELSEN, W. 1902. Neue Oligochaeten und neue Fundorte altbekannter. Mit. Mus. Ham-
burg 19: 1-54.
MICHAELSEN, W. 1918. Die Lumbriciden, mit besonderer Beriicksichtigung der bisher als
Familie Glossoscolecidae zusammengefassten Unterfamilien. Zool. Jb. Syst. 41: 1-398.
MICHAELSEN, W. 1926. Zur Kenntnis einheimischer und ausländischer Oligochaeten. Zool. Jb.
Syst. 51:255-328.
MICHAELSEN, W. 1935. Oligochaeten aus Peru. Capita Zool. 6/2/: 1-12.
MICHAELSEN, W. 1936. On the genus Thamnodrilus Beddard. Proc. Zool. Soc. Lond. 1171-
14172.
PICKFORD, G.E. 1937. A Monograph of the Acanthodriline Earthworms of South Africa. Heffer
and Sons, Cambridge. 612 pp.
RIGHI, G. 1968. Über die Oligochaetengattung Eukerria. Beitr. zur. Neotrop. Fauna. 5:178-185.
RIGHI, G. 1984 a. On a Collection of Neotropical Megadrili Oligochaeta I. Ocnerodrilidae,
Acanthodrilidae, Octochaetidae, Megascolecidae. Stud. Neotrop. Fauna. 19(1):9-31.
RIGHI, G. 1984 b. On a Collection of Neotropical Megadrili Oligochaeta II. Glossoscolecidae,
Lumbricidae. Stud. Neotrop. Fauna. 19(2): 73-78.
RIGHI, G. 1986. Sobre o genero Andiorrhinus (Oligochaeta, Glossoscolecidae). Bolm Zool.
Univ. S. Paulo. 10: 123-151.
RIGHI, G. 1989. Adicao ao conhecimento dos Oligochaeta da Venezuela. Rev. Brasil. Biol.
49(4):1065-1084.
RIGHI, G. 1993. Venezuelan earthworms and consideration on the genus Andiorrhinus Cognetti
1908 (Oligochaeta, Glossoscolecidae). Trop. Zool. 1: 125-139.
608 ANDRAS ZICSI
RIGHI, G. 1994. On a new and old-known Oligochaeta genera from Paraiba State, Brasil. Revue
suisse Zool. 101: 89-106.
RIGHI, G. & J. ROMBKE 1987. Alguns Oligochaeta da Bolivia e do Peru. Rev. Brasil. 47(4):523-
I:
Rosa, D. 1895. Terricoli Neotropicali. Mem. Acad. Torino 45(2):89-152.
Zıcsı, A. 1989. Revision der Gattung Yagansia Michaelsen, 1899 (Oligochaeta, Acanthodri-
lidae) Regenwürmer aus Südamerika 11. Acta zool. Hung. 35:413-430.
Zıcsı, A. 1992. Über weitere neue und bekannte Arten der Gattung Periscolex (Oligochaeta:
Glossoscolecidae) Regenwürmer aus Südamerika 16. Revue suisse Zool. 99(1): 211-
DITE
REVUE SUISSE DE ZOOLOGIE, 102 (3) : 609-613; septembre 1995
Studio dei tipi di Tychiini di Henri Tournier custoditi presso
il Museo di Storia Naturale di Ginevra (Coleoptera, Curculionidae)
Roberto CALDARA
Piazza Bolivar 7, 20146 Milano, Italia.
Study of the Tychiini types of Henri Tournier preserved at the Natural
History Museum of Geneva (Coleoptera, Curculionidae). — The lecto-
types of Tychius depressicollis Tournier and Sibinia rudepilosa Tournier are
designated. Moreover, one holotype and paralectotypes of several Tychius
and Sibinia species described by Tournier and housed at the MHNG are
listed. The following new synonymies are proposed: Tychius brisouti
Tournier, 1873 (= T. galloprovincialis Hustache, 1924, n. syn.); T. fuscipes
Chevrolat, 1859 (= 7. depressicollis Tournier, 1873, n. syn.); 7. seriepilosus
Tournier, 1873 (= 7. terrosus Tournier, 1873, n. syn.); Sibinia tibialis
(Gyllenhal, 1836) (= S. rudepilosa Tournier, 1873, n. syn.).
INTRODUZIONE
Sono più di 60 le descrizioni di Tychiini effettuate da Tournier e riportate per la
maggior parte in un'unica pubblicazione (TOURNIER, 1873). Quando ho effettuato la
revisione dei Tychiini paleartici, una delle parti più impegnative è stata proprio quella
di rintracciare il numero più elevato possibile di tipi delle specie di tale autore. La
collezione di curculionidi di Tournier è attualmente custodita per la massima parte al
Museo di Storia Naturale di Parigi (MHNP), ove è tuttora conservata nelle scatole
originali ed arrangiata secondo la sistematica seguita dall'autore e con le etichette per
ogni specie. E' da notare però che prima di essere acquistata dal MHNP, essa era stata
venduta a Maurice Pic nel 1894. Pic, oltre a mettere sugli esemplari delle serie tipiche
dei cartellini rossi stampati o bianchi scritti a mano con l'indicazione di type (per quanto
riguarda i Tychiini in modo solitamente molto accurato), ha spostato numerosi
esemplari che ora si trovano sparsi confusamente in altre scatole. Nonostante questi
rimaneggiamenti, dopo approfondite ricerche sono riuscito ad esaminare quasi tutti i
tipi delle specie di Tournier. Tuttavia alcune di esse, mai prese in esame da nessun altro
autore dopo la loro descrizione, mi sono rimaste sconosciute.
Per cercare di ovviare a ciò, visto che il Museo di Storia Naturale di Ginevra
(MHNG) ha acquistato la parte rimanente della collezione Tournier alla sua morte ed
anche la collezione Ernest Poncy, al quale Tournier aveva donato numerosi suoi
Manoscritto accettato il 09.11.1994.
610 ROBERTO CALDARA
esemplari, ho chiesto e ottenuto, grazie alla gentilezza del Dr. Claude Besuchet, di
esaminare gli esemplari di Tychius e Sibinia presenti in tali collezioni. L'esame è stato
molto fruttuoso, perchè mi ha permesso di stabilire lo stato tassonomico di tre specie di
Tychius e una di Sibinia descritte da Tournier a me non note in precedenza e di rin-
tracciare molti esemplari appartenenti alle serie tipiche di numerose specie dell'autore.
EISTA DELLE SPECIE
Tychius armatus Tournier
Tychius armatus TOURNIER, 1873: 495; CALDARA 1990: 190 (lectotypus des.). (Syn. di T. grandi-
collis Desbrochers).
Paralectotypi MHNG: 1 d e 1 2 "Algérie, Raffray" el d el 2 "Tanger".
Tychius brisouti Tournier
Tychius brisouti TOURNIER, 1873: 471; CALDARA 1990: 203.
Specie descritta su esemplari della Svizzera (Giura bernese: Saint-Imier) e
rimasta a me sconosciuta. La descrizione di Tournier è piuttosto precisa e ricca di
particolari utili alla determinazione del taxon. Oltre a caratteristiche diagnostiche
minori, i dati più importanti sono costituiti dal rivestimento elitrale, formato sia da
squame strette grigio-dorate che da squame larghe biancastre, e dai femori posteriori
inermi. Solo 4 specie in tale area geografica hanno un simile tipo di rivestimento: il 7.
galloprovincialis Hustache, il T. tridentinus Penecke, il T. alpinus Hustache e il T.
parallelus (Panzer), ma solo la prima specie ha i femori posteriori senza dentino. Al
MHNG è presente 1 esemplare 2, perfettamente conservato e classificato come 7. bri-
souti, che porta i seguenti cartellini "Thoiry, Jura. — Juin. (scritto a mano in inchiostro
nero) / Collection E. Poncy. (scritto a stampa in nero)" e che corrisponde esattamente
alla descrizione originale. Come ipotizzato, tale esemplare appartiene proprio alla
specie chiamata attualmente 7. galloprovincialis e nota di poche località della Francia
meridionale e della Spagna sud-orientale (CALDARA 1990). A questo punto non ho più
dubbi che 7. brisouti Tournier, 1873 sia la stessa cosa di 7. galloprovincialis Hustache,
1924 che pertanto diventa suo sinonimo. Purtroppo, come risulta dal cartellino, l'esem-
plare sembra apparentemente raccolto in una località del Giura differente da quella
riportata da Tournier e non mi è pertanto possibile designarlo come lectotypus, sebbene
non si possa escludere un errore di citazione da parte dell'autore francese come
avvenuto in altre circostanze (Besuchet, in litt.). La località Thoiry in questione
(esistono altre due cittadine francesi con tale nome, una nella Savoia e l'altra nella Seine
et Oise) si trova in territorio francese nel dipartimento dell'Ain, a pochi chilometri dal
confine con la Svizzera. Essa risulta al momento la località più settentrionale in cui è
stata trovata la specie ma, se fosse esatta l'indicazione Saint-Imier riportata da Tournier,
la diffusione di 7. brisouti a nord sarebbe ulteriormente estesa.
TIPI DI HENRI TOURNIER 611
Tychius comptus Tournier
Tychius comptus TOURNIER, 1873: 497; CALDARA 1990: 190 (lectotypus des.). (Syn. di T. tibialis
Boheman).
Paralectotypi MHNG: | © "Sicile/ T. comptus" e 2 4 G "Tanger".
Tychius depressicollis Tournier
Tychius depressicollis TOURNIER, 1873: 468; CALDARA 1990: 204.
Specie descritta su esemplari dell' Algeria senza più precise indicazioni e para-
gonata da Tournier a 7. fuscipes Chevrolat dal quale differirebbe per la forma più
appiattita, per il protorace a lati più arrotondati e per il rostro più lungo. Nella colle-
zione Tournier al MHNP non ho trovato esemplari classificati con tale nome e la specie
mi è rimasta sconosciuta. Al MHNG è custodita 1 © classificata come depressicollis ed
etichettata "Algérie", perfettamente corrispondente alla descrizione originale (lecto-
typus qui designato). Essa non mostra nessuna differenza sostanziale dai tipici esem-
plari di 7. fuscipes e pertanto stabilisco che T. depressicollis Tournier, 1873 è sinonimo
di T. fuscipes Chevrolat, 1859.
Tychius hypaetrus Tournier
Tychius hypaetrus TOURNIER, 1873: 469; CALDARA 1990: 147 (lectotypus des.). (Syn. di 7.
depressus Desbrochers).
Paralectotypi MHNG: | 9 "Bone".
Tychius kiesenwetteri Tournier
Tychius kiesenwetteri TOURNIER, 1873: 485; CALDARA 1985 b: 88 (lectotypus des.); 1990: 138;
(Syn. di 7. trivialis Boheman).
Paralectotypi MHNG: 1 d e 1 2 "Serbie".
Tychius ochraceus Tournier
Tychius ochraceus TOURNIER, 1873: 505; CALDARA 1990: 199 (lectotypus des.).
Paralectotypi MHNG: 1 © "Syrie".
Tychius olcesei Tournier
Tychius olcesei TOURNIER, 1873: 483; CALDARA 1990: 90 (lectotypus des.).
Paralectotypi MHNG: 2 dd e 3 9 "Tanger, Olcèse" e 2 2 2 "Alger, Raffray".
Tychius sericatus Tournier
Tychius sericatus TOURNIER, 1873: 488; CALDARA 1974: 178 (lectotypus des.); 1990: 177. (Syn.
di T. crassirostris Kirsch).
Paralectotypi MHNG: 2 9 9 "Genève".
612 ROBERTO CALDARA
Tychius seriepilosus Tournier
Tychius seriepilosus TOURNIER, 1873: 460; CALDARA 1990: 151 (lectotypus des.).
Paralectotypi MHNG: 2 9 © "Egypte".
Tychius sharpi Tournier
Tychius sharpi TOURNIER, 1873: 506; CALDARA 1990: 197 (lectotypus des.).
Paralectotypi MHNG: 1 9 "Peney, Genève".
Tychius similis Tournier
Tychius similis TOURNIER, 1873: 401; CALDARA 1978: 29 (lectotypus des.). (Syn. di T. multi-
lineatus Desbrochers).
Paralectotypi MHNG: 1 © "Blidah, Algérie" e 1 d "Algérie".
Tychius subsulcatus Tournier
Tychius subsulcatus TOURNIER, 1873: 470; CALDARA 1990: 102 (lectotypus des.).
Paralectotypi MHNG: 1 d e 2 2 2 "Hongrie".
Tychius terrosus Tournier
Tychius terrosus TOURNIER, 1873: 475; CALDARA 1990: 205.
Specie descritta su 1 d della Calabria, da me non reperito al MHNP. Dopo la
descrizione nessun autore si è più occupato della specie, che pertanto è stata da me
posta fra quelle incertae sedis (CALDARA 1990). Al MHNG, sotto questo nome, è
conservato | d etichettato "Italie" perfettamente corrispondente alla descrizione ori-
ginale. Il fatto che sul cartellino non sia riportata anche la dizione "Calabria" non è
inusuale per gli esemplari di Tournier e pertanto ritengo tale esemplare l'holotypus della
specie. Esso non mostra nessuna differenza dai tipici esemplari di 7. seriepilosus
Tournier (anche le due descrizioni, effettuate nella stessa pubblicazione a poche pagine
di distanza, sono molto simili sebbene quella del 7. seriepilosus sia molto più corta) e
pertanto stabilisco che 7. terrosus TOURNIER, 1873 (p. 475) è sinonimo di 7. serie-
pilosus TOURNIER, 1873 (p. 460).
Questa risulta la prima segnalazione della specie per l'Italia, essendo 7. serie-
pilosus noto in precedenza solo del Nord Africa e del Vicino Oriente dove è largamente
distribuito (CALDARA 1990). Pur rimanendo il dubbio che la località di raccolta riportata
sul cartellino sia errata, la presenza di questa specie nell'Italia meridionale, vista la sua
distribuzione, è verosimile.
Sibinia fusca Tournier
Sibinia fusca TOURNIER, 1873: 513; CALDARA 1979: 84 (lectotypus des.).
Paralectotypi MHNG: 2 6 d "Egypte".
TIPI DI HENRI TOURNIER 613
Sibinia hopffgarteni Tournier
Sibinia hopffgarteni TOURNIER, 1873: 514; CALDARA 1985a: 80 (lectotypus des.).
Paralectotypi MHNG: 1 d e 1 9 "Hongrie".
Sibinia reichei Tournier
Sibinia reichei TOURNIER, 1873: 514; CALDARA 1979: 83 (lectotypus des.).
Paralectotypi MHNG: 1 d e 1 © "Calabre, Huet”.
Segnalo che, sempre al MHNG, 1 ® di S. reichei risulta raccolta in Sicilia senza
più precise indicazioni. E' da notare che della specie non avero mai esaminato altri
esemplari raccolti in Italia oltre ai tipi.
Sibinia rudepilosa Tournier
Sibinia rudepilosa TOURNIER, 1873: 520; CALDARA 1985a: 93.
Specie descritta su esemplari della Turchia non presenti al MHNP e rimasta
sconosciuta a tutti gli autori, me compreso. Tournier la paragona alla S. pellucens
(Scopoli), dalla quale differirebbe essenzialmente per le tibie e i tarsi di colore ros-
sastro. Sotto tale nome al MHNG ho trovato 1 d etichettato "Turquie", ben corris-
pondente alla seppur sintetica descrizione originale e che designo come lectotypus della
specie. Non esiste nessuna differenza fra questo e i tipici esemplari di S. tibialis
(Gyllenhal) e pertanto stabilisco che S. rudepilosa Tournier, 1873 è sinonimo di S.
tibialis (Gyllenhal, 1836). E' questa la prima segnalazione di S. tibialis per la Turchia.
La specie comunque risultava largamente diffusa, seppure in modo sporadico, in
Europa centrale ed orientale, con la località più a sud rappresentata da Deliblato
(Serbia).
BIBLIOGRAFIA
CALDARA, R. 1974. Note sinonimiche su alcune specie del genere Tychius Germar. Boll. Soc. ent.
ital. 106: 178-181.
CALDARA, R. 1978. I generi Apeltarius Desbrochers, Xenotychius Reitter e Pseudolignyodes Pic.
Boll. Soc. ent. ital. 110: 23-34.
CALDARA, R. 1979. Revisione delle specie paleartiche di Sibinia vicine a sodalis Germar ed
exigua Faust. Mem. Soc. ent. ital. 57: 65-100.
CALDARA, R. 1985a. Revisione delle Sibinia paleartiche. Mem. Soc. ent. ital. 62/63: 24-105.
CALDARA, R. 19855. Taxonomic notes on three little known species of Tychius Germ. from
Siberia (USSR). Reichenbachia 23: 87-89.
CALDARA, R. 1990. Revisione tassonomica delle specie paleartiche del genere Tychius Germar.
Mem. Soc. ital. Sci. nat. Mus. civ. Stor. nat. Milano 25: 51-218.
TOURNIER, H. 1873. Observations sur les espèces européennes et circumeuropeennes de la tribu
des Tychiides. Annls Soc. ent. Fr. 3(5): 461-509.
REVUE SUISSE DE ZOOLOGIE, 102 (3) : 615-627; septembre 1995
Contributi allo studio delle Crambinae (Lepidoptera: Crambidae).
IX. Note sui generi Sebrus Bteszyfiski e Alphacrambus nov.,
con descrizione di nuove specie
Graziano BASSI
c/o Museo Regionale di Scienze naturali, Via Giolitti 36, I-10123 Torino, Italia.
Contribution to the study of the Crambinae (Lepidoptera, Crambidae).
IX. Notes on genera Sebrus Bteszyfiski and Alphacrambus nov., with
description of new species. - The genus Sebrus Bteszyñski is revised; its
differences towards related genera are discussed, its tympanal organs are
shown; S. perdentellus (Hmps) n. comb. and S. pseudosparsellus (Btesz.) n.
comb. are transferred from Crambus F.; a lectotypus is designated for S.
perdentellus (Hmps.); S. absconditus n. sp. and S. argus n. sp. are described
and illustrated. Alphacrambus gen. nov. is described based on differences in
genitalia to Crambus F. and Crambixon Btesz.. Its tympanal organs are
described. A. prodontellus (Hmps) n. comb., type species, and A. razowskii
(Btesz.) n. comb., are transferred from Crambus F. and their females are
correctly assigned; A. phoeostrigellus (Hmps.) n. comb. is transferred from
Crambixon Bteszyfiski; A. cristatus n. sp. and A. parvus n. sp. are described
and illustrated.
Key-words: Pyraloidea - Crambidae - Crambinae - new genus - new species.
INTRODUZIONE
Lo studio del cospicuo materiale africano appartenente alle Crambinae conser-
vato nei Musei di Budapest, Copenaghen, Ginevra, Londra, Parigi, Pretoria e Tervuren,
mi consente di esporre alcune considerazioni su specie finora assegnate al genere
Crambus F. Quest'ultimo infatti ha da sempre accolto la maggior parte delle specie
ascrivibili alle Crambinae, ed oggi, almeno per quel che riguarda la Fauna Afrotro-
picale, si rende necessaria un'accurata revisione tesa a garantire l'unità filetica dei taxa
specifici all'interno dei gruppi di rango superiore. Questo lavoro segue ad altri (BASSI
1991, 1992 e 1995) e cerca di contribuire con uno studio che considera prioritaria la
morfologia degli apparati genitali, ad una organica revisione della sistematica dei
Crambini (sensu GASKIN 1975) della regione Etiopica.
Manoscritto accettato il 02.12.1994.
616 GRAZIANO BASSI
Abbreviazioni usate:
CB = Collezione Bassi, Torino.
GS...GB = Preparato microscopico .... G. Bassi.
GS...SB = Preparato microscopico .... S. Bteszynski.
MHNG = Muséum d'histoire naturelle, Genève.
MNHNP = Muséum National d'Histoire Naturelle, Paris.
NHML = Natural History Museum, London.
TM = Musée Royal de l'Afrique Centrale, Tervuren.
TMB = Magyar Termèszettudomànyi Mùzeum, Budapest.
TMP = Transvaal Museum, Pretoria.
ZMC = Zoologisk Museum Kobenhavn
Sebrus Bteszynski, 1970.
Specie-tipo: Sebrus amandus Bteszynski, 1970: 12, per designazione originale.
Il genere venne descritto per una specie, S. amandus Btesz., del Madagascar.
Esso si caratterizza, nell'ambito dei Crambini, oltre per quanto evidenziato da
Bteszynski (1970), per la peculiarità dell'anatomia del vinculum e del tipo di giunzione
tra tegumen e vinculum nel genitale 4. L'ostium bursae differenziato ma senza processi
sporgenti dall' VIII segmento addominale, la presenza in quest'ultimo di una placca
urosternale nell'area dell'ostium bursae, il ductus seminalis che si diparte sempre a
livello dell'antrum e l'assenza di signa in una bursa copulatrix debolmente corrugata
differenziano invece il genitale © da quello dei generi più affini, che sembrano essere
Pediasia Hb., Caffocrambus Btesz., Catoptria Hb. e Crambus F.
In tutte le specie conosciute vi è dimorfismo sessuale: Le ali posteriori del 4
sono sempre molto più chiare di quelle della 9.
Organi timpanali (Fig.24): Bulla tympani di medie dimensioni. Pons tympani
ben sviluppato, come pure il processus tympani. Area subtimpanale inferiore, compresa
tra pons tympani ed ala tympani, ampia e ben definita. Angolo tra sclerite tergo-sternale
e base del pons tympani acuto. Placca sternale dell'addome subrettangolare e
notevolmente sviluppata. Venulae secundae ridotte e ben visibili solo al centro della
placca sternale. Le 9 © hanno la bulla tympani di dimensioni ridotte.
Geonemia: sud-est della regione Afrotropicale.
La revisione critica del materiale finora ascritto al genere Crambus F. mi per-
mette di assegnare le seguenti specie al genere qui considerato.
TABELLA DELLE SPECIE
l Specie.bianco-bruner 5.2 ee esiti argus
— Specie bianco-grigier. can. 2eme ee E 2
2 macchia grigio-nera al centro dell'ala anteriore che raggiunge il margine
AL ANE? dics ee ne Bee pseudosparsellus
NOTE SUI GENERI SEBRUS E ALPHACRAMBUS 617
3 genitale d con phallus senza denti apicali, genitale 2 con ostium bursae
SclenimcatovedeantrumemnembranOSOmse se. eerie oe eee ore amandus
— genitale d con phallus dotato di un solo grande dente apicale inferiore e
minuti dentelli perimetrali; genitale 9 con ostium bursae e antrum
Sclenacatindit medic Ginensionite as ae eas. cee ees ne perdentellus
— genitale d con phallus dotato di molti aguzzi denti apicali, genitale 9
con ostium bursae e antrum sclerificati, di piccole dimensioni. . . . . absconditus
Sebrus amandus Bleszynski, 1970: 7.
Holotypus d: [Madagascar] Station Perinet, 149km east of Tananarive, 20.X-
10.X1.1930, (Mme N. d'Olsoufieff), GS 5586SB (11282BM Pyral.), nel NHML, esaminato. Altro
materiale esaminato: 1 ©, paratypus, Madagasc.[ar], GS 5822 SB (16176 BM Pyral.), nel
NHML; 5 ? ©, Madagascar, GS 2388, 2432, 2631 e 2628 GB (Fig. 21), nel MNHNP e in CB.
Sebrus perdentellus (Hampson, 1919). n. comb.
Crambus perdentellus Hampson, 1919: 287.
Lectotypus d (qui designato): [Malawi] Nyasaland, Mlanje Plateau, 6500ft., 18.XII.
1913, S.A. Neave, 1914-171, "Crambus perdentellus, Type 6, Hampson", GS3202GB (19481
BM Pyral.), nel NHML.
Altro materiale esaminato: Paralectotypi: 58 ¢, 22 2, stessa etichetta del lectotypus,
GS3150 (19482 BM Pyral), 3228 e 3229GB, nel NHML; 1 d, Nyasaland ,Mlanje Plateau, 6500
ft., S.A. Neave, 14.XI. 1913, PG 1660 GB, NHML.
Cenni morfologici (Fig. B): ben descritto da Hampson, si può aggiungere che
l'apertura alare varia da 15,5 a 22 mm; i palpi labiali sono lunghi 3 volte il diametro
dell'occhio; gli ocelli e le chetosemae sono presenti; la fronte, tondeggiante, sporge
appena oltre l'occhio; l'addome è bruno bronzeo con ciuffo anale bianco. Si ha un netto
dimorfismo sessuale nella colorazione delle ali posteriori: nel 4 sono bronzee fino alla
venatura M2 esclusa, poi bianche e bronzee solo lungo il margine alare fino Cu2, indi
interamente bianche; nella 9 sono invece interamente brune.
Apparato genitale d (Fig.3): Uncus e gnathos circa della stessa lunghezza.
Tegumen ben sviluppato, nettamente separato dal vinculum. Vinculum con parte
supero-distale che si sovrappone al tegumen e tozza protuberanza infero-distale. Juxta
subconica, con due sclerificazioni indipendenti ed opposte. Pseudosaccus mediamente
sviluppato. Valve simmetriche, con processo costale ben sviluppato, sia verso il
cucullus che, con un processo secondario, verso l'interno della valva. Phallus (Fig. 4) di
medie dimensioni, con un solo cornutus lungo e molto sottile. Il suo apice presenta un
grande dente ventrale e minuti dentelli lungo il perimetro (Fig. 7). La sclerificazione
dell'urosterno dell'VIII segmento addominale (Fig.6) è tozza e subconica.
Apparato genitale 2 (Fig.19): Papillae anales di piccole dimensioni. Apophyses
posteriores di media lunghezza, sottili. L'VIII segmento addominale, molto sottile,
forma una placca urosternale dentellata al di sopra dell'ostium bursae. Apophyses
anteriores subvestigiali. Ostium bursae tondeggiante. Antrum ampio e sclerificato, non
618 GRAZIANO BASSI
FicG. 1-9
Sebrus spp., apparati genitali d d. Scala: 0,5 mm. 1: S. absconditus n. sp., paratypus, GS3064GB,
phallus e valva dx estratti. 2: idem, phallus. 3: S. perdentellus (Hmps.), lectotypus, phallus e valva
dx estratti. 4: idem, phallus. 5: $. absconditus n. sp., paratypus, GS3064GB, placca urosternale
dell'VIII segmento addominale. 6: S. perdentellus (Hmps.), lectotypus, placca urosternale dell'VIII
segmento addominale. 7 - 9: variabilità nell' apice del phallus: 7- S. perdentellus (Hmps.),
lectoparatypus, GS3229GB; 8 - S. absconditus n. sp., paratypus, GS3226GB; 9 - idem, holotypus.
NOTE SUI GENERI SEBRUS E ALPHACRAMBUS 619
differenziato dall'ostium. Il ductus bursae è lunghissimo, membranoso, interamente
corrugato; a 2/3 della sua lunghezza è presente un caratteristico ispessimento debol-
mente sclerificato. Da qui il ductus si allarga via via fino a giungere alla bursa copu-
latrix, che si presenta piccola, lievemente corrugata e senza signa. Il ductus seminalis
prende origine tra l'antrum e il ductus bursae.
Sebrus absconditus n. sp.
Holotypus d , [S. Africa, E. Transvaal] Woodbush, 1670m., 11.1.[19]25, A.J.T. Janse,
GS3240GB, nel TMP.
Paratypi: 14 , Pretoria, 11.3.'15, A.J.T. Janse; 18 , Woodbush, 1670 m., 25.1.11, A.J.T.
Janse, GS 3064 GB; 19, Blouberg, T[rans]v[aa]l, Motlakeng, 5-6,000 ft., 6.15.1.1955,
Transv.Mus.Exp., GS 3336 GB; 14 , Mazoe, S. Rhod.[esia] [Zimbabwe], 6-17 jan. 1920, A.J.T.
Janse, GS3226GB; 12, idem, PG3225GB; 19%, Salisbury, Rhod.[esia] [Zimbabwe], 31.12.'17,
A.J.T. Janse, GS 3280 GB; 19, Chirindo Forest swimmert [Mt. Selinda, Zimbabwe], 12.'18,
Coll. Janse, GS 2554 Janse, TMP e CB.
Cenni morfologici (Fig. C): Molto simile a S. perdentellus (Hmps.). Se ne
differenzia, anche ad un esame superficiale, per la maggior nitidezza e definizione nel
disegno delle ali e nei colori delle stesse, decisamente più grigiastre. Apertura alare
15,5-18 mm. Zampe argentee con articoli tarsali parzialmente dorati e speroni tibiali
minuti. Permane il dimorfismo sessuale osservato in perdentellus (Hmps.).
Apparato genitale d (Figg.1,2): Rispetto a perdentellus (Hmps.): processo
infero-distale del vinculum meno sviluppato, processo costale interno e placca mediale
del sacculus più sviluppati, apice del phallus sempre provvisto di processi lunghi e
acuminati lungo tutto il suo perimetro (Figg. 8, 9). La sclerificazione urosternale
dell'VIII segmento addominale (Fig.5) è significativamente diversa.
Apparato genitale 9 (Fig. 20): La caratteristica che meglio differenzia la n. sp.
da perdentellus (Hmps.) è il complesso ostium bursae-antrum. L'ostium bursae è più
minuto, mentre l'antrum è evidentemente più piccolo e corto.
Derivatio nominis: da absconditus = nascosto, allusione alla facilità con cui la
specie può venire confusa, senza l'esame degli apparati genitali, con S. perdentellus
(Hmps.).
Sebrus argus n.sp.
Holotypus d, [Zaire] Ht. Katanga, Panda, 16.1.[19]30, J. Romieux, GS 2185 GB (1472
MHNG), nel MHNG.
Paratypi: 16, Musée du Congo, Elisabethville, 24.2.1935, Ch. Seydel, GS 2456 GB; 1 4,
idem, 28.2.1935, GS 2450 GB; 1d, idem, IV.1938, GS 6345 SB (15922B.M Pyral.), "Crambus
argus Bteszyñski 1969"; 19, idem, III.1935, GS 6377 SB "argus", nei TM, NHML e CB.
Cenni morfologici (Fig. A): Apertura alare 12-14 mm., palpi lunghi 3,5 volte il
diametro degli occhi, bianchi con esternamente una macchia bruna prossimale ed una
nera distale. Fronte bianca, tondeggiante, sporge appena oltre l'occhio. Antenne debol-
mente serrate, color crema. Ocelli e chetosemae presenti. Capo, collo, tegulae e torace
bianco argentei. Ali anteriori bianco argentee con gruppi di scaglie bruno giallastre, che
formano una delicata doppia banda submarginale. Vi sono inoltre un'ampia macchia
620 GRAZIANO BASSI
Figc. 10-18
Sebrus e Alphacrambus spp., apparati genitali d 4. Scala: 0,5 mm. 10: S. argus n. sp., holotypus,
phallus e valva dx estratti. 11: idem, cornutus ingrandito. 12: idem, phallus. 13: A. prodontellus
(Hmps.), Zaire, Kyala, GS2233GB, phallus e valva dx estratti. 14: idem, phallus. 15: S.
pseudosparsellus (Blesz.), Zaire, Elisabethville, GS2556GB, phallus e valva dx estratti. 16: idem,
phallus. 17: A. parvus n.sp., holotypus, phallus e valva dx estratti. 18: idem, phallus.
NOTE SUI GENERI SEBRUS E ALPHACRAMBUS 621
tondeggiante bruno nerastra al centro dell'ala, due tacche nere traslucide submarginali
tra le venature Cul e Cu2 e una sottile banda marginale nera. Frange argentee. Ali
posteriori traslucide, bianco sporco con frange bianche nel à , brune con frange bianche
a base bruna nella ©. Zampe anteriori dorate anteriormente, argentee posteriormente; le
mediali e le posteriori sono argentee con speroni tibiali minuti.
Apparato genitale & (Fig. 10): E' tipico del genere. Si differenzia dalle altre
specie per la forma del processo costale e del phallus (Fig.12), munito di un ricurvo
dente apicale e di un lungo e sottile cornutus (Fig. 11).
Apparato genitale 9 (Fig. 27): Differisce dalle altre specie nella conformazione
della placca urosternale dell'VIII segmento addominale e nella forma dell'ostium
bursae. Il ductus bursae è mediamente sclerificato. La bursa copulatrix è debolmente
corrugata. i
Derivatio nominis: specie dedicata al mitico figlio di Friso. Mantengo il nome a
suo tempo indicato in litteris sui cartellini di determinazione da Bteszyñski, che già
aveva identificato il d ora nel NHML e la © ora nel TM come appartenenti ad una
specie nuova.
Sebrus pseudosparsellus (Bteszyfiski, 1961) n. comb.
Crambus pseudosparsellus Bteszyfiski, 1961: 189.
Holotypus 2: Salisbury, Mashonaland [Zimbabwe], 27.XII.[18]97, G.A.K. Marshall, 98-
62, GS 1529 SB (5277 B.M.Pyral.), nel NHML, esaminato.
Altro materiale esaminato: Zaire: 16, Musée du Congo, Elisabethville, XI.1936, Ch.
Seydel, GS 2556 GB, TM; 19, Ht. Katanga, Tshinkolobwe, 8.1.[19]31, J. Romieux, GS 2211
GB (1429 MHNG), MHNG; 1 2, idem, Panda, 14.1.[19]30, GS 2210 GB, CB.
Apparato genitale d (Fig.15): E' caratterizzato dalla forma del processo costale,
i cui due lobi terminano entrambe con punte aguzze, e dalla forma del phallus (Fig. 16),
che presenta all'apice due ampi processi simmetrici. L'esemplare esaminato non è
dotato di cornuti.
Apparato genitale © (Fig.26): La bursa copulatrix "densamente corrugata" des-
critta per l'Holotypus è frutto di un'aberrazione del preparato microscopico. In realtà
essa è lunga, stretta e solo debolmente corrugata. La placca urosternale posta al di sopra
dell'ostium bursae è ben definita ma senza dentelli.
Alphacrambus gen. nov.
Specie-tipo: Crambus prodontellus Hampson, 1919. Genere maschile.
Genere caratterizzato dalle piccole dimensioni (mm. 12-15,5 di apertura alare).
Antenne debolmente serrate nel d, lisce nella 9. Ocelli e chetosemae ben sviluppati.
Ali anteriori bianche o gialle variegate da strie e bande dorate e/o brune. Banda
submarginale argenteo-dorata sempre presente. Il frenulum nella 9 è semplice.
Apparato genitale d: si distingue per l'assenza dell'uncus e per la completa
fusione tra tegumen e vinculum. Il vinculum è sempre più o meno profondamente bilo-
bato, la costa alta e sclerificata, lo pseudosaccus ben sviluppato e vi è sempre la
presenza di cornuti nella vesica.
622 GRAZIANO BASSI
Fico. 19-25
Sebrus e Alphacrambus spp., apparati genitali 2 2 e organi timpanali. Scala 0,5 mm. 19: S.
perdentellus (Hmps.), lectoparatypus, GS3228GB, fino a ductus bursae escluso. 20: S.
absconditus n. sp., paratypus, GS3225GB. 21: S. amandus Btesz., Madagascar, GS2628GB, parte
del ductus bursae e bursa copulatrix esclusi. 22-23: variabilità nel complesso ostium bursae -
antrum in A. prodontellus (Hmps.). 22- GS2192GB, Sud Africa, Pretoria. 23- GS3191GB, Sud
Africa, Magunde. 24-25: organi timpanali. 24- S. absconditus n. sp., 6, paratypus, GS3064GB.
25- A. prodontellus (Hmps.), 9, GS3191GB.
NOTE SUI GENERI SEBRUS E ALPHACRAMBUS 623
Apparato genitale 9: VIII segmento addominale completamente membranoso;
apophyses posteriores molto corte; apophyses anteriores assenti; ostium bursae for-
mante ampi processi periostiali di supporto; antrum non differenziato; ductus bursae
mediamente sclerificato nella sua prima metà, poi membranoso; ductus seminalis origi-
nantesi alla metà circa del ductus bursae; bursa copulatrix di media grandezza, dotata di
un signum ed intensamente corrugata e/o cosparsa di microplacche regolarmente di-
sposte.
Gli apparati genitali tendono ad avere lievi variazioni (Figg.22-23) non costanti
anche all'interno di popolazioni simpatriche.
Organi timpanali (Fig.25): caratterizzati dalla bulla tympani, dal pons tympani e
dal processus tympani minuti, dall'area subtimpanale inferiore ampia. Venulae secun-
dae ben sviluppate. Nel d sia il tympanum che la bulla tympani sono più sviluppati.
Il genere è molto vicino a Crambixon Bteszyñski, nel cui genitale d si osserva
l'assenza di cornuti, tegumen esile e giunzione tegumen/vinculum evidente. Nel genitale
2 la bursa copulatrix è priva di signa, ostium bursae ed area periostiale sono solo
debolmente sclerificati. Il ductus seminalis origina nella seconda metà del ductus
bursae.
Geonemia: Africa tropicale, India.
TABELLA DELLE SPECIE
1 Colore di fondo delle ali anteriori giallo.................... phoeostrigellus
— Color dijfondodellealianterion DIanco =. E ee TE 2
2 apice dell'alafanternore acuto le en ee 3
- apice dell'ala anteriore decisamente appuntito, banda submarginale
SITE TARN POE II razowskii
3 ala anteriore con 2 macchie nere marginali distanziate tra loro. ......... parvus
- ala anteriore con 4 macchie nere marginali in sequenza. .................. 4
4 fronte tondeggiante, appena sporgente oltre l'occhio... .......... prodontellus
fronte subconica, decisamente sporgente oltre l'occhio.............. cristatus
Alphacrambus prodontellus (Hampson, 1919) n. comb.
Crambus prodontellus Hampson, 1919: 288.
Holotypus d: [Sud Africa, Transvaal] Pretoria, 17.IX.[19]08, 11, A.J.T. Janse,
GS5576BM Pyral., nel NHML, esaminato.
Altro materiale esaminato: Botswana: 19 Gaborone, XII. 1977, B. Skule, GS2883GB,
ZMC. Kenya: 2 d d, Naro Moru, m. 1950, 18.XI-3.XII.1984, lux, Leg. G. Bassi, GS 1965 e 1969
GB, CB. Nigeria: 1d Jos, 4000 ft., D.M. Roberts, 21. VII.1977, B.M. 1977:377, GS1650GB,
NHML; Sud Africa:14 exx. dd e 2 9 Pretoria, Magunde, I-INI-VII-X-XI-XII, TMP; 19,
Pretoria, I.10.09, A.J.T. Janse, GS 1649 GB, NHML. Tanzania: 11 exx. dd e 8 9, Lake Sereri,
3150 ft., VIII-IX.1965, leg. Dr. Szunyoghy, GS 3252, 3286 e 3296 GB, TMB. Zaire: 1d,Ht.
Katanga, Kyala, 29.VIII.29, J. Romieux ,GS2233GB, MHNG.
Alphacrambus razowskii (Bteszyfiski, 1961) n. comb.
Crambus razowskii Bteszyriski, 1961: 191.
Holotypus d: Cape Province, Mossel Bay, XII.1934, S[outh] Africa, R. E. Turner,
GS1561SB (5265BM Pyral.), nel NHML, esaminato.
624 GRAZIANO BASSI
FIGG. 26-29
Sebrus e Alphacrambus spp., apparati genitali 22. Scala 0,5 mm. 26: S. pseudosparsellus
(Blesz.), Zaire, Panda, GS2210GB. 27: S. argus n. sp., paratypus, GS6677SB. 28: A. prodon-
tellus (Hmps.), Sud Africa, Pretoria, GS1649GB. 29: A. cristatus n. sp., holotypus.
NOTE SUI GENERI SEBRUS E ALPHACRAMBUS 625
Altro materiale esaminato: Kenya: 11 exx. dd e 9 9, Naro Moru, m. 1950, 18.XI-
3.XII.1984, lux, Leg. G. Bassi, GS 1221, 1229, 1234 e 1970 GB, CB. Sud Africa: 6 9 9,
Pretoria, Barberton, Umkomaas, XII e III, TMP; 12, Natal, Coll. Dumont, 10.04, GS 2604GB,
MNHNP. Tanzania: | 6, Mafinga, 20.XI-5.XII.1989, lux, Leg. Curletti, GS 2673 GB, CB. Zaire:
12, Ht. Katanga, Panda, 12.X.29, J.Romieux, GS2234GB e 12, Ht. Katanga, Tshituru, 7.1V.29,
J.Romieux, nel MHNG.
Nota sistematica: le 22 e i dd delle due specie non furono correttamente
accoppiati da Bteszynski (1961: 190-192, figg. 74-75). In realtà sia le descrizioni che le
figure riguardanti le 9 9 vanno, nel citato lavoro, invertite. Esaminando serie di
esemplari in buone condizioni distinguere le due specie è facile: prodontellus (Hmps.) è
leggermente più grande e più scuro, ha l'apice dell'ala anteriore meno appuntito, ha una
grande macchia subtriangolare nera all'apice (bruno argentea in razowskii Btesz.), la
banda subterminale è più ampia con colore di fondo bianco, vi sono 4 grandi tacche
nere submarginali (in razowskii Btesz. la banda subterminale è stretta e con colore di
fondo giallo argenteo, le 3-4 tacche submarginali sono più minute). La fronte è, in pro-
dontellus (Hmps.), più tondeggiante e meno sporgente oltre l'occhio rispetto a razowskii
Btesz.
Alphacrambus phoeostrigellus (Hampson, 1903) n. comb.
Crambus phoeostrigellus Hampson, 1903: 658.
Crambixon phoeostrigellus (Hmps.), Bteszynski 1965: 325.
Lectotypus d (stabilito da Bleszynski, 1965:325): [India] Kashmir, Goorais Valley,
VI.1887, J.H. Leech, nel NHML.
Materiale esaminato: 14 paralectotypo, stessa etichetta del lectotypus, sept. 1887,
GS3411GB, NHML.
Per quanto sopra esplicitato la specie in questione è tipica rappresentante del
genere qui trattato. Dall'apparato genitale (si conoscono solo d d) appare vicina a
prodontellus (Hmps.).
Alphacrambus cristatus n. sp.
Holotypus ©: [Etiopia] Abyssinia, Kovacs, Marako, 1912.IIT. 8, GS 3273 GB, nel TMB.
Cenni morfologici (Fig.D): apertura alare mm. 12,7. Fronte bianca, subconica,
nettamente sporgente oltre l'occhio. Palpi lunghi 3,5 volte il diametro dell'occhio,
bianchi superiormente ed internamente, bronzei esternamente. Antenne lisce, bianche e
bronzee. Capo, collo e torace bianchi. Tegulae bronzee. Colore di fondo delle ali
anteriori bronzeo. Ali posteriori bianche soffuse di bruno.
Apparato genitale © (Fig.29): Simile a prodontellus (Hmps.) (Fig. 28), ma con
placca periostiale molto alta e bilobata.
d sconosciuto.
Derivatio nominis: da cristatus = con cresta, in allusione al processo dell'ostium
bursae, ad essa simile.
Alphacrambus parvus n. sp.
Holotypus d: Fort Crampel, (Congo français), 1920-1932, Coll. L. & J. de Joannis,
Muséum Paris, GS 2436 GB, nel MNHNP.
Cenni morfologici (Fig. E): apertura alare mm. 12,1. Fronte tondeggiante, bian-
ca, appena sporgente oltre l'occhio. Antenne brune, debolmente serrate. Capo e torace
626 GRAZIANO BASSI
Ficc. A - E
Sebrus e Alphacrambus spp., imagini. A: S. argus n. sp., holotypus. B: S. perdentellus (Hmps.),
paralectotypus 9. C: S. absconditus n. sp., paratypus d. D: A. cristatus n. sp., holotypus. E: A.
parvus n. sp., holotypus.
bianchi. Tegulae brune. Ali anteriori con un'ampia e irregolare stria bruna che attraversa
medialmente l'ala in tutta la sua lunghezza. Due sole tacche nere submarginali, dis-
tanziate tra loro. Ali posteriori brune. Simile a prodontellus (Hmps.), ma più piccolo e
più scuro.
Apparato genitale 4 (Fig. 17, 18): Simile a prodontellus (Hmps.) (Fig. 13, 14),
con gnathos più corto, vinculum lievemente più stretto e più ampiamente bilobato e
costa molto più corta ed alta.
2 sconosciuta.
Derivatio nominis: da parvus = piccolo, in allusione alle minute dimensioni
dell'Holotypus.
NOTE SUI GENERI SEBRUS E ALPHACRAMBUS 627
RINGRAZIAMENTI
Sono grato ai Dott. V. Mahnert e D. Burckhardt del Muséum d'histoire naturelle
di Ginevra, al Dott. U. dall'Asta del Musée Royal de l'Afrique Centrale di Tervuren, al
Dott. L. Gozmany del Magyar Termeszettudomanyi Mùzeum di Budapest, al Sig. O.
Karsholt dello Zoologisk Museum di Copenaghen, al Sig. M. Kriiger del Transvaal
Museum di Pretoria, al Dott. G. Luquet del Muséum National d'Histoire Naturelle di
Parigi, ed al Sig. M. Shaffer del Natural History Museum di Londra per avermi
concesso in studio il materiale a loro affidato. Un sentito ringraziamento anche al Prof.
U. Parenti del Dipartimento di Biologia animale dell'Università di Torino per i preziosi
consigli di cui è sempre prodigo.
BIBLIOGRAFIA
Bassi, G., 1991. Contributi alla conoscenza delle Crambinae (Lepidoptera, Crambidae). VI: Note
sul genere Aureocramboides Bteszynski. Boll. Mus. reg. Sci. nat. Torino, 9 (2):391-396.
Bassi, G., 1992. Contributi alla conoscenza delle Crambinae (Lepidoptera, Crambidae). VII:
Note sulle specie africane di Crambus Fabricius presenti nel Muséum d'Histoire Naturelle
di Parigi. Boll. Mus. reg. Sci. nat. Torino, 10 (2): 221-235.
Bassı, G., 1995. Contributi allo studio delle Crambinae (Lepidoptera: Crambidae): VIII. Note sul
genere Caffocrambus Bteszyfiski, con descrizione di nuove specie. Boll. Mus. reg. Sci.
nat. Torino, 13 (1), in stampa.
BLESZYNSKI, S., 1961. Studies on the Crambidae (Lepidoptera). Part XXX. On several species of
the generic group Crambus F. from the Ethiopian Region with the descriptions of new
genera and species. Pol. Pis. ent., 31: 165-218.
BLESZYNSKI, S., 1965. Crambinae. In: Amsel, H.G., Gregor, F., Reisser, H., Microlepidoptera
Palearctica, 1: 1-553.
BLESZYNSKI, S., 1970. New genera and species of tropical Crambinae (Studies on the Crambinae,
Lepidoptera, Pyralidae, Part 48). Tijdschr. Ent.,113, 1: 1-26.
GASKIN, D.E., 1975. Revision of the New Zealand Crambini (Lepidoptera: Pyralidae:
Crambinae). N.Z. Journal of Zoology, 2 (3) :265-363.
Hampson, G.F., 1903. The moths of India. J. Bomb. natur. Hist. Soc., 14: 639-659.
Hampson, G.F., 1919. Description of new Pyralidae of the subfamilies Crambinae and Siginae.
Ann. Mag. Nat. Hist., (9), 3:275-292, 437-457.
REVUE SUISSE DE ZOOLOGIE, 102 (3) : 629-634; septembre 1995
Un Anapleus Horn nouveau de |’ Himalaya
(Coleoptera, Histeridae)
Yves GOMY
116, rue Dalayrac, F-94120 Fontenay-sous-Bois, France.
A new species of Anapleus Horn from the Himalaya (Coleoptera
Histeridae).- Anapleus davidneelae sp.n. is described from Nepal and
North India. Anapleus stigmaticus (Schmidt) is revalided. A key to the
Oriental species of Anapleus is provided. The relationships of Anapleus are
discussed: the genus possesses a combination of characters of the Dendro-
philinae and Onthophilinae.
Key-words: Coleoptera - Histeridae - Anapleus - Taxonomy - Oriental
region.
INTRODUCTION
Dans une première note consacrée aux Histeridae du Népal, MAZUR (1987a)
signale une espèces d'Anapleus Horn comme probablement nouvelle en plus d'Ana-
pleus cyclonotus (Lewis, 1892). Dans sa seconde note sur cette région, le méme
auteur (MAZUR, 1991) signale de nouveau A. cyclonotus (Lewis) mais ne parle plus de
l'existence possible d'une autre espece. Par ailleurs, OLEXA (1982), dans sa révision
des Anapleus paléarctiques ne mentionne pas A. cyclonotus et considére donc que
cette espèce appartient, avec son synonyme Abraeus stigmaticus Schmidt, 1892, a la
faune orientale. Cette synonymie, signalée par BICKHARDT (1913), ne parait pas, a
notre connaissance, avoir fait l'objet d'une nouvelle étude et si le Népal et l'Etat de
Perak (Malaisie) nous semblaient assez éloignés géographiquement pour ne pas trop
craindre la synonymie entre Anapleus davidneelae n.sp. et Abraeus stigmaticus
Schmidt une vérification s'imposait cependant.
Ainsi, grace a l'obligeance de notre collegue M. Uhlig du Zoologisches Mu-
seum de Berlin (Allemagne), nous avons recu pour étude l’unique syntype d'Abraeus
stigmaticus Schmidt conservé a Berlin, ce qui nous permet de désigner un lectotype. Il
s’agit d’un mâle portant les étiquettes suivantes: / mâle (étiquette blanche, manus-
crite). / stigmaticus Type Perak (étiquette blanche, manuscrite, de la main de J.
Schmidt) / Type (petite étiquette orange, carrée, imprimée, caractéristique des types
de la Coll. Schmidt-Bickhardt) / Coll. J. Schmidt (petite étiquette blanche, allongée et
imprimée) / Zool. Mus. Berlin (étiquette jaune clair, imprimée).
Manuscrit accepté le 23.01.1995.
630 YVES GOMY
Ce Muséum possède un second mâle d'Anapleus stigmaticus (Schmidt) déter-
mine A. stigmosus (sic.) provenant d'Indonésie: Mentawei, Sipora, Sereinu, V.VI.1894
(Modigliani) ex. Mus. Civ. de Genova; un troisième exemplaire de même provenance
est conservé au Muséum de Génes (R. Poggi, comm. pers.). Ces exemplaires se
différencient d'Anapleus cyclonotus (Lewis), par la forme générale moins globuleuse,
la taille plus petite, la couleur moins sombre, la ponctuation et la forme de l'édéage.
La synonymie proposée par BICKHARDT (1913) entre A. cyclonotus et A. stigmaticus
n'est pas valable. Anapleus stigmaticus (Schmidt) est donc, à notre avis une espèce
distincte. Nous donnons ci-dessous des figures permetttant de différencier ces deux
espèces ainsi que celle que nous décrivons dans la présente note.
Anapleus davidneelae n.sp.
Espece beaucoup plus proche d'Anapleus stigmaticus (Schmidt) que d'A. cyclo-
notus (Lewis). Elle se distingue immédiatement de ces deux espèces par la synthèse
habituelle des caractères spécifiques distinctifs: taille, ponctuation générale forme du
prosternum forme de l'édéage etc... Elle se rapproche aussi d'A. jelineki Olexa d'Iran
et d'A. monticola Mazur du Pakistan mais s'en sépare principalement par sa ponc-
tuation, la forme de son prosternum et celle de son édéage.
Corps en ovale large, convexe (beaucoup moins arrondi que chez A. cyclo-
notus). Noir, billant. Pattes, scape et funicule antennaire sensiblement de la méme
couleur, parfois très légèrement éclaircis; massues sombres, éclaircies superficielle-
ment par une très fine pubescence claire. Scape renflé, environ trois fois plus long que
large; premier article du funicule allongé, deux fois plus long que large: 2ème et 3ème
articles très petits, les autres s'élargissant légèrement vers la massue.
Téte relativement petite. Front légèrement convexe a ponctuation forte, dense
mais irrégulière car les points (séparés par 1/2 a 1 de leurs diametres environ) ne sont
pas de taille homogène (mélange de gros points et de points plus petits); marqué par
une large impression au milieu; entaillé devant, de chaque côté au dessus des yeux par
une petite encoche d'insertion antennaire. Epistome convexe, sans séparation visible
avec le front, à ponctuation de méme type mais plus petite, plus serrée, donnant un
aspect rugueux. Yeux à peine bombés peu visibles de dessus.
Pronotum un peu plus de deux fois plus large que long à la base (1,14 mm /
0,53 mm); convexe, à ponctuation forte, dense mais irrégulière: les points sont plus
petits et plus régulièrement denses sur le disque, plus gros et espacés a la base et
surtout sur les côtés avec deux impressions post-oculaires qui lui donnent un aspect
un peu plus bombé sur le disque; strie marginale large, complète derrière la téte et
relevée en léger rebord émoussé jusqu'aux angles visibles puis se poursuivant de
même jusqu'à la base en position d'épipleures. Les bords apparents du pronotum
(visibles de dessus) ne correspondent, en fait, qu'à un bourrelet latéral caractéristique
du genre pour les espèces que je connais. Base en arc large et bisinué.
Scutellum très visible, en triangle équilatéral.
UN ANAPLEUS DE L’ HIMALAYA 631
Elytres pris ensemble convexes, plus larges aux épaules que longs au milieu
(1,40 mm / 1,14 mm); caractérisés par leur bourrelet latéral large et émoussé, plus fort
dans la région humérale; à ponctuation forte et irrégulière (mélange de points plus ou
moins gros) mais beaucoup plus lache que celle du pronotum (points séparés par | a 4
de leurs diamètres environ) et plus fine dans la région périscutellaire. Stries dorsales
élytrales obsolètes, réduites a des rudiments latéraux légèrement costiformes plus
longs à l'extérieur qu'à l'intérieur. Apparemment, les deux stries subhumérales se sont
transformées pour donner le bourrelet caractéristique. Strie marginale forte, très
abaissée en position épipleurale, non visible de dessus, ponctuée nettement. Apex
élytral avec une marge lisse précédée par quelques strioles longitudinales un peu plus
visibles sur les côtés qu'au milieu (chez l'holotype mâle) un peu plus nettes et plus
étendues chez le paratype femelle de l'Inde du Nord (dimorphisme sexuel). Epipleures
avec une longue strie complete, ponctuée, bisinuée. Espace entre cette strie épipleu-
rale et la strie marginale avec quelques points nets mais espacés.
Propygidium étroit, densément ponctué (points séparés par 1/2 a 1 de leurs
diamètres environ). Pygidium a ponctuation du méme type, mais nettement plus
grosse.
Lobe prosternal court, à mentonnière trapézoidale et nettement rebordée, gros-
sièrement et rugueusement ponctuée. Prosternum proprement dit plus large que long
(ce qui la sépare à première vue d'A. cyclonotus (fig.2). Stries prosternales sub-
parallèles, très courtement convergentes au sommet, a ponctuation forte et rugueuse le
long de la mentonnière, plus fine et espacée sur le disque et a la base. Séparé de la
mentonnière par une ligne peu nette. Mésosternum (fig.2) très court, rectangulaire, a
strie marginale forte sur les côtés et dans les angles antérieurs, interrompue derrière le
prosternum; ponctué fortement et irrégulièrement dans les 2/3 apicaux, lisse dans le
tiers basal. Suture méso-métasternale droite, ponctuée crénelée. Métasternum avec
une fine ligne longitudinale médiane; a ponctuation forte et irrégulière: beaucoup plus
forte et espacée sur les côtés et dans la moitié apicale, plus fine sur le disque et plus
fine et serrée le long de la ligne longitudinale (fig.2). Strie métasternale latérale en
deux trongons: trongon interne très fort, large et ponctué de très gros points, incom-
plet, ne marquant pas l'arrondi sous les hanches intermédiaires et s'arrétant un peu
avant de rencontrer le trongon externe. Celui-ci rectiligne, oblique, très long, rejoi-
gnant presque les hanches postérieures. Ce tronçon délimite avec le bord épipleural
une zone large et ponctuée de très gros points (séparés par 1/2 à 1 de leurs diamètres
environ) et avec quelques points plus petits. Mésépiméron triangulaire avec une
grosse ponctuation du méme type. Plaque méso-postcoxale sans strie postcoxale,
ponctuée de très gros points.
Premier sternite abdominal à ponctuation forte, dense et double; strie méta-
postcoxale forte et ponctuée, ne marquant pas l'arrondi sous les hanches postérieures.
Tibias antérieurs caractéristiques du genre, à arétes externes (supérieure et
inférieure) subparallèles, doublement coudées, formant une sorte de gouttière. Cette
disposition soulignant l'épaisseur du tarse se retrouve également pour les tarses inter-
médiaires et postérieurs. Toutes les arétes sont frangées de courtes épines.
YVES GOMY
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FIGS 1-6
1-3: Ponctuation et striation schématiques des régions pro-méso-métasternales chez Anapleus
cyclonotus (Lewis) (1), A. davidneelae n.sp. (2) et A. stigmaticus (Schmidt) (3). Echelle = 0,5
mm. 4-6: Edéage (vue ventrale) chez les mêmes espèces et dans le même ordre. (4. sur un ex.
du Népal, long: 0,36 mm); (5. sur holotype, long: 0,38 mm); (6. sur holotype, long: 0,47 mm).
Echelle = 0,1 mm environ.
UN ANAPLEUS DE L’ HIMALAYA 633
Edéage: long. 0,38 mm (fig. 5) bien différent de celui de A. cyclonotus (Lewis)
qui est assymétrique (fig. 4) de celui de A. stigmaticus (Schmidt) (fig. 6) et de A.
monticola (Mazur, 1987b).
Long: 1,67 mm (téte et pygidium exclus); large: 1,40 mm (holotype).
Derivatio nominis: Espèce d'altitude dédiée a la mémoire et aux exploits
d'Alexandra David-Néel (1868-1969), l'une des grandes dames de ce siècle,
aventurière de l'esprit, première européenne a pénétrer le Tibet interdit et sa capitale
Lhassa. Elle fut, sa vie durant fascinée par les Himalayas, ses peuples et ses traditions
et elle participa activement a répandre en France la connaissance du Bouddhisme
tibétain.
Hototype d : Est Nepal: Kosi, Col N-E Mangmaya (2300 m), 6.1V.1984 (I. Löbl et A.
Smetana). Muséum de Genève.
Paratypes: 2 males, Est Nepal: Kosi, Val Induwa Kola (2000 m.), 14.1V.1984 (I. Löbl et
A. Smetana ) Coll. Y Gomy; 1 femelle, Inde: Kumaon (U.P.), Rangarh no 68 (env. 2000m.),
9.X.1979 (I. Löbl), Muséum de Genève; 2 ex. W. Nepal, Dampa Pab, près de Chauta, région du
lac Rara (Pa. 2294), (H. Franz), (Coll. H. Franz, Mödling et Y Gomy, Paris).
CLE DES Anapleus DE LA FAUNE ORIENTALE
1 Prosternum plus long que large (fig. 1). Taille relativement grande (2
mm); forme très arrondie. Birmanie, Inde du Nord, Népal. . cyclonotus (Lewis)
- Prosternum plus large que long (fig. 2 et 3). Taille plus réduite (infé-
Heu) orme un peuplusiovalalre = ea 2
2 Espèce noire. Ponctuation élytrale irrégulière, plus ou moins fine et
espacée (points séparés par un à trois ou quatre de leurs diamètres
environ). Ponctuation du métasternum fine et dense le long de la ligne
médiane (fig. 2). Edéage symétrique (fig. 5). Inde du Nord, Népal
5 SE OS We nl PPS COVE CERI EE, Dear WER RL, davidneelae n.sp.
— Espèce brune. Ponctuation élytrale régulière, forte et dense comme
chez A. cyclonotus. Ponctuation du métasternum fine mais espacée le
long de la ligne médiane. Edéage avec un début de dissymetrie (fig. 6).
Malaisie/((Rerao Indonésie em re PE ee stigmaticus (Schmidt)
CONSIDÉRATIONS SUR LA POSITION SYSTÉMATIQUE DE GENRE Anapleus HORN
La description détaillée d'Anapleus davidneelae nous a permis de constater
qu'un nombre non négligeable de caractères génériques rapproche les Anapleus Horn
(1873) et les Onthophilus Leach (1817).
La forme générale du corps; l'encoche frontale d'insertion antennaire; la forme
de la tête, de l'épistome et des yeux; la légère gibbosité du disque pronotal due aux
fossettes post-oculaires; la configuration très particulière des bords du pronotum; la
634 YVES GOMY
forme de la base du pronotum; la ponctuation générale; les rudiments de striation
élytrale costiformes et les bourrelets latéraux des élytres; la forme de la mentionnière
et du prosternum; la forme du mésosternum; la ligne longitudinale médiane du méta-
sternum; la configuration des côtés du métasternum, du mésépiméron et de la plaque
meso-postcoxale; la forme des tibias antérieurs; la forme de l'édéage et de sa pièce
basale sont des caractères rapprochant Anapleus des Onthophilinae.
Si les Anapleus Horn ont été longtemps placés dans les Dendrophilinae à cause
de leurs tibias antérieurs (entre autres), ce caractère ne résiste pas à un examen
approfondi car l'on trouve chez les différentes espèces d'Onthophilus des tibias
antérieurs grèles certes, mais dont la structure n'est pas foncièrement différente de
celle des Anapleus. Par contre, le lobe prosternal des Anapleus possède une incision
antennaire alors que celle-ci n'existe pas chez les Onthophilus. En fait, il semblerait
que le lobe prosternal ait évolué chez les Onthophilus par tassement et élargissement
de la mentonnière corrélés avec l’évolution d’une alvéole antennaire sous l'angle
antérieur du pronotum. Il serait évidemment intéressant de vérifier ces hypothèses à
partir d'une étude plus complète.
Le genre Anapleus Horn ne répond pas aux définitions des sous-familles des
Dendrophilinae ni des Onthophilinae, telle qu'elles sont acceptées actuellement. Nous
laissons à d'autres histéridologues le soin d'infirmer ou de confirmer nos présomptions
et de redéfinir ces sous-familles.
REMERCIEMENTS
Nous remercions très cordialement nos amis I. Löbl et H. Franz pour le prêt du
matériel ayant servi de base a cette note, ainsi que les Dr. M. Uhlig et R. Poggi
respectivement du Zoologisches Museum de Berlin (Allemagne) et du Muséum de
Génes (Italie) pour nous avoir permis de vérifier les synonymies établies par H.
Bickhardt.
REFERENCES
BICKHARDT, H. 1913. Histeridenstudien (19. Beitrag zur Kenntnis der Histeriden). Dt. ent. Z.:
696-701.
Lewis, G. 1892. Viaggio di Leonardo Fea in Birmania e regioni vicine. XLII Histeridae
(Part.2). Annali Mus.civ. Stor.nat. Giacomo Doria (2), 12 (32): 16-39.
Mazur, S. 1984. World catalogue of Histeridae. Polskie Pismo ent. 54 (3-4): 1-379.
Mazur, S.1987a. Histeridae from the Nepal Himalayas (Insecta: Coleoptera). Cour. For-
schungsinst. Senckenb. 93: 463-470.
MAZUR, S. 1987b. Contribution to the knowledge of the Histeridae of Pakistan (Coleoptera).
Revue suisse Zool. 94(4): 659-670.
MAZUR, S. 1991. Histeridae from the Nepal Himalayas, II. (Insecta: Coleoptera). Stuttgarter
Beitr. Naturk., Ser.A. 467: 1-12.
OLEXA, A. 1982. Revision der paläarktischen Arten der Gattung Anapleus (Coleoptera, His-
teridae). Acta ent. bohemoslav. 79: 37-45.
SCHMIDT, J. 1882. Neue Histeriden (Coleoptera). Ent.Nachr. 18: 17-30.
REVUE SUISSE DE ZOOLOGIE, 102 (3) : 635-653; septembre 1995
Oswaldocruzia (Nematoda, Trichostrongylina, Molineoidea)
parasites d'Amphibiens du Brésil et de l'Equateur, avec redéfinition
de l'espèce-type O. subauricularis (Rudolphi, 1819)
et d'O. mazzai Travassos, 1935
Badreddine BEN SLIMANE & Marie-Claude DURETTE-DESSET
Laboratoire de Biologie parasitaire, Protistologie, Helminthologie, associé au
C.N.R.S., Muséum national d'histoire naturelle, 61 rue de Buffon,
F-75231 Paris cedex 05, France.
Oswaldocruzia (Nematoda, Trichostrongylina, Molineoidea) parasitic
in Brazilian and Ecuadorian Amphibians, with redefinition of the type
species O. subauricularis (Rudolphi, 1819) and O. mazzai Travassos,
1935.- O. subauricularis (Rudolphi, 1819), type species of Oswaldocruzia
Travassos, 1937, parasite from Brazilian and Ecuadorian Bufonidae and O.
mazzai Travassos, 1935, parasite from Argentinian Bufonidae are redes-
cribed using new morphological characters, particularly the relative arran-
gement of the bursal rays 6,8 and 9 and the pattern of the synlophe in the
oesophagal region. Oswaldocruzia dlouhyi n.sp. coparasite of O. subau-
ricularis from Bufo sp. and Oswaldocruzia taranchoni n.sp. fom Bufo
marinus are described from Brasil. O. dlouhyi belongs to the Oswaldo-
cruzia species group having cervical alae and rays 8 arising on the dorsal
ray and overlapped by rays 6 along their median part (type II).The species
is characterised by a cephalic vesicle in two parts and by rays 4 longer than
rays 5. O. taranchoni belongs to the Oswaldocruzia species group having
no cervical alae and rays 8 arising on the root of the dorsal ray and joined
to rays 6 along the two thirds (type III). The species is characterised by a
cephalic vesicle in one part, by the spicular externo-lateral branch (blade)
not divided on its tip and by rays 8 joined to the common trunk of rays 4 to
6. With just one exception, the neotropical Oswaldocruzia have spicula
divided in three main branches: blade, shoe and fork as is the case in
holarctic Oswaldocruzia but the division of the fork always occurs before
the distal third of its length in neotropical species and beyond the distal
third in holarctic ones. A new taxon is proposed: Oswaldocruzia proencai
n.sp.(= O. mazzai sensu Lent et al., 1946 nec Travassos, 1935).
Key-words: Oswaldocruzia spp. - Nematoda - Trichostrongylina - Bufo-
nidae - Neotropics - Taxonomy.
Manuscrit accepté le 12.12.1994.
636 BADREDDINE BEN SLIMANE & MARIE-CLAUDE DURETTE-DESSET
INTRODUCTION
Le genre Oswaldocruzia parasite cosmopolite d'Amphibiens et de Reptiles a
été créé par TRAVASSOS (1917) avec pour espèce type Oswaldocruzia subauricularis.
Cette espèce avait été décrite du Brésil, de façon très succincte par Rudolphi (1819)
chez "Rana musica", c'est-à-dire Bufo americanus dans la nomenclature actuelle. Les
neuf espèces connues actuellement en zone néotropicale se répartissent en trois
groupes d'après la disposition des côtes 6, 8 et 9 de la bourse caudale (DURETTE-
DESSET et al., 1992). L'objet principal de ce travail est d'identifier dans la faune néo-
tropicale l'espèce correspondante à celle de Travassos, premier redescripteur moderne
puisque comme nous le verrons dans la discussion, les spécimens types n'ont pas pu
être localisés. Grâce à du matériel provenant de Bufonidae du Brésil et d'Équateur, il a
été possible de trouver une espèce qui s'accorde avec la redescription d'O. subauri-
cularis de Travassos. Celle-ci est redécrite ainsi que O. mazzai en utilisant de nou-
veaux critères morphologiques, en particulier les caractéristiques du synlophe dans la
région oesophagienne. Enfin deux nouvelles espèces sont décrites chez des Bufonidae
du Brésil.
MATÉRIEL ET MÉTHODES
Les Nématodes proviennent de l'intestin grêle d'un Bufo sp. originaire du
Brésil et de 3 Bufo marinus dont l'un est originaire du Brésil et les 2 autres d'Équa-
teur.
La nomenclature des Strongylida utilisée au-dessus du groupe famille est celle
de DURETTE-DESSET & CHABAUD (1993).
Le synlophe est étudié selon la méthode de DURETTE-DESSET (1985); la
nomenclature utilisée pour l'étude du synlophe dans la région oesophagienne est celle
de BEN SLIMANE & DURETTE-DESSET (1993). En particulier, l'aile cervicale est définie
comme formée d'une ou de plusieurs crêtes latéro-ventrales qui peuvent s'observer
immédiatement en vue médiane lorsqu'elles sont très développées ou bien n'apparaître
qu'en coupe transversale lorsqu'elles sont petites mais dans ce cas, elles sont toujours
plus développées que les crêtes adjacentes.
La nomenclature utilisée pour l'étude de la bourse caudale est celle de
DURETTE-DESSET & CHABAUD (1981); celle concernant la naissance et le trajet des
côtes 8 par rapport à la côte dorsale et aux côtes 6 est celle suivie par DURETTE-
DESSET, NASHER & BEN SLIMANE (1992) modifiée (fig. 1).
L'étude complète des spicules est faite sur du matériel disséqué et la
nomenclature utilisée celle de BEN SLIMANE & DURETTE-DESSET (1993).
Le matériel étudié, conservé dans l'alcool à 70°, est déposé dans les
Collections du Muséum national d'Histoire naturelle de Paris (MNHN), du Muséum
d'Histoire naturelle de Genève (MHNG) et de l'Institut Oswaldo Cruz.
Les mensurations sont exprimées en micromètres.
OSWALDOCRUZIA DU BRÉSIL ET DE L’EQUATEUR Ga
Bb
FIG. 1
Disposition relative des côtes 6, 8 et de la côte dorsale. A: type I, les côtes 8 naissent sur la côte
dorsale et sont séparées des côtes 6 sur tout leur trajet; B: type II, les côtes 8 naissent apicale-
ment sur la côte dorsale et apparaissent, en vue ventrale, chevauchées par les côtes 6 dans leur
portion médiane (a) ou exceptionnellement sont légèrement écartées des côtes 6 et suivent un
trajet parallèle à ces dernières (b); C: type III, les côtes 8 naissent a la racine de la côte dorsale
et sont soit jointives aux côtes 6 soit chevauchées par ces dernières dans leur deux tiers
proximaux (d'après Durette-Desset et al.1992, modifié).
Oswaldocruzia subauricularis (Rudolphi, 1819)
Matériel étudié: Un d MNHN 882 MD parasite de l'intestin grêle de Bufo sp., Gavioes
(134 km au Nord de Rio de Janeiro), Brésil, 10.09.1985, coparasite d'O. dlouhyi n.sp., leg.
C.Dlouhy. Trois 4,4 2 MNHN 712 MD, 2 6,4 © MHNG 18768 INVE, parasites de l'intestin
gréle d'un Bufo marinus, San Lorenzo (Esmaraldas), Equateur, 2.03.1987, leg. J.M. Touzet.
Pour permettre une éventuelle révaluation des caractères, les mensurations du
mâle originaire du Brésil sont séparées de celles des mâles originaires d'Équateur.
Description: Nématodes ne présentant pas d'enroulement. Vésicule céphalique
formée d'une partie antérieure enflée et d'une partie postérieure fine (fig. 2, A; 3, A).
Pore excréteur toujours situé dans le tiers postérieur de l'oesophage. Deirides de forme
triangulaire, situées postérieurement au pore excréteur (fig.3,A). Glandes excrétrices
très développées. Séparation oesophage musculaire et glandulaire très nette (fig. 2, A;
3, A). Présence de minuscules ailes cervicales, visibles seulement en coupe trans-
versale du corps. Crêtes cuticulaires pourvues d'un soutien chitinoide (fig. 2, D; 3, E).
Tête: (fig.3,B). Bouche triangulaire, arrondie aux angles. En vue apicale, pré-
sence de 6 papilles labiales externes dont les latérales sont accolées aux 2 amphides et
de 4 papilles céphaliques. Petite dent oesophagienne dorsale présente.
Synlophe: (étudié chez 3 mâles et une femelle. Les chiffres relatifs au mâle
originaire du Brésil sont indiqués en premier, ceux d'Équateur entre parenthèses).
Dans les deux sexes, corps parcouru longitudinalement par des crêtes cuticulaires
continues. Chez le mâle, la grande majorité des crêtes dorsales et les deux tiers des
crêtes ventrales naissent dans la région oesophagienne. Chez la femelle, les deux tiers
des crêtes dorsales et la moitié des crêtes ventrales naissent dans cette région. Les
autres crêtes naissent donc principalement sur la face ventrale puisqu'au milieu du
638 BADREDDINE BEN SLIMANE & MARIE-CLAUDE DURETTE-DESSET
Fic. 2
Oswaldocruzia subauricularis (Rudolphi, 1819) chez Bufo sp. du Brésil. Male. A, extrémité
antérieure, vue latérale droite. B, téte, vue apicale. C-E, coupes transversales du corps. C, au
niveau du pore excréteur. D, id. détail de l'aile cervicale droite. E, au milieu du corps. F, détail
du cône génital, vue ventrale. G, bourse caudale, vue ventrale. H, I, spicule droit disséqué,
successivement vues ventrale et externo-latérale. Toutes les coupes sont orientées comme la
pure 27€. I: lame; f: fourche, s: sabot. A:téch° 50> im. BS DEE TEN snai
E: éch. 40 um. G: éch. 60 um.
OSWALDOCRUZIA DU BRÉSIL ET DE L’EQUATEUR 639
corps, le nombre de crétes dorsales est sensiblement équivalent à celui des ventrales.
Les crétes disparaissent en avant de la bourse caudale chez le male (fig. 3, I) et
progressivement en arrière de la vulve chez la femelle jusqu'au niveau des phasmides.
(fig. 3, k).
Chez le male originaire du Brésil, le nombre de crétes est de 33 (20 dorsales,
13 ventrales) au niveau du pore excréteur (fig. 1, C) de 41 (21 d, 20 v), au niveau de
la jonction oesophago-intestinale et de 55 (28 d, 27 v) au milieu du corps (fig. 2, E).
Chez les deux mâles originaires d'Equateur, le nombre de crétes est de 39 (23 d, 16 v)
et 40 (22 d, 18 v) au niveau du pore excréteur (fig. 3, C); de 40 (22 d, 18 v) et 43 (22
d, 21v) au niveau de la jonction oesophago-intestinale, de 47 (24 d, 23 v,) et SI (26 d,
25 v) au milieu du corps (fig. 3, G). Chez la femelle, le nombre de crétes est de 39 (23
dorsales, 16 ventrales) au niveau du pore excréteur (fig. 3, D), de 43 (25 d, 18 v) au
niveau de la jonction oesophago-intestinale, de 61 (30 d, 31 v), au milieu du corps
(fig. 3, F) et de 63 au niveau du vestibule (les cordons latéraux n'étant pas visibles à
ce niveau, le nombre exact de crétes dorsales et ventrales ne peut ètre donné).
Les crétes sont espacées de facon régulière mais, dans la région du pore
excréteur, elles sont plus serrées sur la face dorsale, puisque plus nombreuses.
Les crétes sont de taille équivalente sauf dans la région oesophagienne où les
deux crétes ventrales adjacentes aux cordons latéraux sont légèrement plus grandes que
les autres crétes et forment des ailes cervicales (fig. 2, D; 3, E).
A l'exception des ailes cervicales, légèrement orientées ventralement, les autres
crétes sont orientées perpendiculairement à la paroi du corps.
Male: (les mensurations concernant le mâle originaire du Brésil sont données
en premier, les extrêmes des mensurations concernant les 5 mâles originaires d'Équa-
teur sont données entre parenthèses). Longueur: 8200 (8450-9600). Largeur dans la
partie moyenne du corps: 170 (160-180). Vésicule céphalique haute de 85 (85-95) sur
40 (40-50) de large dans sa partie enflée et de 40 (40-45) dans sa partie fine. Anneau
nerveux, pore excréteur et deirides situés respectivement à 230 (185-220), 425 (380-
445) et 470 (420-495) de l'apex. Oesophage long de 520 (480-560). (fig. 2, A; 3, A).
Bourse caudale de type 2-3 à tendance 2-1-2 c'est-à-dire que les extrémités des
côtes 4 étant coudées vers l'avant elles sont plus rapprochées de celles des côtes 3 que
de celles des côtes 5 (fig. 2, G; 3, O). Côtes 2-3 d'une part, 5-6 de l'autre jointives.
Côtes 6,8 et dorsale de type III avec côtes 8 chevauchées par les côtes 6 sauf dans leur
tiers distal. Cote dorsale divisée distalement en 2 ou 3 rameaux, les rameaux externes
(côtes 9) se détachant avant la division de la côte dorsale. Absence de gubernaculum.
Cone génital haut de 15 (15-15) sur 17 (17-17) de large a sa base. Il porte sur sa lèvre
antérieure une papille zéro bien développée et deux minuscules papilles 7 sur sa lèvre
postérieure. (fig. 2, F; 3, 0). Spicules ailés, longs de 210 (190-205) a pointes com-
plexes, se divisant au tiers proximal de leur hauteur en trois branches: lame, fourche
et sabot. La lame se divise distalement en 8(7-7) rameaux; la fourche se divise en
deux branches de longueur équivalente à 20 (25-25) % de sa hauteur. La branche
externe est effilée, la branche interne est recourbée et enflée distalement. (fig. 2, H, I:
3 EIN):
640 BADREDDINE BEN SLIMANE & MARIE-CLAUDE DURETTE-DESSET
FIG. 3
Oswaldocruzia subauricularis (Rudolphi,1819) chez Bufo marinus d'Équateur. A, màle, extrémité
antérieure, vue ventrale. B, mâle, tête, vue apicale. C-G, coupes transversales du corps. C, mâle, au
niveau du pore excréteur. D, femelle, id. E, femelle, détail de l'aile cervicale droite au niveau du
pore excréteur. F, femelle, au milieu du corps. G, male, id. H, femelle, ovéjecteur, vue latérale droi-
te. I, male, bourse caudale, vue latérale gauche. J, male, extrémité antérieure, naissance des crétes
cuticulaires, vue ventrale. K, femelle, queue, vue latérale droite. L-N, male, spicule droit disséqué.
L, vue externo-latérale. M, vue ventrale. N, vue interne. O, male, bourse caudale, vue ventrale.
Toutes les coupes sont orientées comme la figure 3,D. 1: lame, f: fourche, s: sabot. A, I, K, O, éch.:
60 um. B, E, L, M, N, éch.: 50 um. C, D, F, G, éch.: 40 um. H, éch: 100 um. J, éch: 80 pm.
OSWALDOCRUZIA DU BRÉSIL ET DE L’EQUATEUR 641
Femelle: (les mensurations concernent 8 spécimens originaires d'Équateur; le
premier chiffre représente la moyenne, les chiffres entre parenthèses, les extrêmes):
Longueur: 16025 (14450 - 18000). Largeur: 211 (190-230) dans leur partie moyenne.
Vésicule céphalique haute de 104 (95-130) sur 52 (45-60) de large dans sa partie
enflée et de 47 (45-50) dans sa partie fine. Anneau nerveux, pore excréteur et deirides
situés respectivement à 244 (225-280), 459 (390-530) et 509 (420-580) de l'apex.
Oesophage long de 590 (500-640).
Didelphie. La vulve s'ouvre à 5362 (4450-6500) de la pointe caudale soit dans
le tiers postérieur du corps. Vagina vera long de 59 (50-75) divisant le vestibule long
de 396 (310-470) en deux parties équivalentes. Sphincters et trompes longs respec-
tivement de 49 (40-55) et de 36 (30-45). (fig. 3, H). Branche utérine antérieure longue
de 3297 (2860-3700), contenant 119 (85-140) oeufs; branche utérine postérieure lon-
gue de 3420 (2550-4050) contenant 122 (98-140) oeufs. Oeufs au stade morula, hauts
de 92,5 (85-100) sur 54 (50-60) de large. Queue longue de 196 (185-210) sur 86 (75-
100) de large au niveau de l'anus. Pointe caudale longue de 17,6 (15-20) (fig. 3, K).
Discussion: O. subauricularis a été très brièvement décrite par RUDOLPHI
(1819) chez "Rana musicae" au Brésil. La détermination de l'hôte parait eronnée
puisque dans la nomenclature actuelle cette espèce est synonyme de Bufo americanus
qui n'est connu que d'Amérique du Nord.
TRAVASSOS (1917) donne une description plus détaillée de l'espèce mais
toujours sans 1llustration. Il signale la présence de l'espèce chez différents Amphi-
biens qui, dans la nomenclature actuelle, correspondent aux taxa suivants: Cerato-
phrys cornuta, Leptodactylus ocellatus (= Cystignatus ocellatus) pour les Leptodacty-
lidae, Bufo americanus (= Rana musica), Bufo ceratophrys, Bufo marinus (= Bufo
agua), Bufo terrestris (=Bufo musica, dont la distribution est uniquement nord-améri-
caine) pour les Bufonidae. Il précise que sa description est basée sur le matériel ré-
colté chez Ceratophrys cornuta et Bufo agua.
TRAVASSOS (1921) illustre l'espèce chez Ceratophrys cornuta et Bufo agua et
reprend la description de 1917. D'après cette description, l'espèce est caractérisée par
une vésicule céphalique en deux parties, un pore excréteur situé juste en arrière de
l'annneau nerveux, une queue femelle tronquée avec une pointe fine, une vulve sans
lèvres saillantes, un ovéjecteur relativement court (460um) et une bourse caudale où
les côtes 8 naissent à la racine de la côte dorsale et sont recouvertes par les côtes 6 sur
les deux tiers antérieurs de leur trajet (type III). Aucune mention n'est faite du
synlophe.
TRAVASSOS (1937) ajoute à la liste d'hôtes, Bufo marinus, B. crucifer, Hyla
faber, H. mesophaea, et Phyllomedusa barmeisteri, tous du Brésil. Il redécrit l'espèce
chez Bufo crucifer et donne des illustrations aussi bien de la bourse caudale que des
spicules pour mettre en évidence la variabilité de l'anatomie de la côte dorsale et des
pointes spiculaires chez différents hôtes. Il précise qu'il existe des crêtes cuticulaires
espacées de 9 a 12 um et qu'il n'existe pas d'ailes cervicales.
642 BADREDDINE BEN SLIMANE & MARIE-CLAUDE DURETTE-DESSET
Le matériel original de l'Institut Oswaldo Cruz ne comporte pas d'Oswaldo-
cruzia de Ceratophrys cornuta ni de Bufo marinus (= Bufo agua). Il s'agit de 4
spécimens provenant d'Angra dos Reis, Estado do Rio: un male et une femelle n°
5038, parasites de Bufo crucifer, un male n° 5008, parasite d'Hyla faber et une
femelle n°4910 également parasite d'Hyla faber. Aucun des spécimens mâles ne
correspond aux dessins de TRAVASSOS (1921). Le spécimen parasite du Bufo n'a pas
une bourse caudale d'Oswaldocruzia; celui parasite d'Hyla a une bourse caudale de
type II. En ce qui concerne les femelles, celle parasite du Bufo a une vésicule cépha-
lique simple, celle parasite d’Hyla ne présente aucun élément contradictoire majeur
avec la description de Travassos mais nous savons que dans le genre Oswaldocruzia,
les femelles d'une région biogéographique déterminée sont morphologiquement très
proches les unes des autres et que seule la connaissance du synlophe oesophagien
permet de façon sûre de rapporter un spécimen femelle à une espèce déterminée. Dans
le cas de la femelle parasite d'Hyla, l'état du matériel ne nous a pas permis de la
déterminer.
En revanche, les spécimens que nous décrivons ci-dessus chez des Bufonidae,
nous paraissent pouvoir être identifiés au matériel décrit par TRAVASSOS (1917, 1921)
chez Ceratophrys cornuta et Bufo marinus (= Bufo agua) au Brésil. Tous les carac-
tères concordent à l'exception de la position du pore excréteur plus postérieure chez
nos spécimens mais nous savons par ailleurs (cf. BEN SLIMANE & DURETTE-DESSET,
1993) que ce caractère est dans ce genre très variable au sein de la même population,
a fortiori chez un hôte différent. L'absence d'ailes cervicales signalée par Travassos
peut s'expliquer par le fait que ces dernières sont minuscules et ne peuvent être obser-
vées qu'en coupe transversale du corps.
En tenant compte des critères actuels, O. subauricularis serait donc carac-
térisée par les éléments suivants:
— Ailes cervicales présentes mais minuscules et visibles seulement en coupe
transversale du corps. Synlophe avec crêtes cuticulaires pourvues d'un soutien
chitinoïde tout le long du corps. Nombreuses crêtes ventrales naissant en
arrière de la région oesophagienne.
— Disposition des côtes bursales 6, 8 et dorsale de type III.
— Vésicule céphalique en deux parties.
— Vulve dépourvue de lèvres saillantes.
— Spicules formés de trois branches principales: lame, fourche, sabot. Lame divi-
see à son extrémité distale en "peigne"; fourche divisée en-deçà de son tiers
postérieur en deux branches de longueur équivalente mais de forme différente.
La seule espèce néotropicale, sur les 9 espèces décrites, à posséder également
des côtes 6 et 8 de type III, est O. chambrieri Ben Slimane & Durette-Desset, 1993.
Cette espèce se distingue facilement par l'absence d'ailes cervicales, des crétes
dépourvues de soutien chitinoide et une vésicule céphalique simple.
Le spectre d' hòtes d'O. subauricularis tel que nous l'identifions, serait actu-
ellement réduit a Ceratophrys cornuta et Bufo marinus. En effet, la disposition des
côtes bursales 6,8 et 9 décrites par TRAVASSOS (1937), chez B. crucifer et Hyla faber
OSWALDOCRUZIA DU BRÉSIL ET DE L’EQUATEUR 643
est clairement de type II, c'est-à-dire que les côtes 8 naissent a la racine de la côte
dorsale mais ne sont recouvertes par les 6 que dans leur partie médiane. Bufo ameri-
canus (= Rana musica) et Bufo musicus (= Bufo terrestris) ne peuvent faire partie de
la liste d'hôtes puisqu'il s'agit d'espèces strictement néarctiques. En ce qui concerne
les parasites trouvés chez les autres hôtes par Travassos, nous manquons d'éléments
pour les rattacher à une espèce donnée. Enfin, en accord avec TRAVASSOS (1937),
nous pensons que les spécimens femelles parasites de Rana pipiens aux U.S.A et
identifiés comme ©. subauricularis par WALTON (1929), ne peuvent appartenir à cette
espèce car le vagina vera est d'un type très particulier.
Oswaldocruzia dlouhyi n.sp.
Matériel étudié: & holotype, £ allotype 715 MD a, 1 4,4 ® paratypes 715 MD b, 1 à,
1 2 MHNG 18769 INVE, coparasites d'O. subauricularis, parasites de l'intestin grêle d'un Bufo
sp., Gavioes (134 km au Nord de Rio de Janeiro), Brésil., 10.09.1985, leg. C. Dlouhy.
Description: Nématodes ne présentant pas d'enroulement. Vésicule céphalique
formée d'une partie antérieure enflée et d'une partie postérieure fine (fig. 4, B). Pore
excréteur toujours situé dans le tiers postérieur de l'oesophage. Deirides de forme
triangulaire, situées postérieurement au pore excréteur (fig. 4, D). Glandes excrétrices
très développées. Séparation oesophage musculaire et glandulaire très nette (fig. 4,
A). Présence d'ailes cervicales (fig. 4, F, G). Crétes cuticulaires dépourvues d'un sou-
tien chitinoide.
Téte: (fig. 4, C).Bouche triangulaire, arrondie aux angles. En vue apicale, pré-
sence de 6 papilles labiales externes dont les latérales sont accolées aux 2 amphides et
de 4 papilles céphaliques. Petite dent oesophagienne dorsale présente.
Synlophe: (étudié a différents niveaux chez | male et 1 femelle et au milieu du
corps chez un male et 3 femelles). Dans les deux sexes, corps parcouru longitu-
dinalement par des crêtes cuticulaires continues. Chez le mâle, les quatre cinquièmes
des crétes dorsales et les trois cinquièmes des crétes ventrales naissent dans la région
oesophagienne. Chez la femelle, presque toutes les crétes dorsales et les trois quart
des crétes ventrales naissent dans cette région. Au milieu du corps, le nombre de
crétes dorsales reste légèrement supérieur a celui des ventrales. Chez la femelle, il y a
naissance de crétes dorsales supplémentaires dans la région vestibulaire (fig. 4, J). Les
crêtes disparaissent en avant de la bourse caudale chez le mâle (fig. 4, N) et pro-
gressivement en arriere de la vulve chez la femelle jusqu'au niveau des phasmides
(fig. 4, P).
Chez le male, le nombre de crétes est de 30 (18 dorsales,12 ventrales) au
niveau du pore excréteur (fig. 4, F), de 31 (19 d,12 v) au niveau de la jonction oeso-
phago-intestinale et de 32 (17 d, 15 v) au milieu du corps (fig. 4, H). Chez le deu-
xième mâle, le nombre de crêtes est de 32 au milieu du corps dont 17 dorsales et 15
ventrales.
Chez la femelle, le nombre de crétes est de 34 (21 dorsales, 13 ventrales) au
niveau du pore excréteur, de 35 (21 d et 14 v), au niveau de la jonction oesophago-
644 BADREDDINE BEN SLIMANE & MARIE-CLAUDE DURETTE-DESSET
Fic. 4
Oswaldocruzia dlouhyi n.sp. A, male, extrémité antérieure, vue latérale droite. B, femelle, détail de
la vésicule céphalique, vue ventrale. C, femelle, téte, vue apicale. D, femelle, détail du pore excré-
teur et des deirides, vue ventrale. E, femelle, ovéjecteur, vue latérale gauche. F-J, coupes transver-
sales du corps. F, male, au niveau du pore excréteur. G, femelle, id. H, male, au milieu du corps. I,
femelle, id. J, femelle, au niveau du vestibule. K-L, male, spicule droit disséqué. K, vue ventrale. L,
vue externo-latérale. M, mâle, cône génital, vue ventrale. N-O mâle, bourse caudale. N, vue latérale
gauche. O, vue ventrale. P, femelle, queue, vue latérale droite. Toutes les coupes sont orientées
comme la figure 4,G. 1: lame, f: fourche, s: sabot. A, E, éch.: 50um. B, C, F-I, K, L, éch.: 30 um. D,
éch: 20 um, J, éch: 40 um, M, éch: 10 um, N-P, éch: 60 um
OSWALDOCRUZIA DU BRÉSIL ET DE L’EQUATEUR 645
intestinale, de 41 (22 d, 19 v) au milieu du corps (fig. 4, J) et de 47 au niveau du
vestibule. (Les cordons latéraux n'étant pas visibles à ce niveau, le nombre exact de
crétes dorsales et ventrales ne peut étre donné). Chez les trois autres femelles, le nom-
bre de crétes est de 29 (16 d, 13 v), 44 (24 d, 20 v) et 42 (23 d, 19 v) au milieu du corps.
Au milieu du corps, les crêtes sont espacées de façon régulière mais dans la
région oesophagienne les crétes latéro-ventrales sont plus espacées que les crétes
ventrales (fig. 4, F, G).
Les crétes sont de taille équivalente sauf dans la région oesophagienne où les
crétes ventrales présentent un léger gradient décroissant chez le male (fig. 4, F). Les
ailes cervicales sont formées d'une créte unique, de forme triangulaire et située
ventralement juste en arrière du cordon latéral. Les ailes naissent à la base de la vési-
cule céphalique et disparaissent a environ 300 en arrière de la jonction oesophago-
intestinale.
A l'exception des ailes cervicales, légèrement orientées ventralement, les autres
crêtes sont orientées perpendiculairement à la paroi du corps (fig. 4, F-I).
Male holotype: Longueur: 6450. Largeur dans la partie moyenne du corps:
120. Vésicule céphalique haute de 90 sur 55 de large dans sa partie enflée et 45 dans
sa partie fine. (fig. 4, B). Anneau nerveux, pore excréteur et deirides situés respec-
tivement à 235, 390 et 420 de l'apex. Oesophage long de 550 (fig. 4, A).
Bourse caudale de type 2-3 à tendance 2-1-2 c'est-à-dire que les extrémités des
côtes 4 étant coudées vers l'avant sont plus rapprochées de celles des côtes 3 que de
celles des côtes 5 (fig. 4, O). Côtes 2-3 d'une part, 5-6 de l'autre jointives. Côtes 6,8 et
dorsale de type II avec court tronc commun aux côtes 8 et 9; côtes 6 recouvrant les
côtes 8 dans leur partie médiane. Côte dorsale divisée distalement en 3 rameaux, les
rameaux externes (côtes 9) se détachant avant la division de la côte dorsale (fig. 4, 0).
Absence de gubernaculum. Cône génital haut de 25 sur 25 de large à sa base. Il porte
sur sa lèvre antérieure une papille zéro bien développée et deux minuscules papilles 7
sur sa lèvre postérieure (fig. 4, M). Spicules ailés, longs de 225 à pointes complexes, se
divisant au tiers proximal de leur hauteur en trois branches: lame, fourche et sabot.
Chez un paratype dont les spicules ont été disséqués, la lame se divise distalement en 9
rameaux; la fourche se divise en deux branches de longueur inégale à 27% de sa hau-
teur. La branche externe est nettement plus courte que la branche interne (fig. 4, K).
Femelle allotype: Longueur: 10200. Largeur: 200 dans sa partie moyenne.
Vésicule céphalique haute de 100 sur 60 de large dans sa partie enflée et 50 dans sa
partie fine. Anneau nerveux, pore excréteur et deirides situés respectivement à 265,
455 et 480 de l'apex. Oesophage long de 600.
Didelphie. La vulve s'ouvre à 3750 de la pointe caudale soit aux deux tiers
postérieurs du corps. Vagina vera long de 55 divisant le vestibule long de 420 en deux
parties inégales, l'antérieure étant légèrement plus longue. Sphincters et trompes longs
respectivement de 50 et 30 (fig. 4, E). Branche utérine antérieure longue de 2200,
contenant 64 oeufs, branche utérine postérieure longue de 2120 contenant 47 oeufs.
Oeufs au stade morula, hauts de 85 sur 50 de large. Queue longue de 220 sur 80 de
large au niveau de l'anus. Pointe caudale longue de 16 (fig. 4, P).
646 BADREDDINE BEN SLIMANE & MARIE-CLAUDE DURETTE-DESSET
Discussion: Comme la grande majorité des espèces d'Oswaldocruzia néotro-
picaux, les spécimens ci-dessus possèdent des spicules à 3 branches principales dont
la fourche est divisée dans son tiers postérieur et dont la lame se termine en "peigne"
c'est-à-dire que son extrémité distale est divisée en plusieurs rameaux.
Parmi ces espèces, seules O. touzeti parasite d'Eleutherodactylus variabilis et
O. vaucheri parasite d'Ischnocnema quixensis toutes deux décrites d'Équateur se
rapprochent de nos spécimens par la présence d'ailes cervicales et une disposition des
côtes bursales 6, 8 et 9 de type II.
O. dlouhyi n. sp. s'éloigne de O. touzeti par l'anatomie des ailes cervicales, des
crêtes oesophagiennes dépourvues de soutien chitinoïde et des côtes 4 plus longues
que les côtes 5. L'espèce la plus proche est O. vaucheri avec une forme identique des
ailes cervicales et des crétes cuticulaires dépourvues de soutien chitinoide tout le long
du corps. Les spécimens du Brésil se distinguent par une vésicule céphalique en deux
parties, des crêtes oesophagiennes plus nombreuses et plus marquées et par des côtes
4 aussi longues que les côtes 3.
Nous dédions cette nouvelle espèce à Carlo Dlouhy, Correspondant du
Muséum de Genève au Paraguay, qui a récolté le matériel.
Oswaldocruzia mazzai Travassos,1935
Matériel étudié: Un d, 1 2 MNHN 714 MD, 1 d, 1 2 MNHG 18770 INVE et 1 ©
MNHN 713 MD parasites de l'intestin gréle de deux Bufo marinus, San Pablo (Napo), Equa-
teur., 23.02.1987, leg. J.M. Touzet
Description: Nématodes ne présentant pas d'enroulement. Males au moins
deux fois plus petits que les femelles. Vésicule céphalique simple (fig. 5,A). Pore
excréteur situé environ aux deux tiers de la hauteur de l'oesophage. Deirides de forme
triangulaire, situées postérieurement au pore excréteur (fig. 5, A). Glandes excrétrices
très développées. Séparation oesophage musculaire et glandulaire très nette. Présence
de minuscules ailes cervicales, visibles seulement en coupe transversale du corps (fig.
5, E). Crétes cuticulaires pourvues d'un soutien chitinoïde.
Téte: (fig. 5, B). Bouche triangulaire, arrondie aux angles. En vue apicale, pré-
sence de 6 papilles labiales externes dont les latérales sont accolées aux 2 amphides et
de 4 papilles céphaliques. Petite dent oesophagienne dorsale présente.
Synlophe: (étudié chez 1 male et 2 femelles). Dans les deux sexes, corps par-
couru longitudinalement par des crêtes cuticulaires continues. Les quatre cinquièmes
des crétes dorsales et les deux tiers des crétes ventrales naissent dans la région oeso-
phagienne. Au milieu du corps, le nombre de crétes dorsales reste équivalent ou légè-
rement supérieur a celui des ventrales. Les crétes disparaissent en avant de la bourse
caudale chez le male (fig. 5, I) et progressivement en arrière de la vulve chez la
femelle jusqu'au niveau des phasmides (fig. 5, J).
Chez le male, le nombre de crétes est de 38 (22 dorsales, 16 ventrales) au
niveau du pore excréteur (fig. 5, C), 44 (23 d, 21 v) au niveau de la jonction
oesophago-intestinale et de 56 (29 d, 27 v) au milieu du corps (fig. 5, G). Chez les
OSWALDOCRUZIA DU BRÉSIL ET DE L’EQUATEUR 647
FIG. 5
Oswaldocruzia mazzai Travassos, 1935. A, male, extrémité antérieure, vue ventrale. B, femelle,
téte, vue apicale. C-G, coupes transversales du corps.C, male, au niveau du pore excréteur. D,
femelle, id. E, male, détail de l'aile cervicale gauche au niveau du pore excréteur. F, femelle, au
milieu du corps. G, male, id. H, femelle, ovéjecteur, vue latérale droite. I, male, bourse caudale,
vue latérale droite. J, femelle, queue, vue latérale droite. K-M, mâle, spicule gauche disséqué.
K, vue externo-latérale. L, vue interne. M, vue sub-ventrale. N, male, bourse caudale, vue
ventrale. Toutes les coupes sont orientées comme la figure 5,D. 1: lame, f: fourche, s: sabot. A,
celia 50 pm. B, E, éch::) 308mm: CD ME G} ech: 160 im; H, éch: 100) my; I N, ech: 70
um, J, éch: 80 um, K, L, M, éch: 40 um.
648 BADREDDINE BEN SLIMANE & MARIE-CLAUDE DURETTE-DESSET
deux femelles, le nombre de crétes est de 36 (21 d, 15 v) et 44 (25 d, 19 v) au niveau
du pore excréteur, 46 (25 d, 21v) et 49 (27 d, 22 v) au niveau de la jonction
oesophago-intestinale, 68 (34 d, 34 v) et 69 (36 d, 33 v) au milieu du corps (fig.5,F) et
de 67, 64 (33 d, 31 v) au niveau du vestibule.
Les crêtes sont espacées de façon régulière mais dans la région du pore
excréteur, elles sont plus serrées sur la face dorsale puisque plus nombreuses.
Les crétes sont de taille équivalente sauf dans la région oesophagienne où la
créte ventrale adjacente aux cordons latéraux est légèrement plus grande que les
autres crétes et forme une aile cervicale (fig. 5, E).
Toutes les crétes sont orientées perpendiculairement a la paroi du corps y
compris les ailes cervicales (fig. 5, C-G).
Male: (2 spécimens). Longueur: 7200-10650. Largeur dans la partie moyenne
du corps: 170-180. Vésicule céphalique haute de 65-85 sur 45-45 de large. Anneau
nerveux, pore excréteur et deirides situés respectivement à 175-200, 360-450 et 390-
480 de l'apex. Oesophage long de 540-600 (fig. 5, A).
Bourse caudale de type 2-3 a tendance 2-1-2, les extrémités des côtes 4 étant
coudées vers l'avant et plus rapprochées de celles des côtes 3 que de celles des côtes 5
(fig. 5, N). Côtes 2-3 d'une part, 5-6 de l'autre jointives. Côtes 6,8 et dorsale de type II
avec court tronc commun aux côtes 8 et 9 : côtes 6 recouvrant les côtes 8 dans leur
partie médiane. Cote dorsale divisée distalement en 3 rameaux, les rameaux externes
(côtes 9) se détachant avant la division de la côte dorsale (fig. 5, N). Absence de
gubernaculum. Cone génital haut de 20-20 sur 20-20 de large a sa base. Il porte sur sa
lèvre antérieure une papille zéro bien développée et deux minuscules papilles 7 sur sa
lèvre postérieure. Spicules ailés, longs de 210-210 a pointes complexes, se divisant au
tiers proximal de leur hauteur en trois branches: lame, fourche et sabot. La lame se
divise distalement en 10 rameaux; la fourche se divise en deux branches de longueur
équivalente à 23% de sa hauteur. La branche externe est effilée, la branche interne est
enflée distalement (fig. 5, M).
Femelle: (3 spécimens. Les mensurations entre parentheses correspondent a la
femelle 713 MD, ce qui permet de donner les mensurations réelles de chaque
spécimen étudié): Longueur: 19300-19350 (13800). Largeur: 260-260 (210) dans la
partie moyenne. du corps. Vésicule céphalique haute de 80-90 (95) sur 55-55 (50) de
large. Anneau nerveux, pore excréteur et deirides situés respectivement a 250-250
(255), 470-510 (470) et 570-580 (530) de l'apex. Oesophage long de 650-670 (620).
Didelphie. La vulve s'ouvre a 6000-6300 (4150) de la pointe caudale soit dans
le tiers postérieur du corps. Vagina vera long de 55-65 (55) divisant le vestibule long
de 500-520 (370) en deux parties inégales, la partie antérieure étant la plus longue.
Sphincters et trompes longs respectivement de 40-60 (45) et 40-45 (35) (fig. 5, H).
Branche utérine antérieure longue de 4150-4390 (2290) contenant 200-260 (85) oeufs,
branche utérine postérieure longue de 4430-4650 (1950) contenant 190-250 (70)
oeufs. Oeufs au stade morula, hauts de 85-85 (85) sur 50-50 (50) de large. Queue
longue de 180-280 (180) sur 110-110 (80) de large au niveau de l'anus. Pointe caudale
longue de 18 -18 (17) (fig. 5, J).
OSWALDOCRUZIA DU BRÉSIL ET DE L’EQUATEUR 649
FIG. 6
Oswaldocruzia taranchoni n.sp. A, male, extrémité antérieure, vue latérale droite. B, femelle,
vésicule céphalique, vue ventrale. C, femelle, téte, vue apicale. D, femelle, détail du pore excré-
teur et des deirides, vue ventrale. E, femelle, ovéjecteur, vue latérale droite. F-H, femelle, coupes
transversales du corps. F, au niveau de la jonction oesophago-intestinale. G, au milieu du corps.
H, au niveau du vestibule. I, male, naissance des crétes cuticulaires, vue latérale droite. J, femelle,
queue, vue latérale droite. K, id. naissance des crétes cuticulaires, vue ventrale. L, M, male,
spicule droit disséqué. L, vue ventrale. M, vue externo-latérale. N, O, male, bourse caudale. N,
vue latérale droite. O, vue ventrale. Toutes les coupes sont orientées comme la figure 6, F. |:
lame, f: fourche, s: sabot. A, E éch.: 60 um. B, D, F-H, éch.: 50 pm. C, L, M, éch: 30 um. I-K,
éch: 80 um. N, éch: 40 pm, O, éch: 20 um.
650 BADREDDINE BEN SLIMANE & MARIE-CLAUDE DURETTE-DESSET
Discussion: Parmi les espèces néotropicales, seuls O. mazzai Travassos (1935)
parasite de Bufonidae d'Argentine et O. bonsi Ben Slimane & Durette-Desset, 1993
parasite de Bolitoglossa equatoriana et d'Ischnonema quixensis en Équateur, pré-
sentent comme nos spécimens une disposition des côtes bursales 6, 8 et 9 de type II.
De plus, chez O. bonsi, le synlophe est dépourvu d'ailes cervicales et chez O. mazzai,
au moins in toto, les ailes cervicales paraissent absentes.
Les spécimens décrits plus haut s'éloignent de O. bonsi par des crétes cuti-
culaires pourvues de soutien chitinoide tout le long du corps, une bourse caudale avec
des côtes 2 et 3 d'une part, 5 et 6 de l'autre, jointives et par une lame spiculaire se
divisant en un nombre plus élevé de rameaux. Par contre il nous paraît possible de les
identifier a O. mazzai décrit pour la première fois par TRAVASSOS (1935) chez Bufo sp.
et redécrit par le méme auteur en 1937 chez B. marinus. Bien que nous n'ayons pas de
certitude absolue concernant les caractères du synlophe (en particulier la présence
d'un soutien chitinoide), la vésicule céphalique est simple, les bourses caudales sont
identiques et les branches de la fourche spiculaire ont la méme forme.
Par contre, O. mazzai sensu LENT et al., (1946) décrit chez B. paracnemis,
Leptodactylus ocellatus et L. bufonius du Paraguay nous paraît étre une autre espèce
qui pourrait étre rapprochée de O. dlouhyi par la présence d'une vésicule céphalique
composée de deux parties. Elle s'en distingue cependant par des còtes 8 relativement
courtes, par la division plus distale de la fourche spiculaire et par le fait que les deux
branches de cette fourche sont de longueur équivalente. Nous proposons le nom de
Oswaldocruzia proencai n.sp. (= O. mazzai sensu Lent, Freitas & Proenga, 1946, nec
Travassos, 1935) pour désigner les spécimens paraguayens.
Oswaldocruzia taranchoni n.sp.
Matériel examiné: d holotype, © allotype MNHN 317 Ua, 1 femelle paratype MNHN
317 U b, 2 9 9 MHNG 18771 INVE., parasites de l'intestin grêle d'un Bufo marinus, Pernam-
buco, Brésil, 3.07.1967, leg. J. C. Quentin.
Description: Nématodes ne présentant pas d'enroulement. Vésicule céphalique
simple (fig. 6, B). Pore excréteur situé au niveau de la région oesophago-intestinale.
Deirides de forme triangulaire, situées postérieurement au pore excréteur (fig. 6, D).
Glandes excrétrices tres développées. Séparation oesophage musculaire et glandulaire
très nette (fig. 6, A). Absence d'ailes cervicales. Crétes cuticulaires dépourvues d'un
soutien chitinoide.
Téte: (fig. 6, C). Bouche triangulaire, arrondie aux angles. En vue apicale,
présence de 6 papilles labiales externes dont les latérales sont accolées aux 2 am-
phides et de 4 papilles céphaliques. Petite dent oesophagienne dorsale présente.
Synlophe: (en coupe transversale, étudié chez 2 femelles). Dans les deux
sexes, corps parcouru longitudinalement par des crétes cuticulaires continues. Plus
des deux tiers des crétes dorsales et la moitié des crétes ventrales naissent dans la
région oesophagienne. Au milieu du corps, le nombre de crétes dorsales reste trés
légèrement supérieur a celui des ventrales. Les crétes disparaissent en avant de la
bourse caudale chez le male (fig. 6, N) et progressivement en arrière de la vulve chez
la femelle jusqu'au niveau des phasmides (fig. 6, J).
OSWALDOCRUZIA DU BRÉSIL ET DE L’EQUATEUR 651
Chez les deux femelles, le nombre de crétes est de 34 (17 dorsales, 17
ventrales), 29 (16 d, 13 v) au niveau du pore excréteur, de 43 (22 d, 21 v) 31 (18 d, 13
v) au niveau de la jonction oesophago-intestinale (fig. 6, F) de 75, (38 d, 37 v),53 (28
d, 25 v) au milieu du corps (fig.6,G) et de 59 (38 d, 26 v), 52 au niveau du vestibule
(fig. 6, H).
Les crêtes sont de taille équivalente, espacées de façon régulière et orientées
perpendiculairement à la paroi du corps (fig. 6, F-H).
Mâle holotype: Longueur: 6450. Largeur dans la partie moyenne du corps:
120. Vésicule céphalique haute de 80 sur 35 de large. Anneau nerveux, pore excréteur
et deirides situés respectivement à 180, 370 et 390 de l'apex. Oesophage long de 405
(fig. 6, A).
Bourse caudale de type 2-3 à tendance 2-1-2, les extrémités des côtes 4 étant
coudées vers l'avant et plus rapprochées de celles des côtes 3 que de celles des côtes 5
(fig. 6, O). Côtes 2-3 d'une part, 5-6 d'autre part jointives. Côtes 6,8 et dorsale de type
III avec côtes 8 chevauchées par les côtes 6 sauf dans leur tiers distal. Côte dorsale
divisée distalement en 3 rameaux, les rameaux externes (côtes 9) se détachant avant la
division de la côte dorsale (fig.6,0). Absence de gubernaculum. Cône génital haut de 20
sur 25 de large à sa base. Il porte sur sa lèvre antérieure une papille zéro bien déve-
loppée et deux minuscules papilles 7 sur sa lèvre postérieure (fig.6,0). Spicules ailés,
longs de 175 à pointes complexes, se divisant au tiers proximal de leur hauteur en trois
branches: lame, fourche et sabot. La lame est spatulée sans division distale; la fourche
se divise en deux branches de longueur équivalente à 18% de sa hauteur (fig. 6, L, M).
Femelle allotype: Longueur: 7350. Largeur: 100 dans sa partie moyenne. Vési-
cule céphalique haute de 90 sur 40 de large. Anneau nerveux, pore excréteur et
deirides situés respectivement à 180, 370 et 410 de l'apex. Oesophage long de 390.
Didelphie. La vulve s'ouvre à 2300 de la pointe caudale soit aux deux tiers
postérieurs du corps. Vagina vera long de 30 divisant le vestibule long de 240 en deux
parties équivalentes. Sphincters et trompes longs respectivement de 30 et 20 (fig. 6,
E). Branches utérines de même longueur soit 1030, contenant 10 oeufs pour la
branche antérieure, 14 pour la branche postérieure. Oeufs au stade morula, hauts de
85 sur 40 de large. Queue longue de 175 sur 50 de large au niveau de l'anus. Pointe
caudale longue de 20 (fig. 6, J).
Discussion: Ces spécimens se distinguent de tous les Oswaldocruzia néo-
tropicaux connus dont les spicules sont formés de trois branches, par une lame spicu-
laire ne se terminant pas en peigne”. Ils sont caractérisés par une vésicule céphalique
simple, l'absence d'ailes cervicales et par une bourse caudale dont les côtes bursales 6,
8 et 9 sont de type III. Seule O. chambrieri Ben Slimane et Durette-Desset, 1993,
décrite chez Bufo typhonius en Équateur présente l'ensemble de ces caractères mais
peut être différenciée des spécimens brésiliens non seulement par sa lame spiculaire
se terminant en "pinceau" mais également par la présence d'un tronc commun aux
côtes 4 à 8 et par un sabot spiculaire possédant une branche supplémentaire.
Nous rangeons donc les spécimens brésiliens dans une nouvelle espèce que
nous proposons de nommer Oswaldocruzia taranchoni n.sp. en hommage à Mr. Pierre
Taranchon, Associé au Muséum, pour l'aide précieuse qu'il nous apporte.
652 BADREDDINE BEN SLIMANE & MARIE-CLAUDE DURETTE-DESSET
CONCLUSION
A l'exception d'O. neghmei Puga, 1981, parasite de Leptodactylidae au Chili et
dont les spicules seraient divisés en 2 et non pas 3 branches, les 11 autres espèces
néotropicales actuellement connues possèdent la méme anatomie spiculaire que les
espèces holarctiques à savoir division du manche au tiers de sa hauteur proximale en
3 branches principales, lame (branche externo-latérale), sabot (branche interno-dor-
sale) et fourche (branche interno-ventrale). Mais les espèces néotropicales se diffé-
rencient immédiatement des espèces holarctiques par deux éléments: la hauteur plus
distale de division de la fourche, et par le fait que chez 10 espèces sur 11, la lame se
divise en de nombreux rameaux ce que nous avons nommé une division en "peigne".
Dans la zone néotropicale, nous retrouvons les trois principaux types d'arran-
gement des côtes 8 par rapport à la côte dorsale et aux côtes 6, types déjà décrits dans
la faune de l'Ancien Monde (DURETTE-DESSET et al., 1992):
O. brasiliensis, O. lopesi et O. neghmei appartiennent au type I qui peut étre
considéré comme le plus primitif puisque c'est celui rencontré chez la grande majorité
des Trichostrongles.
O. bonsi, O. dlouhyi, O. mazzai, O. proencai, O. touzeti et O. vaucheri appar-
tiennent au type II plus évolué que le précédent puisque le tronc commun aux côtes 8
et à la côte dorsale est très court et que les côtes 6 recouvrent où sont accolées aux
côtes 8 sur une partie de leur trajet.
O. chambrieri, O. taranchoni et O. subauricularis appartiennent au type III le
plus évolué puisqu'il n'existe plus de tronc commun entre les côtes 8 et la côte dorsale.
A l'intérieur de chaque groupe, les espèces sont morphologiquement très
proches entre elles. Elles diffèrent par la présence ou l'absence d'ailes cervicales,
l'anatomie de ces dernières (nombre de crétes formant l'aile, forme de l'aile, hauteur,
etc.), l'absence ou la présence de soutien chitinoïde des crêtes, la forme de la vésicule
céphalique, la présence ou l'absence de becs vulvaires, les variations qui peuvent
exister sur la lame et la fourche spiculaires.
La valeur des caractères spécifiques apparaît très différente de celle reconnue
chez les autres Trichostrongles. Ainsi les spicules sont caractéristiques d'une région
biogéographique déterminée et, à l'intérieur de cette zone varient peu d'une espèce à
l'autre. A l'opposé, certains caractéres comme la position du pore excréteur par
rapport a la longueur de l'oesophage ou le nombre de crétes cuticulaires au milieu du
corps présentent une très grande variabilité. Ceci joint à une absence de spécificité
étroite entre l'hôte et le parasite traduit probablement l'évolution très récente de ce
groupe.
REMERCIEMENTS
Les auteurs remercient tout particulièrement Carlo Dlouhy, Ascuncion, Para-
guay et Jean-Marc Touzet, Quito, Equateur qui ont récolté une partie de ce matériel
mis à notre disposition par le Dr. Claude Vaucher, Muséum d'histoire naturelle de
OSWALDOCRUZIA DU BRÉSIL ET DE L’EQUATEUR 653
Genève, que nous remercions également pour ses conseils. La nomenclature actuelle
des Amphibiens néotropicaux a été mise à jour par le Dr. Annemarie Ohler du
laboratoire des Reptiles et Amphibiens du Muséum national d'Histoire naturelle de
Paris. L'Institut Oswaldo Cruz à Rio de Janeiro à mis à notre disposition le matériel
original de O. subauricularis. Nous les en remercions très vivement.
BIBLIOGRAPHIE
BEN SLIMANE, B. & M.C. DURETTE-DESSET. 1993. Quatre nouvelles espèces du genre Oswaldo-
cruzia Travassos,1917 (Nematoda: Trichostrongyloidea) parasites d'Amphibiens d'Equa-
teur. Revue suisse Zool. 100: 113-136.
BEN SLIMANE, B., DURETTE-DESSET, M.C. & A.G. CHABAUD. 1993. Oswaldocruzia (Tricho-
strongyloidea) parasites d'Amphibiens des Collections du Muséum de Paris. Annis
Parasit. hum. comp. 68:88-100.
DURETTE-DESSET, M.-C., 1985. Trichostrongyloid nematodes and their Vertebrate hosts: re-
construction of the phylogeny of a parasitic group. Adv. Parasitol., 24: 239-306.
DURETTE-DESSET, M.-C. & A. G. CHABAUD. 1981. Nouvel essai de classification des Néma-
todes Trichostrongyloidea. Ann/s Parasit. hum. comp., 56: 297-312.
DURETTE-DESSET, M.-C. & A. G. CHABAUD. 1993. Nomenclature des Strongylida au-dessus du
groupe famille. Ann/s Parasit. hum. comp., 68: 111-112.
DURETTE-DESSET, M.-C., K. NASHER & B. BEN SLIMANE. 1992. Oswaldocruzia arabica n.sp.
(Nematoda, Trichostrongyloidea) parasite d'un Bufonidae de la péninsule arabique et
remarque sur des espèces proches. Bull. Mus. natl. Hist. nat., Paris, 4ème sér., 14,
section A: 693-703.
LENT, H. TEIXEIRA DE FREITAS, J.F. & M.C. PROENCA. 1946. Alguns helmintos de batraquios
colecionados no Paraguai. Mems Inst. Oswaldo Cruz. 44: 195-214.
PUGA, S.R. 1981. Oswaldocruzia negmei nov.sp. (Trichostrongylidae) un nuevo nematodo
parasito del anuro chileno Hylorina sylvatica (Leptodactylidae). Studies neotrop. Fauna
Envir. 16: 107-111.
RUDOLPHI, C.A. 1819. Entozoorum synopsis. Berolini, 81 1p.
TRAVASSOS, L. 1917. Trichostrongylinas brazileiras (Se nota previa). Oswaldocruzia n. gen.
Brazil medico, 31: 9.
TRAVASSOS, L. 1921. Contribution a la faune helminthologique du Brésil XIII. Essai mono-
graphique sur la faune des Trichostrongylidae Leiper, 1909. Mems Inst. Oswaldo Cruz.
1325-185:
TRAVASSOS, L. 1935. Alguns novos generos e especies de Trichostrongylidae. Revta Med.
Cirurg. Brasil. 43:345-361.
TRAVASSOS, L. 1937. Revisao da familia Trichostrongylidae Leiper, 1912. Monogr. Inst.
Oswaldo Cruz. 1: 512p.
WALTON, A.C. 1929. Studies on some nematodes of North american frogs. J. Parasitol. 15:
227-240.
REVUE SUISSE DE ZOOLOGIE, 102 (3) : 655-761; septembre 1995
Revision of the Afrotropical rove-beetles of the genus
Megarthrus (Coleoptera, Staphylinidae, Proteininae)!
Giulio CUCCODORO2 & Ivan LOBL3
2,3 Muséum d'histoire naturelle, Case postale 6434, CH-1211 Genève 6, Switzerland.
2 The Natural History Museum, Cromwell Road, London SW7 5BD, UK.
Revision of the Afrotropical rove-beetles of the genus Megarthrus (Cole-
optera, Staphylinidae, Proteininae). - Megarthrus species from Africa
South of Sahara are revised. In addition to 11 species previously described,
the following 27 new species are recognised: M. bantu, M. clarkei, M. domi-
nicae, M. falasha, M. horticola, M. hutu, M. magnicaudatus, M. mahnerti,
M. major, M. maniwaata, M. merabet, M. mukankundiyeorum, M. mwami,
M. nanus, M. negus, M. niloticus, M. panga, M. ras, M. rougemonti, M.
scotti, M. selenitus, M. spinosus, M. stylifer, M. twa, M. vanschuytbroecki,
M. watutsi and M. zulu. Lectotypes are designated for M. abessinus Bern-
hauer and M. africanus Eichelbaum. Descriptions are provided and diag-
nostic characters are figured for all species, except for M. kamerunensis
Bernhauer which is not represented in the collections. Keys to species are
given separately for males and females. Most of these species occurs in the
highlands and mountains of East Africa (27 spp.) and in Ethiopia (9 spp.).
Only one species has been found South of 11°S latitude. The genus appears
to be absent from Madagascar, the Mascarene Archipelago and Africa west
of Cameroon.
Key-words: Staphylinidae - Proteininae - Megarthrus - taxonomy - Africa.
INTRODUCTION
The Proteininae are one of the staphylinoid taxa with a distinct bipolar
distribution (NEWTON, 1985), though with several tropical members. With the
Neophoninae, Micropeplinae, Dasycerinae and Pselaphinae, it shares atrophied spi-
racles on the abdominal segments 4 to 6, which suggests that these groups may form a
monophyletic group. Neophoninae differ from the last three taxa and from the
! This paper forms part of the work by G. Cuccodoro towards the degree of Ph. D. at the
University of Geneva, funded in part by the Swiss National Science Foundation, project no. 31-
2331.91.
Manuscript accepted 20.03.1995.
656 GIULIO CUCCODORO & IVAN LOBL
Proteininae in the undivided male 9th tergite (THAYER, 1987). Thus the Proteininae are
important for the understanding of the relationships within the other, more derived
members of the omaliine group.
STEEL (1966) defined within the Proteininae the south temperate tribes Anepiini
and Nesoneini with 10 species in 8 genera, and the predominantly north temperate
Proteinini which go as far south as New Guinea and, in the New World, north
Argentina. Apart from keys to genera and a few regionally restricted revisions of
Proteinus Latreille and individual species descriptions (e.g. COIFFAIT, 1982; HAYASHI,
1988), no taxonomic work has been done recently on the Proteinini. The lack of
taxonomic information is particularly striking in Megarthrus which may be the most
diverse goup within the subfamily, in terms of species number, ecology and distri-
bution.
Some 80 species of Megarthrus are recognised to date, from which 11 have been
described from the Afrotropical region. The revision of unstudied material, however,
shows that the diversity in this region (38 species) is much greater than previously
estimated. The present paper diagnoses the Afrotropical species and provides keys and
illustrations for identification. Phylogenetic and biogeographical relationships are
briefly discussed.
MATERIALS AND METHODS
The present study is based exclusively on adults. Unless specified, all material
(776 specimens) mentioned in the text has been examined. For detailed examination,
specimens were dissected, cleared in 0.1 N potassium hydroxide and mounted in Eukit
or Canada balsam on acetate slides. Drawings were made using a drawing tube.
Detailed locality data are reproduced according to labels, except for elevations which
are given in m. Major administrative units are given in English.
The term frons, as used in the present study refers to the area anterior of the U-
shaped impression, the vertex to the area behind. Patches of sensilla on antennomeres 6
to 10 were detected by examining slide preparations. Abdominal sternites and tergites
are counted from the first morphological segment. Measurements and ratios are defined
as follows: length of specimens = interval from middle of anterior pronotal margin to
inner apical angle of elytron; width of specimens = maximum pronotal width; AL =
antennal length / pronotal length: EL = elytral sutural length / pronotal length; ET =
elytral sutural length / shortest interval between sutural margin and lateral edge of
elytron in dorsal view; EW = shortest interval between sutural margin and outer apical
angle of elytron in dorsal view / shortest interval between sutural margin and lateral
edge of elytron in dorsal view; EY = interval between posterior ocular margin and apex
of frons in dorsal view / interval between anterior and posterior ocular margins in dorsal
view; GT = posterior width of gula / median length of gula; GW = width of neck / pos-
terior width of gula; HW = maximum pronotal width / interval between posterior ocular
margins in dorsal view; ML = median metasternal length / median mesosternal length;
MP = length of segment 4 of maxillary palpus / length of segment 3 of maxillary
AFROTROPICAL MEGARTHRUS 657
palpus; PT = maximum pronotal width / pronotal length; SP = maximum width of
abdominal sternite 8 / width of the basal projection; TPF = interval between basal angle
and tip of medioapical projection of female abdominal tergite 8 / lateral length of
medioapical projection of female abdominal tergite 8. The absence of that projection is
indicated as “abs”.
Material examined is deposited in the following collections: BMNH = The
Natural History Museum, London; CNCI = Canadian National Collection of Insects,
Ottawa; FMNH = Field Museum of Natural History, Chicago; MHNG = Muséum
d'histoire naturelle, Geneva; MRAC = Musée Royal de l'Afrique Centrale, Tervuren;
SEMC = Snow Entomological Museum, University of Kansas, Lawrence; TMSA =
Transvaal Museum, Pretoria; ZMHB = Museum fiir Naturkunde der Humboldt-Uni-
versitàt, Berlin.
NATURAL HISTORY AND ECOLOGY
Little is known about the life history and biology of Megarthrus spp. The water
loading behaviour noted by Cuccoporo (1995) has been also observed in two
Afrotropical species M. horticola and M. spinosus. The Afrotropical members of
Megarthrus possess fully developed wings and are found in a wide range of habitats
(savannas, forests, in leaf litter, humus, and under stones). According to locality labels
they have been found in dung of various mammal species, in fungi and decaying
vegetational matter. They aiso have been collected in carrion, meat and faeces traps.
TAXONOMY
Within the Proteinini, adult members of the genus Megarthrus may be distin-
guished by the absence of a vertexal ocellus, the pronotum medianly impressed or
sulcate, and the lateral pronotal edges denticulate or crenulate. Compared to the so far
examined congeners from other regions, the Afrotropical Megarthrus appear quite
homogeneous. They share the following characters: mesal portion of frons with setae
orientated backwards; pronotal and elytral pubescence recumbent; abdominal pubes-
cence parallel, except for tergite 3 bearing short setae converging toward base; frons
with mesal portion slightly arcuate anteriorly and straight posteriorly in lateral view;
supra-ocular margin sinuate in dorsal view; occipital ridge indistinct; antennal scape not
flattened; pronotal disc weakly convex in frontal view, with mesal portion almost
straight in lateral view; hypomeral groove and median prosternal ridge absent; elytron
with discal swellings low, lateral edge finely carinate; metasternum with the femoral
line arcuate in middle; median apophysis of abdominal sternite 3 with apical portion
straight; male sternite 9 lacking subbasal protuberance.
The keying of Afrotropical Megarthrus is difficult without reference to sexual
characters. As several species are represented by are sex only, separate keys are
provided for males and females. At this stage in our investigations of Megarthrus no
attempt has been made to analyse phylogenetic relationships. Therefore, no species
658
GIULIO CUCCODORO & IVAN LOBL
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Distributional pattern of the Afrotropical species of Megarthrus. a: Ethiopian area (9 spp); b:
Equatorial area (28 spp); c: South-African area (1 sp). Scale square = 63'550 km?.
a
AFROTROPICAL MEGARTHRUS
659
g DA h can I N k N
2 7
N GA Rowan / \
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eni a a aed
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FIG. 2
Scutellum, a-f; Temple, g-k; Median processes of abdominal sternites 2-3 (left to right), I-n;
schematic. Megarthrus abessinus: a, g, 1; M. africanus: f, i, n; M. clarkei: c; M. congoensis: d, k:
M. falasha: b, h; M. gigas: m; M. mwami: e.
660 GIULIO CUCCODORO & IVAN LOBL
groups are defined and the species are listed alphabetically. However, some of the
species are linked by particularly noteworthy characters: 1) M. nanus and M. zulu
possess a protrochanteral ridge; 2) M. panga and M. stylifer have a large ventral process
on the male abdominal tergite 8; 3) M. africanus, M. basilewskyi, M.gigas, M. major,
M. mukandudiorum, M. selenitus, and M. spinosus lack a medioapical projection of the
female abdominal tergite 8 and adhesive setae on male protarsi. Among these species,
M. basilewskyi, M.gigas and M. major have a conspicuously projecting inner apical
angle of the elytra in the female; 4) M. mahnerti and M. monticola are linked by a
bilobed apical margin of the female sternite 8.
KEY TO MALES
(not included are M. apicicornis, M. major, M. scotti, M. selenitus, and M. spinosus in
which only females are known).
1 Apical margin of 6th abdominal sternite incised. . . . M. kamerunensis Bernhauer
- Apicalimarein of 6th-abdominal sternite truncate: ... . 2.2... ee 2
2 Eighth abdominal tergite bearing a ventral process projecting ventrally (Fig. 59f)
RO II ROLO PR CR e can ee 60 0000: 3
- Eighth abdominal tergite lacking a ventral process projecting ventrally....... 4
3 [ipiosfacdea gus broadi(FIS4S9 DETTE e PER M. stylifer sp. n.
_ iproraedeaeusnarowi(Bier49b)e 2.02 0 M. panga sp. n.
4 Protarsal'sesment I bearing adhesive setae (Fig. 30). "Eee er rer 8
— Protarsal'segment 1 lacking adhesive setae (Fig. 228)... sn eee 5
5 Aedeasusisymmetricali 22 ee M. africanus Eichelbaum
- Aédeagus'asymmetrical.s.. 25.521220 CO 6
6 Protibiavlackime peg-like setaen nn. 2... M. mukankundiyeorum sp. n.
~ Protibia bearing pes-like setae (Fig. 98). SE 7
7 Ventral wall of aedeagal median lobe notched on the left side (Fig. 9b)
5 EN ER O SIA III M. basilewskyi Fagel
_ Ventral wall of aedeagal median lobe not notched (Fig. 20c). ...M. gigas Fagel
8 Nedeagusrasy imme th Cale singe sa). PER ET LEE M. ovalis Cameron
- Aedeasus symmetrical. 7.1: dat nn SSR 9
9 Tip of aedeagus hook-shaped (Fig. 63d)........... M. vanschuytbroecki sp. n.
— IMplonaedeasusnmothook-shaped EP PE EEE EE 10
10 Protibia/bearing;peg-like'setae(Fig-3d)} 1.2... ERP EEE 11
- Rrotibialackine:pes-like setae 51552 2... 20 see see ee eee 2
1] Aedeagal median lobe, in ventral view, with apical portion slender, near
tip somewhat narrower than middle (Fig. 3a)......... M. abessinus Bernhauer
= Aedeagal median lobe, in ventral view, with apical portion wide at base
andicvenlystapenines (Rie. 2//c) ine ee eee M. magnicaudatus sp. n.
12 Dorsoapical sclerite of aedeagal median lobe projecting proximally of
level'ofparameres:(Fig. 16c). ... ..... 2.4. de see E 13
— Dorsoapical sclerite of aedeagal median lobe not projecting proximally
of leveliof parameres); (Fig.l 8b). nae 2222 eee eee eee 23
AFROTROPICAL MEGARTHRUS 661
Metatrochantenlackingipegikeisetaer rn ner M. rougemonti sp. n.
Metatrochantenibearingape slike setaeyemaree RR eect rete EEE ou 14
IMetanbiallackimnegpeg=MKeiSetA ROERO PEER ae 13
Metatibiaybearineipes-like, Setac PEN PRE Eee: 16
Ventral wall of aedeagal median lobe notched basally (Fig. 16c)
IRE DO I DATI ASSI RE EEE A M. dominicae sp. n.
Ventral wall of aedeagal median lobe not notched (Fig. 23b). M. horticola sp. n.
Mesotrochanteral peg-like setae arranged in a single row................. 17
Mesotrochanteral peg-like setae arranged in a double row or grouped in a field 19
hipiofsacdeazusitruncateobliguelyé Giles 29) aan sn one 18
ihiprotracdcacusmounded(Ric#4 0b) Meese eas ee M. mwami sp. n.
Nedeagusistraishtnearup(i 29e) = eee ae M. mahnerti sp. n.
Aedeagus inflexed dorsally near tip (Fig. 36d)......... M. monticola Cameron
Mipoteacdcagus:broad (Fig FI) RCE ee RE RARES ER OSE 20
Miprotacdeagusmarrowa(Eigad/ a) ERRE en Ae re nee ne DA
Metatibial peg-like setae partly arranged in a row (Fig. 45g). . M. niloticus sp. n.
All metatibial peg-like setae grouped in a field (Fig. 65e)..... M. watutsi sp. n.
Metatibial peg-like setae grouped in area exceeding half of tibial length
(Me 1410). ath Moses ee AME AEA ante oes M. congoensis Cameron
Metatibial peg-like setae grouped on area smaller than half of tibial
ngi (BEE eee ia Mes cat) eee or con tee are à à Ra ER
Internal sac of aedeagus with strongly sclerotized tooth-like structures.
TE a ee ae ee ee NT M. bantu sp. n.
Internal sac of aedeagus lacking strongly sclerotized tooth-like structures
GODITI IA ILA EE M. hutu sp. n.
Netatrochanterlackin s;pes-like;seta MB Re Be RR 24
Metatrochantenbeanne spes-like;setaen an PER RR O 27
Eighth abdominal sternite 4.5-7.0x as long as its median projection M. twa sp. n.
Eighth abdominal sternite 2.0-4.0x as long as its median projection......... 25
Mesotrochanteral peg-like setae arranged in a single row................. 26
Mesotrochanteral peg-like setae arranged in a double row. ....M. clarkei sp. n.
All metatibial peg-like setae arranged in a single row (Fig. 43d). M. negus sp. n.
Some metatibial peg-like setae grouped in a field........ M. simienensis Fagel
Mesotrochanteral peg-like setae arranged in a single row................. 28
Mesotrochanteral peg-like setae arranged in a double row or grouped in a field 31
Mesotibia broader in middle than near tip (Fig. 69c)..................... 30
Mesotibia slender, near tip about as broad as in middle (Fig. 67c). ......... 29
Apex of 8th abdominal tergite truncate (Fig. 51f). .............. M. ras sp. n.
Apex of 8th abdominal tergite pointed (Fig. 18e)............ M. falasha sp. n.
Aedeagal median lobe, in lateral view, with apical portion slender, near
tip somewhat narrower than in middle (Fig. 69b)............... M. zulu sp. n.
Aedeagal median lobe, in lateral view, with apical portion wide at base
andievenlyacape nino, (182420) ae 2 RP eee M. nanus sp. n.
GIULIO CUCCODORO & IVAN LOBL
Mesotrochanteral peg-like setae arranged in a double row. . . M. wittei Cameron
Mesotrochanteral peg-like setae grouped in a field. ..................... 32
Ventral outline of apical portion of aedeagal median lobe straight to
ventrallysrecurveditipi(E19234D) SSR SAS ee ee eee M. merabet sp. n.
Ventral outline of apical portion of aedeagal median lobe sinuate (Fig.
DD) A ede aaa dhe, AN a ENS M. maniwaata sp. n.
KEY TO FEMALES
(not included are M. africanus, M. kamerunensis, M. nanus, M. panga and M. ras in
which only males are known)
N |
|
œ |
Eighth abdominal tergite lacking medioapical projection (Fig 10f). ......... 2
Eighth abdominal tergite bearing a medioapical projection (Fig. 33g). ....... 7
Inner apical angle of elyton projecting conspicuously (Fig. 10b)............ 3
Inner apical angle of elytron not projecting (Fig. 30g)..................... 5
Mediodorsal suture of 9th abdominal sternite arcuate (Fig. 1 1b)
SE TER cones RE PRO MER Ce aire ME ANNEES M. basilewskyi Fagel
Mediodorsal suture of 9th abdominal sternite angulate (Fig. 22b). .......... 4
Eighth abdominal sternite 4.5-4.6x as wide as its median projection (Fig.
SD D cs a has I eae N I RE o M. major sp. n.
Eighth abdominal sternite 3.0-3.5x as wide as its median projection (Fig.
DAC) iret Ser geet, Be Seah ee tae, eae gt RIO ER M. gigas Fagel
Eighth abdominal sternite 2.2-2.3x as long as width of its median
PLOJECHOM(RISPIDO) SE Sac caste eee eases er RE M. selenitus sp.n.
Eighth abdominal sternite 1.7-1.9x as long as width of its median
projection: (Fig. 39)... 2... . en. Seen re N 6
Antenna about 2.2x as long as pronotum. ................. M. spinosus sp. n.
Antenna about 2.6x as long as pronotum......... M. mukankundiyeorum sp. n.
Apical margin of 8th abdominal sternite sinuate (Fig. 37b)................. 8
Apical margin of 8th abdominal sternite straight or arcuate (Fig. 4c)......... 9
Dorsobasal edge of coxites V-shaped (Fig. 30c). .......... M. mahnerti sp. n.
Dorsobasal edge of coxites U-shaped (Fig. 37f)........ M. monticola Cameron
Eighth abdominal tergite 1.8-2.5x as long as its medioapical projection. ..... 10
Eighth abdominal tergite 3.2-10.0x as long as its medioapical projection. . . . . 11
Eighth abdominal tergite about 2.5x as long as its medioapical projection
CPS) PE EIN dre Done ko Io M. abessinus Bernhauer
Eighth abdominal tergite about 1.8 as long as its medioapical projection
Mie SD) SIT RER LEE Al Ria misent M. magnicaudatus sp. n.
Antennomere 4 strongly asymmetrical (Fig. 48d). ...............---.---- 12
Antennomere 4 symmetrical or slightly asymmetrical (Fig. 44f)........... 14
Protrochanter withtaimansverse midges, erase eee M. zulu sp.n.
Protrochanter without a transverse ridges a. RR 18
Elytron deeply depressed along lateral edge.............. M. ovalis Cameron
Blytrontlatalong;lateraliedee=. TERRE M. twa sp. n.
AFROTROPICAL MEGARTHRUS 663
Eighth abdominal tergite 10.0x as long as its medioapical projection........ 15
Eighth abdominal tergite 3.2-7.0x as long as its medioapical projection. ..... 11
Pronotum flat along apical portion of lateral edge. .......... M. merabet sp. n.
Bronotumidepressedhlonsienture lateraledee pre em euren 16
Dorsobasal edge of coxites V-shaped (Fig. 19e)............. M. falasha sp. n.
Dorsobasal edge of coxites U-shaped (Fig. 68b)........... M. wittei Cameron
leadmotwidened behindsthereyes (Rigs 2h) Eee 18
Keadswidenedibehindithegeyesiliis AR) eee SEE 26
Bodyal#8-2 OMRON ANS AREA Er oe Bok M. dominicae sp. n.
BINOMIO Lit AI rel 19
Eye, in frontal view, with highest point reaching the level of the vertex. . .... 20
Eye, in frontal view, with highest point below the level of the vertex........ 22
hiprotscutellum:pomtedi(Eiete2 b) mr ee. SR M. horticola sp. n.
Uiprosscutellumirounded' (BOTTE ORO ere 21
Dorsobasal edge of coxites V-shaped (Fig. 41c)............. M. mwami sp. n.
Dorsobasal edge of coxites transverse, except for median portion pointed
(LEIS SISA ac RI AI en Nong RA M. maniwaata sp. n.
Scutellanip rounded (RIE 2) TERE ee TRA ONERI EE 23
Scucellantipspomtedi (ie a) EEE AE E ARR IPO EA RE oa 24
Dorsobasal edge of coxites V-shaped (Fig. 54d). ............. M. scotti sp. n.
Dorsobasal edge of coxites transverse (Fig. 13d)............. M. clarkei sp. n.
Antennomere 5 shorter than antennomere 4 (Fig. 52c)..... M. rougemonti sp. n.
Antennomere 5 as long as antennomere 4 (Fig. 44f). .................... DS
Blyton Oley aslometas: wide aces nm ee M. negus sp. n.
Bilytonyles=2iOxeasslongeas wider. ae Set ARE M. simienensis Fagel
Medianametastemalird ge conspicuous... RIE ae ae ene 27
Medianimetastemal ridge tine or absent: tt 22. wen RE TT PERRET 28
Eighth abdominal tergite with medioapical projection longer than wide
GROG) QE A UT ERA wh craked bol git ERI SRC ARE ons M. apicicornis Cameron
Eighth abdominal tergite with medioapical projection wider than long
(BIONOLE DRE APRES RAA TE MANS ARRET M. vanschuytbroecki sp. n.
Iipjoßsthtabdommalitersitespomted( Ei So) a. ERRE 29
Tip’of 8th abdominal tergite rounded (Fig: 462). ............2.......... 31
Eighth abdominal tergite, in lateral view, with apical projection of flat
(VERSION is i suoi Lies M. bantu sp. n.
Eighth abdominal tergite, in lateral view, with apical projection raised,
MODE (1G 32 60) 863 RL ETUI EEE at 30
Eighth abdominal tergite, in lateral view, with apical projection slightly raised
Wi ZOU) ay re Rie heen BL Pe. coso ppi ri hei M. hutu sp. n.
Eighth abdominal tergite, in lateral view, with apical projection strongly
RUSSE SID) En seperate cheese oa M. congoensis Cameron
Dorsobasal edge of coxites V-shaped (Fig. 66a)............. M. watutsi sp. n.
Dorsobasall edge of coxites U-shaped (Fic: 46c).. 7. 42 an. see 32
Eighth abdominal tergite 5.0x as long as its medioapical projection (Fig.
ALS) Oe Pen Rene ES EEN DE M. niloticus sp. n.
Eighth abdominal tergite 4.0x as long as its medioapical projection (Fig.
CUOIO ENI e LARIO ni M. stylifer sp. n.
664 GIULIO CUCCODORO & IVAN LOBL
Megarthrus abessinus Bernhauer (Figs 2a, g, |, 3a-1, 4a-g)
Megarthrus abessinus BERNHAUER, 1931: 566.
Megarthrus abessinicus; BERNHAUER, 1942: 349 (incorrect spelling).
Type material. Lectotype 9: Ethiopia, Djem-Djem Forest (= Shewa prov., Jem Jem?),
c.2700m, 24.1x.1926 (H. Scott) BMNH, by present designation.
Additional material (18): Ethiopia, Kefa prov., Foja, Mt. Bor, 2950m, i1.1971 (R. O. S.
Clarke) 18 in BMNH; Gonder (= Simén) prov., Arghine, c. 3500m or higher, 24.xi.1952 (H.
Scott) ex roots of tufted grass in ravine, | 2 in BMNH (mislabelled paratype of M. simienensis);
Lori, c. 3500m or higher, 27.x1.1952 (H. Scott) in precincts of church, ex tall yellow composite
(Senecio myriocephalus), 19 in BMNH; near Mindigabsa, c. 3500m, 29.xii.1952 (H. Scott) 7d,
19 in BMNH (mislabelled paratypes of M. simienensis); Shewa prov., Addis Abeba, 1971 (G. de
Rougemont) 42 in BMNH and 16,29 in MHNG.
Distribution. Ethiopia.
Description. Length 1.55-1.65 mm; width 1.05-1.15 mm. Head and metasternum
dark brown; antenna, pronotum, elytron and abdomen yellow-brown or reddish-brown,
except for antennomeres 1-4 paler and sutural margin of elytron darkened; mouth parts
and legs paler than metasternum. Dorsal pubescence fairly uniform, on abdomen
shorter, on humeral area of elytron denser. Metasternal setae becoming sparser
medianly, shorter than prosternal setae. Pubescence of abdominal sternites 4-7 uniform,
except for a pair of long setae near medioapical margins. Puncturation fine on anterior
portion of hypomeron; medioposterior portion of metasternum impunctate. Frons on
level with vertex. Anterior frontal edge not carinate, evenly convex. Entire frontal
impression shallow or indistinct. Eye moderatly convex, with highest point below level
of vertex. Temple as in Fig. 2g. Submentum flat. Antenna (Fig. 4g) without patches of
sensilla; antennomeres 3 and 4 slightly asymmetrical. Pronotal disc (Fig. 4a) flat along
lateral edge, shallowly depressed along median groove, latter shallow, parallel-sided.
Anterior prosternal margin bordered by an irregular row of fine longitudinal ridges.
Protrochanter without transverse ridge. Lateral portion of prepectal ridge straight.
Scutellum as in Fig. 2a. Elytron not narrowed at base; base gradually inclined,
overhanging. Elytral disc flat along lateral edge, latter straight in dorsal view; apical
margin convex near suture; inner apical angle obtuse. Median metasternal ridge fine,
low. Abdominal tergite 3 slightly vaulted transversally. Sternites 2 and 3 with median
processes as in Fig. 21. Sternite 4 flat at base, then slightly vaulted transversally.
Ratios: AL 2.5; EL 15; ETE1.6557 EW 12EY62:8=2:9> Gi 2 OIG Wales SA
DOME A MPA Pil 2:0: SPS 2-3 4 PES"
d. Protarsal segment | bearing adhesive setae (Fig. 3d). Mesofemur (Fig. 3g) as
long as metafemur (Fig. 3h). Mesotibia (Fig. 3e) shorter than metatibia (Fig. 3f).
Protibia (Fig. 3d) with one or two peg-like setae. Peg-like setae on mesotrochanter (Fig.
3g) and mesotibia arranged in a single row, on metatibia arranged in a double row, and
grouped in a field on metatrochanter (Fig. 3h). Apex of abdominal tergite 8 as in Fig.
31. Sternite 8 as in Fig. 3c. Aedeagus as in Fig. 3a, b.
2. Abdominal tergite 8 (Fig. 4e, f) with medioapical projection. Sternite 8 as in
Fig. 4c. Genital segment as in Fig. 4b, d.
Comments. The remaining paralectotypes examined are not conspecific; see
under M. falasha and M. rougemonti. This species may be easily distinguished by the
AFROTROPICAL MEGARTHRUS 665
coloration, the elytral base not narrowed and the sexual characters. It resembles M.
magnicaudatus, but has the number of protibial peg-like setae in male strongly reduced
and a much shorter apical projection of the abdominal tergite 8 in female.
Megarthrus africanus Eichelbaum (Figs 2f, 1, n, Sa-h)
Megarthrus africanus EICHELBAUM, 1913: 114.
Type material. Lectotype d: Tanzania, Tanga distr., E Usambara Range, Mt. Bomole,
Amani, 11.x.1903 (F. Eichelbaum) ZMHB, by present designation.
Distribution. Tanzania: Usambara Range.
Description. Length 1.45 mm; width 1.0 mm. Body predominantly red-brown,
with darkened head and sutural margin of elytron, and paler legs, mouth parts and
antennomere 11. Dorsal pubescence fairly uniform, on abdomen shorter, sparser near
lateral pronotal edge, and denser along sutural margin and on humeral areal. Meta-
sternal setae becoming denser medianly, about as long as prosternal setae. Pubescence
on abdominal sternites 4-7 becoming denser medianly. Puncturation coarse on anterior
portion of hypomeron and on medioposterior portion of metasternum. Frons on level
with or raised above level of vertex. Anterior frontal edge finely and evenly carinate,
weakly convex in middle and oblique laterally. Entire frontal impression deep. Eye
almost hemispherical, with highest point reaching level of vertex. Temple as in Fig. 21.
Submentum flat. Antenna (Fig. 5c) without patches of sensilla; antennomeres 3 and 4
symmetrical. Pronotal disc (Fig. 5a) shallowly depressed along entire lateral edge and
flat along median groove; latter deep, parallel-sided. Anterior prosternal margin
bordered by a regular row of conspicuous longitudinal ridges. Protrochanter without
transverse ridge. Lateral portion of prepectal ridge sinuate. Scutellum as in Fig. 2f.
Elytron not narrowed at base; base abruptly inclined, overhanging. Elytral disc flat
along lateral edge; latter straight in dorsal view; apical margin convex near suture; inner
apical angle obtuse. Metasternal median ridge absent. Abdominal tergite 3 strongly
vaulted transversally. Abdominal sternites 2 and 3 with processes as in Fig. 2n. Sternite
4 flat at base, then strongly vaulted transversally.
Ratios: AL 2.6; EL 1.5; ET 1.8; EW 1.15; EY 2.85; GT 2.25; GW 1.8; HW 1.6-
leavin MIP 2.0: Pl 129: SP. 3:3.
3. Protarsal segment 1 lacking adhesive setae. Mesofemur as long as meta-
femur. Mesotibia (Fig. 5b) shorter than metatibia. Peg-like setae grouped in a field on
mesotrochanter (Fig. 5h) and mesotibia, absent from protibia, metatrochanter and
metatibia. Apex of abdominal tergite 8 as in Fig. 5g. Sternite 8 as in Fig. Sd. Aedeagus
asim Fig, Se, fi
2. Unknown.
Comments. Although the shape of the anterior portion of the frons is probably a
male sexual character, in the absence of females it is listed among the general
characters. This species share with M. mukankundiyeorum, M. selenitus and M. spi-
nosus a regularly ridged prosternal margin. It is characterised by the presence of a
symmetrical aedeagus in combination with the absence of protarsal adhesive setae.
666 GIULIO CUCCODORO & IVAN LOBL
Megarthrus apicicornis Cameron (Fig. 6a-g)
Megarthrus apicornis; CAMERON, 1950: 5 (incorrect spelling).
Megarthrus apicicornis CAMERON, 1950: 6.
Type material. Holotype 2: Zaire, Kivu prov. (Albert NP) V. Mikeno, near Rweru,
2400m, 3.vii.1934 (G. F. de Witte) ex bamboo, #469, MRAC.
Distribution. Zaire: V. Mikeno.
Description. Length 1.55 mm; width 1.0 mm. Body uniformly red-brown or
dark brown, except for paler legs, mouth parts and antennomeres 10 and 11, and some-
what darkened head and sutural margin of elytron. Dorsal pubescence fairly uniform,
elytral sparser on disc and denser on humeral area. Metasternal setae becoming sparser
medioposteriorly and longer anteriorly, shorter than prosternal setae. Pubescence of
abdominal sternites 4-7 uniform, except for a pair of long setae near medioapical
margins. Puncturation fine on anterior portion of hypomeron; median area of meta-
sternum impunctate. Frons raised above level of vertex. Anterior frontal edge not
carinate, strongly arcuate in middle, oblique laterally. Frontal impression indistinct in
middle and shallow laterally. Eye moderatly convex, with highest point below level of
vertex. Temple similar to that in Fig. 2k. Submentum weekly convex. Antenna (Fig. 6c)
without patches of sensilla; antennomeres 3 and 4 slightly asymmetrical. Pronotal disc
(Fig. 6f) shallowly depressed along basal portion of lateral edge and deeply depressed
along median groove; latter deep, parallel-sided. Anterior prosternal margin not
bordered by longitudinal ridges. Protrochanter without transverse ridge. Lateral portion
of prepectal ridge straight. Scutellum similar to that in Fig. 2d. Elytron not narrowed at
base; base gradually inclined, overhanging. Elytral disc flat along lateral edge; latter
straight in dorsal view; apical margin slightly sinuate near suture; inner apical angle
right-angled. Metasternal median ridge large, conspicuous. Abdominal tergite 3 slightly
vaulted transversally. Sternites 2 and 3 with median processes similar to that in Fig. 21.
Sternite 4 flat at base, then slightly vaulted transversally.
Ratios: AL 2.1; EL 1.5; ET 1.7; EW 1.2; EY 2.8-2.9; GT 2.0; GW 2.0; HW 1.6-
CAMES MP PN 18 SPE 7: TPE 0;
2. Medioapical projection of abdominal tergite 8 as in Fig. 6e, g. Sternite 8 as in
Fig. 6d. Genital segment as in Fig. 6a, b.
3. Unknown.
Comments. Nine species (M. apicicornis, M. bantu, M. congoensis, M. hutu, M.
niloticus, M. panga, M. stylifer, M. vanschuytbroecki and M. watutsi) possess a head
distinctly widened behind the eyes. Among these species, M. apicicornis is distin-
guished by its conspicuous metasternal median ridge and elongate, narrow apical
projection of abdominal tergite 8. The former character is shared with M. vanschuyt-
broecki, which has the tergal projection much wider.
Megarthrus bantu sp. n. (Figs 7a-i, 8a-g)
Type material. Holotype d: Zaire, Ruwenzori Range (Albert NP) Kalonge, river
Nyamwamba, tributary of Butahu, 2010m, 2-3.11.1953 (P. Vanschuytbroeck & J. Kekenbosch)
#2214-21, MRAC.
AFROTROPICAL MEGARTHRUS 667
Paratypes (85): same data as holotype, 55, 59 in MHNG and 294, 192 in MRAC;
Zaire, Ruwenzori Range (Albert NP) Kalonge, 2210m, 1.1x.1952 (P. Vanschuytbroeck & J.
Kekenbosch) #846-51, 16 in MRAC; same data, but 7.viii.1952, ex humus in forest, #695, 19 in
MRAC; Kalonge (Albert NP) Nyamwamba-Ihongero, 2480m, 25-29.v111.1952 (P. Vanschuyt-
broeck & J. Kekenbosch) ex humus in bamboo, #860-63, 16 in MRAC; same data, but 26-
28.viii.1952, #874, 16 in MRAC; Kalonge (Albert NP) river Katauleko, tributary of Butahu,
2180m, 1.vii.1952 (P. Vanschuytbroeck & J. Kekenbosch) ex humus, #615, 16, 19 in MRAC;
same data, but 1-2.viii.1952, #665-67, 16 in MRAC; Kalonge (Albert NP) river Katsambu, tri-
butary of Butahu, 2000m, 26.1-19.11.1953 (P. Vanschuytbroeck & J. Kekenbosch) #2155-2200,
19 in MRAC; Kalonge (Albert NP) river Kiondyo ya Kwnanza, tributary of Butahu, 2130m,
2.viii.1952 (P. Vanschuytbroeck & J. Kekenbosch) ex humus, #634-35, 16, 19 in MHNG and
1d, 22 in MRAC; same data, but 5.viii.1952, under bark, #657, 12 in MRAC; Ruwenzori
Range (Albert NP) Kyandolire, river Mulaku, tributary of Kakalari, 1750m, 14.x.1952 (P. Van-
schuytbroeck & J. Kekenbosch) ex humus, #1241-43, 16,29 in MHNG and 3d, 42 in MRAC;
Kivu, 39km S Lubero, Mombassa, 25.viii.1932 (L. Burgeon) R. Det. L2556, 16 in MRAC; Bu-
rundi, Bururi terr., Nyamurenbe, 900m, 7.111.1953 (P. Basilewsky) 16 in MRAC; Uganda, Ru-
wenzori Range, Toro prov., Mahoma River, 2700m, 13-16.viii.1952 (D. S. Fletcher) 1d in
BMNH.
Distribution. Burundi, Uganda and Zaire.
Description. Similar to M. congoensis from which it may be distinguished as
follows: Length 1.3-1.6 mm; width 0.85-1.05 mm. Antenna reddish-brown, antenno-
mere 11 paler. Eye strongly convex. Antenna as in Fig. 8c. Pronotum as in Fig. 8f.
Elytral disc shallowly depressed along lateral edge; apical contour somewhat convex
near suture; inner apical angle obtuse. Metasternal median ridge fine, low. Abdominal
tergite 3 almost flat. Ratios: AL 2.0-2.2; EL 1.5; ET 1.65-1.80; EY 2.8-2.9; GW 1.8;
ML 1.4; SP 2.5-2.9; TPF 5.0-6.8.
d. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 7g) longer than
metafemur (Fig. 7h). Mesotibia (Fig. 7c) shorter than metatibia (Fig. 7d). Peg-like setae
arranged in a double row on mesotrochanter (Fig. 7g), grouped in a field on mesotibia,
metatrochanter (Fig. 7h) and metatibia, absent from protibia. Apex of abdominal tergite
8 as in Fig. 7e, i. Aedeagus as in Fig. 7a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 8e, g. Sternite 8 as in
Fig. 8d. Genital segment as in Fig. 8a, b.
Comments. Megarthrus bantu differs from other species with the head widened
behind the eyes (see discussion under M. apicicornis) by the strongly sclerotised tooth-
like structures of the aedeagal internal sac and by the contour of the female abdominal
tergite 8.
Megarthrus basilewskyi Fagel (Figs 9a-g, 10a-g, 11a-e)
Megarthrus basilewskyi FAGEL, 1957: 27.
Type material. Holotype d: Rwanda, Kibuye terr., Yanina, 2300m, 12.11.1953 (P.
Basilewsky) MRAC.
Paratypes (49): Zaire, Kivu prov., Mwenga terr., Luiko, 2050m, 21.1.1952 (N. Leleup)
ex humus in montane forest, 1 in MHNG and 3 in MRAC.
Distribution. Rwanda; Zaire: Kivu.
Description. Similar to M. gigas from which it may be distinguished as follows:
Length 1.9 mm; width 1.4 mm. Body red-brown. Metasternal median ridge absent.
Ratios: EL 1.6; ET 1.8; EY 2.5; GT 2.0; GW 2.0; TPF abs.
668 GIULIO CUCCODORO & IVAN LOBL
d. Inner apical angle of elytron right-angled. Protarsal segment | lacking adhe-
sive setae. Mesofemur (Fig. 9c) shorter than metafemur (Fig. 9a). Mesotibia (Fig. 9f)
shorter than metatibia (Fig. 9e). Peg-like setae arranged in a single row on protibia (Fig.
9g) and mesotrochanter (Fig. 9c); grouped in a field on mesotibia, metatrochanter (Fig.
9a) and metatibia. Abdominal tergite 8 as in Fig 10d, e. Sternite 8 as in Fig. Ile.
Aedeagus as in Fig. 9b, d.
9. Apical contour of elytron as in Fig. 10b. Abdominal tergite 8 (Fig. 10f, g)
without medioapical projection. Sternite 8 as in Fig. 11d. Genital segment as in Fig.
11a-c; mediodorsal suture of sternite 9 arcuate.
Comments. Megarthrus basilewskyi, M. dominicae, M. gigas and M. major may
be distinguished by their large size. Megarthrus basilewskyi is characterised by the
shape of the aedeagus and, in the female, the tergite 8 lacking apical projection in
combination with the presence of the arcuate mediodorsal suture of the sternite 9.
Megarthrus clarkei sp. n. (Figs 2c, 12a-h, 13a-g)
Type material. Holotype ¢: Ethiopia, Shewa prov., Addis Abeba, 1971 (G. de Rouge-
mont) BMNH.
Paratypes (10): same data as holotype, 26,39 in BMNH and 2d, 29 in MHNG; Shewa
prov., Managasha Forest, c. 2900m, 9.1.1971 (R. O. S. Clarke) ex silt of dry stream bed, 19 in
BMNH.
Distribution. Central Ethiopia.
Description. Length 1.45-1.60 mm; width 1.0-1.1 mm. Body red-brown or dark
brown, with darkened head, metasternum and abdomen, and paler legs, mouth parts and
antennomeres 1-4. Dorsal pubescence fairly uniform, becoming denser along median
pronotal groove and near apical margin of abdominal tergite 7. Metasternal setae
becoming sparser on median area; shorter than those on prosternum. Pubescence of
abdominal sternites 4-7 uniform, except for a pair of long setae near medioapical
margins. Puncturation fine on anterior portion of hypomeron; median area of
metasternum impunctate. Frons raised above level of vertex. Anterior frontal edge not
carinate, evenly arcuate. Entire frontal impression shallow. Eye strongly convex, with
highest point below level of vertex. Temple similar to that in Fig. 2g. Submentum
weakly convex. Antenna (Fig. 13b) without patches of sensilla; antennomeres 3 and 4
slightly asymmetrical. Pronotal disc (Fig. 13a) shallowly depressed along entire lateral
edge and along median groove; latter usually deep, parallel-sided. Anterior prosternal
margin not bordered by longitudinal ridges. Protrochanter without transverse ridge.
Lateral portion of prepectal ridge straight. Scutellum as in Fig. 2c. Elytron abruptly
narrowed at base; base gradually inclined, overhanging. Elytral disc shallowly
depressed along lateral edge; latter slightly convex in dorsal view; apical margin
somewhat sinuate near suture; inner apical angle obtuse. Metasternal median ridge fine,
low. Abdominal tergite 3 almost flat. Sternites 2 and 3 with median processes similar to
that in Fig. 21. Sternite 4 flat at base, then slightly vaulted transversally.
Ratios: AL 2.0-2.1; EL 1.4; ET 1.53 BW 122: EY 2:8-2.9: |G 2 25GWa les EMI
1.6-1.7; ML 1.4; MP 1.7; PT 1.9; SPé 2.5; SP? 3.4-3.9; TPF 4.0.
AFROTROPICAL MEGARTHRUS 669
3d. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 12e) shorter
than metafemur. Mesotibia (Fig. 12c) shorter than metatibia (Fig. 12d), with peg-like
setae arranged in a single row, and grouped in a field near apex. Peg-like setae on meta-
tibia arranged in a single row, and in a double row near apex; forming a double row on
mesotrochanter (Fig. 12e); absent from protibia and metatrochanter. Apex of abdominal
tergite 8 as in Fig. 12f, g. Sternite 8 as in Fig. 12g. Aedeagus as in Fig. 12a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 13c, f. Sternite 8 as
in Fig. 13g. Genital segment as in Fig. 13d, e.
Comments. Megarthrus clarkei and M. scotti may be distinguished from other
African congeners by the scutellar shape. The shape of the female abdominal tergite 8
is diagnostic for each of these two species.
The species is dedicated to one of the collectors, Mr. Robin O. S. Clarke.
Megarthrus congoensis Cameron (Figs 2d, k, 14a-1, 15a-g)
Megarthrus congoensis CAMERON, 1950: 4.
Type material. Holotype 9: Rwanda, V. Visoke (Albert NP) Bishoke, 2800-3300m, 13-
14.11.1935 (G. F. de Witte) #1128, MRAC.
Paratypes (13): same data as holotype, 12 in MRAC; same data, but #1125, 32 in
BMNH: same data, but #1127, 16 in MHNG and 12 in MRAC; same data, but #1129, 16,19
in BMNH, 19 in MHNG and 26,22 in MRAC.
Additional material (19): Rwanda (Albert NP) at foot of V. Karisimbi, Ilega, 2400m,
12.11.1935 (G. F. de Witte) #1314, MRAC (paratype of M. apicicornis).
Distribution. Rwanda: V. Visoke and V. Karisimbi.
Description. Length 1.65-1.75 mm; width 0.95-1.05 mm. Body red-brown or
dark brown, with darkened head, sutural margin of elytron, metasternum and abdomen,
and paler legs, mouth parts and antennomeres 1-4. Dorsal pubescence fairly uniform;
that on elytron sparser, but becoming denser on humeral area. Pubescence denser near
posterior margin of abdominal tergite 7. Metasternal setae becoming sparser mediopos-
teriorly and longer anteriorly; shorter than those on prosternum. Pubescence of abdo-
minal sternites 4-7 uniform, except for a pair of long setae near medioapical margins.
Puncturation fine on anterior portion of hypomeron; median area of metasternum im-
punctate. Frons raised above level of vertex. Anterior frontal edge not carinate, evenly
arcuate. Entire frontal impression shallow. Eye moderatly convex, with highest point
below level of vertex. Temple similar to that in Fig. 2k. Submentum weakly convex.
Antenna (Fig. 15a) without patches of sensilla; antennomeres 3 and 4 slightly asym-
metrical. Pronotal disc (Fig. 15c) shallowly depressed along basal portion of lateral
edge and deeply depressed along median groove; latter shallow, parallel-sided. Anterior
prosternal margin bordered by an irregular row of fine longitudinal ridges. Protro-
chanter without transverse ridge. Lateral portion of prepectal ridge straight. Scutellum
as in Fig. 2d. Elytron not narrowed at base; base gradually inclined, overhanging.
Elytral disc flat along lateral edge; latter straight in dorsal view; apical margin straight
or convex near suture. Metasternal median ridge absent. Abdominal tergite 3 slightly
vaulted transversally. Sternites 2 and 3 with median processes as in Fig. 21. Sernite 4
flat at base, then slightly vaulted transversally.
670 GIULIO CUCCODORO & IVAN LÒBL
RatrosssAle 12922705 BES IS EL S 192 ED 17-182 ENAIM ENS 8 Gil
2.02GW7 2:0; HW 1:6-1.7; MIE1:5=1°6; MP)1.7;, PT 187 SPS 23522 ASP
TPF 4.0.
d. Inner apical angle of elytron obtuse. Protarsal segment 1 bearing adhesive
setae. Mesofemur (Fig. 14h) as long as metafemur (Fig. 141). Mesotibia (Fig. 14f)
shorter than metatibia (Fig. 14g). Peg-like setae grouped in a field on mesotrochanter
(Fig. 14h), mesotibia, metatrochanter (Fig. 141) and metatibia; absent from protibia.
Apex of abdominal tergite 8 as in Fig. 14a, e. Sternite 8 as in Fig. 14b . Aedeagus as in
Fig. 14c, d.
9. Inner apical angle of elytron right-angled. Medioapical projection of abdo-
minal tergite 8 as in Fig. 15b, e. Sternite 8 as in Fig. 15g. Genital segment as in Fig.
dit
Comments. Megarthrus congoensis differs from other species with the head
widened behind the eyes (see discussion under M. apicicornis), M. hutu excepted, by
the shape of the male and female abdominal tergite 8. It may be distinguished from M.
hutu by the aedeagal shape and, in female, by the ratio of the elytral / pronotal lengths.
Megarthrus dominicae sp. n. (Figs 16a-1, 17a-f)
Type material. Holotype d: Uganda, Toro prov., Ruwenzori NP, John Mate Camp,
3350m, 14-15.v.1993 (G. Cuccodoro & D. Erne) ex decaying stems of giant Lobelia, MHNG.
Paratypes (10): same data as holotype, 18, 19 in BMNH, 38, 39 in MHNG and là,
19 in MRAC.
Distribution. Uganda: Ruwenzori Range.
Description. Length 1.8-2.0 mm; width 1.1-1.3 mm. Head, metasternum and
abdomen blackish; pronotum and elytron dark brown; sutural margin of elytron
darkened; mouth parts and legs yellowish; antenna dark brown, antennomeres 1-4
somewhat paler. Dorsal pubescence fairly uniform; longer and denser along pronotal
median groove and becoming longer and denser on humeral area of elytron; denser near
posterior margin of abdominal tergite 7. Metasternal setae shorter than those on
prosternum, becoming sparser medioposteriorly and longer anteriorly. Pubescence on
abdominal sternites 4-7 uniform, except for a pair of long setae near medioapical
margins. Anterior portion of hypomeron and median area of metasternum impunctate.
Frons raised above level of vertex. Anterior frontal edge not carinate, evenly arcuate.
Frontal impression indistinct in middle and shallow laterally. Eye moderatly convex,
with highest point below level of vertex. Temple similar to that in Fig. 2g. Submentum
convex. Antenna (Fig. 16a) without patches of sensilla; antennomeres 3 and 4 slightly
asymmetrical. Pronotal disc (Fig. 17a) flat along lateral edge, shallowly depressed
along median groove; latter shallow, widened at base. Anterior prosternal margin
bordered by an irregular row of fine longitudinal ridges. Protrochanter without trans-
verse ridge. Lateral portion of prepectal ridge straight. Scutellum similar to that in Fig.
2b. Elytron abruptly narrowed at base; base abruptly inclined, overhanging. Elytral disc
deeply depressed along lateral edge; latter slightly convex in dorsal view; apical margin
straight or arcuate near suture; inner apical angle obtuse. Metasternal median ridge fine,
AFROTROPICAL MEGARTHRUS 671
low. Abdominal tergite 3 flat. Sternites 2 and 3 with median processes as in Fig. 21.
Sternite 4 flat at base, then slightly vaulted transversally.
Ratios A2 /0SEEU"S EMI EN 1:22EY332347G2236W21:6-1.75HW.
18687 AMIE MP8 Pie? "0 SP 225.0 PK 5:0)
3d. Apical contour of elytron convex near suture. Protarsal segment 1 bearing
adhesive setae. Mesofemur (Fig. 16e) longer than metafemur. Mesotibia (Fig. 16d)
longer than metatibia. Peg-like setae arranged in a double row on mesotibia and
metatrochanter (Fig. 16g), grouped in a field on mesotrochanter (Fig. 16e), absent from
protibia and metatibia. Apex of abdominal tergite 8 as in Fig. 16f, h. Sternite 8 as in
Fig. 161. Aedeagus as in Fig. 16b, c.
2. Apical contour of elytron somewhat sinuate near suture. Medioapical pro-
jection of abdominal tergite 8 as in Fig. 17b, c. Sternite 8 as in Fig. 17f. Genital
segment as in Fig. 17d, e.
Comments. Megarthrus dominicae may be distiguished easily of the others
species of similar size (see discussion under M. basilewskyi) by its uniformly dark
colour and impunctate prothoracic hypomera.
The species is dedicated to Mrs Dominique Cuccodoro.
Megarthrus falasha sp. n. (Figs 2d,k, 18a-i, 19a-g)
Type material. Holotype 3: Ethiopia, Gojam prov., 8km W Falega, Birham <10°46'N;
38°03'E> 2820m, xi.1972 (R. O. S. Clarke) BMNH.
Paratypes (7): same data as holotype, 19 in BMNH; Jimma (= Kefa prov., Jima?)
vi.1971 (R. O. S. Clarke) ex grass cuttings, 29 in BMNH and 1° in MHNG; Shewa prov., Addis
Abeba, 1971 (G. de Rougemont) 16, 19 in MHNG; Mt. Zuquäla (=Mt. Ziq’wala), in crater, c.
2700m, 26.x.1926 (J. Omer-Cooper) box 77, 1 © in FMNH (paralectotype of M. abessinus Bern-
hauer).
Distribution. Central Ethiopia.
Description. Similar to M. wittei from which it may be distinguished as follows:
Frons raised above level of vertex. Antenna as in Fig. 19a. Anterior prosternal margin
not bordered by longitudinal ridges. Elytron with lateral contour somewhat convex and
apical contour straight near suture. Ratios: AL 1.9; EL 1.45; ET 1.45-1.55; EY 2.8-2.9;
GW 1.6-1.7; HW 1.6-1.7; ML 1.4; PT 1.9; SPé 2.8-2.9; SP9 3.3-3.4; TPF 10.0.
d. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 18f) somewhat
shorter than metafemur. Mesotibia (Fig. 18c) shorter than metatibia (Fig. 18d). Meta-
trochanter (Fig. 18h) bearing 1-3 peg-like setae. Peg-like setae on mesotibia arranged in
a single row, and in a double row near apex; arranged in a single row on meso-
trochanter (Fig. 18f) and metatibia; absent from protibia. Apex of abdominal tergite 8
as in Fig. 18e, g. Sternite 8 as in Fig. 181. Aedeagus as in Fig. 18a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 19d, f. Sternite 8 as
in Fig. 19g. Genital segment as in Fig. 19c, e.
Comments. Megarthrus falasha is one of the seven Afrotropical species (M.
falasha, M. nanus, M. ovalis, M. rougemonti, M. twa, M. wittei and M. zulu) possessing
moniliform antennomeres 7 to 10. It may be distinguished from these species, M. wittei
excepted, by the almost uniformly brown color of the body in combination with the
672 GIULIO CUCCODORO & IVAN LOBL
aedeagus symmetrical. The pattern of the mesotrochanteral peg-like setae is diagnostic
for each of these two species.
Megarthrus gigas Fagel (Figs 2m, 20a-f, 21a-e, 22a-i)
Megarthrus gigas FAGEL, 1957: 29.
Type material. Holotype d: Burundi, Ngozi terr., Samutuku, 2600m, 24.ii.1953 (P.
Basilewsky) MRAC.
Paratypes (29 ): Zaire, Kivu prov., Kabare terr., foothills SE Kahuzi, 2000m, vii.1951 (N.
Leleup) ex humus in montane forest, MRAC.
Additional material (26): Zaire, Kivu prov., Mwenga terr., Lake Lungwe, 2700m,
viti.1953 (N. Leleup) ex humus in forest of bamboo and Hagenia, MRAC (mislabelled paratype);
Ruwenzori Range (Albert NP) Kalonge, stream Katsambu, tributary of Butahu, 2000m, 27.1-
9.11.1953 (P. Vanschuytbroeck & J. Kekenbosch) #2203-10, MHNG.
Distribution. Burundi; Zaire: Kivu, Ruwenzori Range.
Description. Length 2.0 mm long; width 1.4 mm wide. Body uniformly dark
brown, with paler legs, mouth parts and antennomere 11. Dorsal pubescence fairly
uniform, becoming sparser on lateral edges of pronotum. Humeral area of elytron with
denser pubescence. Metasternal setae becoming denser near median area; about as long
as those on prosternum. Pubescence of abdominal sternites 4-7 uniform. Puncturation
coarse on anterior portion of hypomeron and on median area of metasternum. Frons on
level with vertex. Anterior frontal edge finely carinate, evenly arcuate. Entire frontal
impression indistinct. Eye almost hemispherical, raised above level of vertex. Temple
similar to that in Fig. 21. Submentum flat. Antenna (Fig. 21c) without patches of sen-
silla; antennomeres 3 and 4 symmetrical. Pronotal disc (Fig. 21a) deeply depressed
along entire lateral edge, flat along median groove; latter shallow, widened at base.
Anterior prosternal margin bordered by an irregular row of fine longitudinal ridges.
Protrochanter without transverse ridge. Lateral portion of prepectal ridge sinuate. Scu-
tellum similar to that in Fig. 2f. Elytron abruptly narrowed at base; base abruptly incli-
ned, overhanging. Elytral disc deeply depressed along lateral edge; latter almost straight
in dorsal view; apical margin sinuate near suture. Metasternal median ridge fine, low.
Abdominal tergite 3 slightly vaulted transversally. Abdominal sternites 2 and 3 with
median processes as in Fig. 2m. Sternite 4 flat at base, then slightly vaulted trans-
versally.
Ratios: AL 2.6; EL 1:4; ET 1.6; EW 1.2; EY 2.7-2.8-GT2.22GW EGE
DD DS NEA MP" TE 2020 SPS'0SSSTPF ADS
d. Inner apical angle of elytron right-angled. Protarsal segment 1 (Fig. 22g)
lacking adhesive setae. Mesofemur (Fig. 20d) shorter than metafemur (Fig. 20a).
Mesotibia (Fig. 22f) shorter than metatibia (Fig. 22e). Peg-like setae arranged in a
single row on mesotrochanter (Fig. 20d), metatrochanter (Fig. 20a) and metatibia;
grouped in a field on protibia (Fig. 22g) and mesotibia. Apex of abdominal tergite 8 as
in Fig. 20e, f. Sternite 8 as in Fig. 21d. Aedeagus as in Fig. 20b, c.
2. Apical contour of elytron and inner apical angle as in Fig. 21b. Abdominal
tergite 8 (Fig. 22h, i) without medioapical projection. Sternite 8 as in Fig. 21d. Genital
segment as in Fig. 22a-d; mediodorsal suture of sternite 9 forming an angle not
projecting anteriorly.
AFROTROPICAL MEGARTHRUS 673
Comments. Megarthrus gigas may be distinguished from other species of similar
size (see discussion under M. basilewskyi) by the aedeagus lacking a lateral notch. This
species shares with M. major the angulate median suture of the abdominal sternite 9,
but differs by the much wider basal projection of that sternite.
Megarthrus horticola sp. n. (Figs 23a-h, 24a-g)
Type material. Holotype d: Uganda, Bugisu prov., Mt. Elgon, Sipi, 1750m, 31.v.1993
(G. Cuccodoro & D. Erne) ex moist leaf litter at foot of tree in banana plantation, MHNG.
Paratypes (8): same data as holotype, 19 in BMNH and 19 in MHNG; same data, but
1.vi.1993, ex moist vegetational debris in garden, 12 in MHNG and 1% in MRAC; Kenya,
Central prov., Mt. Aberdares, near NP entrance, 2300m, 25.x1.1974, (V. Mahnert & J. -L. Perret)
ex leaf litter and rotten wood, 16, 12 in MHNG; Rift Valley prov., Mau for., between Mau
Saummit and Kedowa, near Londiani Rd., 7.x1.1974 (V. Mahnert & J. -L. Perret) under bark and
stones, 19 in MHNG; Zaire, Kivu prov., Mt. Kahuzi, 2000m, 27.v.1985 (H. Mühle) ex Lobelia,
16 in ZMHB.
Distribution. Kenya: Mt. Aberdares; Uganda: Mt. Elgon; Zaire: Kivu.
Description. Similar to M. monticola from which it may be distinguished as
follows: Length 1.35-1.45 mm; width 0.95-1.05 mm. Sutural margin of elytron darker
than disc; antenna reddish-brown, antennomere 11 paler. Antenna as in Fig. 24a.
Pronotal disc (Fig. 24f) flat along lateral edge. Elytron with apical contour arcuate near
suture and inner apical angle obtuse. Abdomen with tergite 3 weakly convex and
sternite 4 flat. Ratios: AL 2.2; EL 1.45; EW 1.2; HW 1.6-1.7; TPF 5.0.
3. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 23f) as long as
metafemur. Mesotibia (Fig. 23c) shorter than metatibia. Peg-like setae grouped in field
on mesotrochanter (Fig. 23f), mesotibia and metatrochanter (Fig. 23h), absent from
protibia and metatibia. Apex of abdominal tergite 8 as in Fig. 23d, e. Sternite 8 as in
Fig. 23g. Aedeagus as in Fig. 23a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 24e, g. Sternite 8 as
in Fig. 24d. Genital segment as in Fig. 24b, c.
Comments. This species is similar to M. mahnerti and M. monticola. It may be
distinguished easily by the apex of the female sternite 8 truncate and, in the male, by
the absence of metatibial peg-like setae.
The water loading behaviour has been reported in this species (Megarthrus sp. A
in Cuccoporo, 1995).
Megarthrus hutu sp. n. (Figs 25a-h, 26a-h)
Type material. Holotype 6: Rwanda (Albert NP) V. Visoke, Bishoke, 2800-3300m, 13-
14.11.1935 (G. F. de Witte) #1127, MRAC (paratype of M. congoensis).
Paratypes (13): same data as holotype, but #1128, 12 in MRAC (paratype of M.
congoensis), same data, but #1129, 16 in MHNG and 29 in MRAC (paratypes of M.
congoensis); at foot of V. Karisimbi (Albert NP) Lake n'Gando, 2400m, 6.111.1935 (G. F. de
Witte) #1243, 13 in BMNH (paratype of M. apicicornis); same data, but 8.iii.1935, #1216, 19 in
MRAC (paratype of M. congoensis); Zaire, Kivu (Albert NP) Tshiaberimu sect., Kirungu,
2720m, 8-9.111.1954 (P. Vanschuytbroeck & H. Synave) #7928-31, 12 in MHNG.
Distribution. Rwanda; Zaire: Kivu.
674 GIULIO CUCCODORO & IVAN LOBL
Description. Similar to M. congoensis from which it may be distinguished as
follows: Length 1.35-1.60 mm; width 0.85-1.00 mm. Elytron with apical contour
somewhat convex near suture and inner apical angle obtuse. Abdomen with pubescence
uniform and tergite 3 weakly convex. Ratios: EL 1.4-1.5; ET 1.7-1.8; EY 2.8-2.9; GT
PIDASE EIZO SO?! 23 APRO!
d. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 25e) as long as
metafemur (Fig. 25h). Mesotibia (Fig. 25c) with peg-like setae arranged in a double
row; somewhat shorter than metatibia (Fig. 25d). Peg-like setae grouped in a field on
metatrochanter (Fig. 25h), mesotrochanter (Fig. 25e) and metatibia, absent from
protibia. Apex of abdominal tergite 8 as in Fig. 25f, g. Sternite 8 as in Fig. 26f.
Aedeagus as in Fig. 25a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 26b, d. Sternite 8 as
in Fig. 26c. Genital segment as in Fig. 26e, g.
Comments. See discussion under M. congoensis.
Megarthrus kamerunensis Bernhauer
Megarthrus kamerunensis BERNHAUER, 1942: 348.
Type material. Type: Cameroon, SE Mt. Cameroun, 1800-2000m, 1939 (P. Lepesme, R.
Paulian & A. Villiers) ex fungi, FMNH (except for the remains of tarsi, the type specimen is
missing and was apparently lost in transit when returned to the FMNH after W. O. Steel's death in
October 1969; A. F. Newton Jr., pers. com.).
Distribution. Cameroon: Mt. Cameroon.
Comments. According to the original desciption, M. kamerunensis is charac-
terised by a rounded incision of the posterior margin of the male abdominal sternite 6.
No material could be assigned to this species.
Megarthrus magnicaudatus sp. n. (Figs 27a-k, 28a-f)
Type material. Holotype d: Ethiopia, Balé prov., Adelay Forest, 3100m, 19.xii.1971 (R.
O. S. Clarke) ex plant roots in Juniper wood, BMNH.
Paratypes (59): same data as holotype, 2 in BMNH and 2 in MHNG; Ethiopia, Gamo
Gofa prov., Mt. Gughé, 3200-3350m, 20.x11.1948 (H. Scott) ex roots of fern, 1 in BMNH.
Distribution. Central and southern Ethiopia.
Description. Similar to M. abessinus from which it may be distinguished as
follows: Length 1.6-1.7 mm; width 1.1-1.2 mm. Frons raised above level of vertex.
Frontal impression indistinct in middle, shallow laterally. Antenna as in Fig. 28c and
pronotum as in Fig. 271. Ratios: AL 2.0; EL 1.4; ET 1.75; HW 1.6-1.7; TPF 1.8.
3d. Apical contour of elytron sinuate near suture. Protarsal segment 1 (Fig. 27f)
bearing adhesive setae. Mesofemur (Fig. 27c) shorter than metafemur. Metatibia (Fig.
27h) with peg-like setae arranged in a double row; longer than mesotibia (Fig. 27g).
Peg-like setae arranged in a single row on mesotrochanter (Fig. 27c); grouped in a field
on protibia (Fig. 27f) and mesotibia; lacking on metatrochanter. Abdominal tergite 8 as
in Fig. 27b, k. Sternite 8 as in Fig. 27a. Aedeagus as in Fig. 27d, e.
AFROTROPICAL MEGARTHRUS 675
2. Apical contour of elytron convex near suture. Abdominal tergite 8 as in Fig.
28b, f. Sternite 8 as in Fig. 28e. Genital segment as in Fig. 28a, d.
Comments. See discussion under M. abessinus.
Megarthrus mahnerti sp. n. (Figs 29a-1, 30a-g)
Type material. Holotype d: Kenya, Central prov., Mt. Aberdares, near NP entrance,
2300m, 25.x1.1974, (V. Mahnert & J. -L. Perret) ex leaf litter and rotten wood, MHNG.
Paratypes (8): same data as holotype, 16, 12 in BMNH, 14,29 in MHNG and 16,19
in MRAC; Kenya, Western prov., Mt. Elgon NP, slope of Koitobos Pk., c. 2700m, 15.x1.1974 (V.
Mahnert & J. -L. Perret) ex litter under bamboo, 1 2 in MHNG;
Additionnal material (1 2). D. O. Afrika, Langenburg (= Tanzania, Iringa prov., Living-
stone Range, Ukenju?) 19.vii.1899 (Fülleborn), ZMHB. The specimen is likely to be conspecific.
It is smaller and differs in the pronotal relief.
Distribution. Kenya: Mt. Aberdares; Tanzania: Livingstone Range.
Description. Similar to M. monticola from which it may be distinguished as
follows: Antenna as in Fig. 29d and abdominal sternite 4 flat. Ratios: AL 2.0; EL 1.8;
Mg WA EP E SV.
d. Apical contour of elytron convex near suture; inner apical angle obtuse.
Protarsal segment | bearing adhesive setae. Mesofemur (Fig. 29b) as long as meta-
femur. Mesotibia (Fig. 29g) shorter than metatibia (Fig. 29a), both with peg-like setae
arranged in a single row, and grouped in a field near apex. Peg-like setae arranged in a
single row on mesotrochanter (Fig. 29b) and metatrochanter (Fig. 29h), absent from
protibia. Apex of abdominal tergite 8 as in Fig. 29c. Sternite 8 as in Fig. 291. Aedeagus
as in Fig. 29e, f.
2. Apical contour of elytron and inner apical angle as in Fig. 30g. Medioapical
projection of abdominal tergite 8 as in Fig. 30d, e. Sternite 8 as in Fig. 30b. Genital
segment as in Fig. 30a, c.
Comments. This species is similar to M. horticola and M. monticola. As M.
monticola, it differs from M. horticola by the bilobed apex of the female sternite 8. It
may be distinguished easily from M. monticola by the shape of the apex of the
aedeagus.
The species is dedicated to one of the collectors, Mr. Volker Mahnert.
Megarthrus major sp. n. (Fig. 31a-k)
Type material. Holotype 9 : Zaire, Kivu (Albert NP) Tshiaberimu sect., Kirungu, 2720m,
8-9.11.1954 (P. Vanschuytbroeck & H. Synave) #7928-31, MRAC.
Distribution. Zaire: Kivu.
Description. Similar to M. gigas from which it may be distinguished as follows:
Ratios: EW 1.3; EY 3.3-3.4; SP 4.5-4.6; TPF abs.
2. Apical contour of elytron as in Fig. 31k. Abdominal tergite 8 (Fig. 31c, g)
without medioapical projection. Sternite 8 as in Fig. 31i. Genital segment as in Fig. 31a,
b, d, f; mediodorsal suture of sternite 9 forming an angle projecting anteriorly.
3. Unknown.
Comments. See discussions under M. basilewskyi and M. gigas.
676 GIULIO CUCCODORO & IVAN LOBL
Megarthrus maniwaata sp. n. (Figs 32a-h, 33a-g)
Type material. Holotype dé: Zaire, Ruwenzori Range (Albert NP) Kalonge, river
Nyamwamba, tributary of Butahu, 2010m, 2-3.11.1953 (P. Vanschuytbroeck & J. Kekenbosch)
#2214-21, MRAC.
Paratypes (22): same data as holotype, 34,29 in MHNG and 86, 49 in MRAC; Zaire,
Ruwenzori Range (Albert NP) Kalonge, river Katsambu, tributary of Butahu, 2000m, 26.1-
19.11.1953 (P. Vanschuytbroeck & J. Kekenbosch) #2155-2200, 16 in MRAC; Ruwenzori Range
(Albert NP) Kyandolire, river Mulaku, tributary of Kakalari, 1750m, 14.x.1952 (P. Van-
schuytbroeck & J. Kekenbosch) ex humus, #1241-43, 16 in MRAC; same data, but #1302, 16 in
MRAC; Kivu (Albert NP) Nyasheke, V. Nyamuragira, 1820m, 14-26.vi.1935 (G. F. de Witte)
#1481, 16 in MRAC; Rwanda, Cyangugu pref., around Nyakabuye, 9.vi.1985 (H. Mühle) 14 in
ZMHB.
Distribution. Rwanda; Zaire: Kivu, Ruwenzori Range.
Description. Similar to M. mwami from which it may be distinguished as fol-
lows: Length 1.15-1.40 mm; width 0.80-0.95 mm. Antenna as in Fig. 33c. Pronotum as
in Fig. 33f. Elytron abruptly narrowed at base. Metasternal median ridge fine or absent.
RAOS EEE TS; Biel 6-1-7; EY 2.8-2.9: Pl 1.9-2.0:5P 2.523.0:20BEI6327720:
3. Protarsal segment | bearing adhesive setae. Mesofemur (Fig. 32d) as long as
metafemur (Fig. 32f). Metatibia (Fig. 32e) with peg-like setae arranged in a double row;
longer than mesotibia (Fig. 32c). Protibia not modified. Metatrochanter (Fig. 32f) with
1-3 peg-like setae. Mesotrochanter (Fig. 32d) and mesotibia with peg-like setae grou-
ped in a field. Apex of abdominal tergite 8 as in Fig. 32h. Aedeagus as in Fig. 32a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 33e, g. Sternite 8 as
in Fig. 33d. Genital segment as in Fig. 33a, b.
Comments. Megarthrus maniwaata, M. merabet and M. mwami have in commun
an evenly and broadly rounded apical margin of the scutellum. They may be distin-
guished by the sexual characters, in particular by the shape of the apical portion of the
aedeagal median lobe and of the basal portion of the female coxites.
Megarthrus merabet sp. n. (Figs 34a-h, 35a-g)
Type material. Holotype 6: Rwanda, Cyangugu pref., around Nyakabuye, 9.vi.1985 (H.
Mühle) ZMHB.
Paratypes (3): Zaire, Ruwenzori Range (Albert NP) Kyandolire, river Mulaku, tributary
of Kakalari, 1750m, 14.x.1952 (P. Vanschuytbroeck & J. Kekenbosch) ex humus, #1241-43, 16,
12 in MRAC; Mutsora (Albert NP) river Katunda, 1600m, 10.11.1953 (P. Vanschuytbroeck & J.
Kekenbosch) #2432, 14 in MHNG.
Distribution. Rwanda; Zaire: Ruwenzori Range.
Description. Similar to M. mwami from which it may be distinguished as
follows: Length 1.15-1.35 mm; width 0.80-0.95 mm. Antenna as in Fig. 35c and prono-
tum as in Fig. 35f. Elytron abruptly narrowed at base; apical contour convex near
suture. Ratios: EL 1.4-1.5; ET 1.4-1.5; ML 1.4; SP 2.2-2.5; TPF 10.0.
3. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 34e) somewhat
longer than metafemur (Fig. 34f). Metatibia (Fig. 34d) longer than mesotibia (Fig. 34c).
Metatrochanter (Fig. 34f) with 1-4 peg-like setae. Peg-like setae grouped in a field on
mesotrochanter (Fig. 34e), absent from protibia; those on metatibia arranged in a single
row, and in a double row near apex, on mesotibia forming in a double row, and grouped
AFROTROPICAL MEGARTHRUS 677
in a field near apex. Apex of abdominal tergite 8 as in Fig. 34h. Aedeagus as in Fig.
34a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 35e, g. Sternite 8 as
in Fig. 35d. Genital segment as in Fig. 35a, b.
Comments. See discussion under M. maniwaata. The apical projection of the
abdominal tergite 8 is particularly small in this species.
Megarthrus monticola Cameron (Figs 36a-1, 37a-h)
Megarthrus monticola CAMERON, 1942: 322.
Type material. Holotype d : Kenya, Rift Valley prov., Chyulu Hills, 1700m, vi.1938,
BMNH.
Paratypes (10): same data as holotype, 14,89 in BMNH and1@ in MHNG.
Additional material (2): same data as holotype, but vii.1938, 16, 12 in MHNG.
Distribution. Kenya: Chyulu Hills.
Description. Length 1.6-1.7 mm; width 1.0-1.2 mm. Head, metasternum and
abdomen dark brown; pronotum and entire elytron reddish-brown; mouth parts and legs
yellowish-brown; antenna reddish-brown, antennomeres 1-4 and 11 paler. Pubescence
on pronotum denser along median groove; elytral pubescence sparser, but becoming
denser on humeral area; that on abdomen shorter. Metasternal setae shorter than those
on prosternum, becoming sparser on median area. Pubescence of abdominal sternites 4-
7 uniform, except for a pair of long setae near medioapical margins. Puncturation fine
on anterior portion of hypomeron; median area of metasternum impunctate. Frons on
level with vertex. Anterior frontal edge not carinate, evenly arcuate. Entire frontal
impression indistinct. Eye strongly convex, with highest point reaching level of vertex.
Temple similar to that in Fig. 2h. Submentum weakly convex. Antenna (Fig. 37a)
without patches of sensilla; antennomeres 3 and 4 slightly asymmetrical. Pronotal disc
(Fig. 37d) shallowly depressed along entire lateral edge, flat along median groove;
latter shallow, parallel-sided. Anterior prosternal margin not bordered by longitudinal
ridges. Protrochanter without transverse ridge. Lateral portion of prepectal ridge
straight. Scutellum similar to that in Fig. 2b. Elytron abruptly narrowed at base; base
abruptly inclined, overhanging. Elytral disc shallowly depressed along lateral edge;
latter slightly convex in dorsal view. Metasternal median ridge fine, low. Abdominal
tergite 3 slightly vaulted transversally. Sternites 2 and 3 with median processes as in
Fig. 21. Sternite 4 flat at base, then slightly vaulted transversally.
RAHOS AIMÉ EE AS ERI EW 133END27IGE22ZGWEIE6- 173 2240)
MESIE6FMEPT.8; PE 1.9: SP 25; 1 PF: 5.0.
3. Apical contour of elytron arcuate near suture; inner apical angle obtuse.
Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 36g) shorter than
metafemur (Fig. 36h). Mesotibia (Fig. 36e) shorter than metatibia (Fig. 36f), both with
peg-like setae arranged in a double row and grouped in a field near apex. Peg-like setae
arranged in a single row on mesotrochanter (Fig. 36g), double row on metatrochanter
(Fig. 36h); absent from protibia. Apex of abdominal tergite 8 as in Fig. 36a, i. Sternite 8
as in Fig. 36b. Aedeagus as in Fig. 36c, d.
678 GIULIO CUCCODORO & IVAN LÒBL
9. Apical contour of elytron and inner apical angle as in Fig. 37h. Medioapical
projection of abdominal tergite 8 as in Fig. 37c, g. Sternite 8 as in Fig. 37b. Genital
segment as in Fig. 37e, f.
Comments. See discussions under M. horticola and M. mahnerti.
Megarthrus mukankundiyeorum sp. n. (Figs 38a-1, 39a-g)
Type material. Holotype d: Rwanda, 25km N Kibuye, Kayove, 2100m, 15.v.1973 (P.
Werner) ex vegetational debris in secondary forest, MHNG.
Paratypes (29): Rwanda, Cyangugu pref., around Nyakabuye, 22.11.1985 (H. Miihle) in
MHNG and ZMHB.
Distribution. Rwanda.
Description. Length 1.7-1.8 mm; width 1.1-1.2 mm. Body predominantly red-
brown or dark brown, with darkened head and sutural margin of elytron, and paler legs,
mouth parts and antennomere 11. Dorsal pubescence fairly uniform, shortened on
abdomen, becoming sparser near lateral pronotal edge, denser along sutural margin and
on humeral area of elytron. Metasternal setae about as long as those on prosternum,
becoming denser near median area. Pubescence on abdominal sternites 4-7 becoming
denser near medioapical margins. Puncturation coarse on anterior portion of hypomeron
and on median area of metasternum. Frons raised above level of vertex. Anterior frontal
edge finely carinate; frontal impression distinct. Eye almost hemispherical, with highest
point reaching level of vertex. Temple similar to that in Fig. 21. Submentum flat.
Antenna (Fig. 39e) without patches of sensilla; antennomeres 3 and 4 symmetrical.
Pronotal disc (Fig. 39g) slightly depressed along entire lateral edge, flat along median
groove; latter deep, parallel-sided. Anterior prosternal margin bordered by a regular row
of conspicuous longitudinal ridges. Protrochanter without transverse ridge. Lateral
portion of prepectal ridge sinuate. Scutellum similar to that in Fig. 2f. Elytron not
narrowed at base; base abruptly inclined, overhanging. Elytral disc flat along lateral
edge; latter straight in dorsal view; apical margin somewhat sinuate near suture; inner
apical angle almost right-angled. Metasternal median ridge absent. Abdominal tergite 3
slightly vaulted transversally. Sternite 4 flat at base, then slightly vaulted transversally.
Ratios: AL 2.67 EL 1.5; ET 1-7; EW 1.27 EY 2.8-2:97 Git 2753 GWS Wale
eM IPO SP tr 1292 SP2-7-310- PE abs
d. Anterior frontal edge subtruncate. Protarsal segment 1 lacking adhesive
setae. Mesofemur (Fig. 38h) shorter than metafemur. Mesotibia (Fig. 38f) shorter than
metatibia (Fig. 38g), both with peg-like setae grouped in a field. Metatrochanter (Fig.
38e) with a single peg-like seta. Peg-like setae arranged in a single row on meso-
trochanter (Fig. 38h), absent from protibia. Apex of abdominal tergite 8 as Fig. 38c, d.
Sternite 8 as Fig. 381. Aedeagus as in Fig. 38a, b.
2. Anterior frontal edge evenly arcuate. Abdominal tergite 8 (Fig. 39c, d)
lacking medioapical projection, with dark tip. Sternite 8 (Fig. 381) about 1.8x as long as
width of its basal projection. Genital segment as in Fig. 39a, b.
Comments. Among the species possessing a ridged prosternal margin (see dis-
cussion under M. africanus), M. mukankundiyeorum may is characterised by the asym-
metrical aedeagus, the long antennae and the length of the female abdominal sternite 8.
AFROTROPICAL MEGARTHRUS 679
The species is dedicated to the family of our friend Miss Consolée Mukan-
kundiye, who suffered recently in Rwanda.
Megarthrus mwami sp. n. (Figs 2e, 40a-i, 41a-g)
Type material. Holotype dé: Burundi, Bururi terr., Nyamurenbe, 900m, 7.111.1953 (P.
Basilewsky) MRAC.
Paratype (2): same data as holotype, 12 in MRAC; Rwanda, Cyangugu pref., around
Nyakabuye, 1-30.xii.1982 (H. Mühle) 1d in ZMHB.
Distribution. Burundi; Rwanda.
Description. Length 1.2-1.3 mm; width 0.9-1.0 mm. Head, metasternum and
abdomen dark brown, pronotum and elytron somewhat paler; sutural margin of elytron
darkened; mouth parts and legs yellowish-brown; antenna reddish-brown, antenno-
meres 11 paler. Dorsal pubescence fairly uniform, on pronotum denser along median
groove. Elytral pubescence sparser, but becoming denser on humeral area. Abdomen
with denser pubescence near posterior margin of tergite 7. Metasternal setae shorter
than those on prosternum, becoming sparser medioposteriorly and longer anteriorly.
Pubescence of abdominal sternites 4-7 uniform, except for a pair of long setae near each
medioapical margin. Puncturation fine on anterior portion of hypomeron; median area
of metasternum impunctate. Frons raised above level of vertex. Anterior frontal edge
finely carinate, evenly arcuate. Entire frontal impression shallow. Eye strongly convex,
with highest point reaching level of vertex. Temple similar to that in Fig. 2h. Sub-
mentum weakly convex. Antenna (Fig. 41a) without patches of sensilla; antennomeres
3 and 4 slightly asymmetrical. Pronotal disc (Fig. 41f) shallowly depressed along basal
portion of lateral edge, deeply depressed along median groove; latter deep, parallel-
sided. Anterior prosternal margin bordered by an irregular row of fine longitudinal
ridges. Protrochanter without transverse ridge. Lateral portion of prepectal ridge
straight. Scutellum as in Fig. 2e. Elytron somewhat narrowed at base; base abruptly
inclined, overhanging. Elytral disc shallowly depressed along lateral edge; latter
subangulate in dorsal view; apical margin straight or convex near suture; inner apical
angle obtuse. Metasternal median ridge fine, low. Abdominal tergite 3 slightly vaulted
transversally. Sternites 2 and 3 with median processes similar to that in Fig. 2m.
Sternite 4 flat at base, then slightly vaulted transversally.
Ratios:e Ale ONE PAPAS EIMECRE WAP TES GER 0G Wires IW 1e6=
fey Mile als: MPA: PP 18-19: SP2.022 520 PFi6.5-7.0:
3. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 40g) as long as
metafemur (Fig. 40h). Mesotibia (Fig. 40c) shorter than metatibia (Fig. 40d). Meta-
trochanter (Fig. 40h) with 1-3 peg-like setae. Peg-like setae arranged in a single row on
mesotrochanter (Fig. 40g); absent from protibia; those on mesotibia arranged in a single
row, and in a double row near apex; those on metatibia arranged in a single row, and
grouped in a field near apex. Apex of abdominal tergite 8 as in Fig. 40e,i. Aedeagus as
in Fig. 40a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 41d, g. Sternite 8 as
in Fig. 41e. Genital segment as in Fig. 41b, c.
Comments. See discussions under M. maniwaata and M. merabet.
680 GIULIO CUCCODORO & IVAN LOBL
Megarthrus nanus sp. n. (Fig. 42a-k)
Type material. Holotype d: Africa or., Katona (= Tanzania, Moshi prov.), Mt. Kiliman-
jaro, BMNH.
Distribution. Tanzania: Mt. Kilimanjaro.
Description. Similar to M. zulu from which it may be distinguished as follows:
Length 1.0 mm; width 0.7 mm. Body dark brown, head blackish, elytron reddish-brown
with darkened sutural margin; legs, mouth parts and antennomeres 1-4 paler. Antenna
as in Fig. 42h. Pronotum as in Fig. 42a. Apical contour of elytron straight near suture.
RatiossE 4: EW IS EN 2.8-2.9: GT 2:0; GW 15-16; PM 8 SP 28229}
d. Protarsal segment | bearing adhesive setae. Mesofemur (Fig. 421) longer than
metafemur (Fig. 42k). Metatibia (Fig. 42f) with peg-like setae arranged in a single row,
and in a double row near apex; shorter than mesotibia (Fig. 42e). Peg-like setae
arranged in a single row on mesotrochanter (Fig. 421), grouped in a field on meta-
trochanter (Fig. 42k) and mesotibia, absent from protibia. Apex of abdominal tergite 8
as in Fig. 42c. Aedeagus as in Fig. 42d, g.
9. Unknown.
Comments. This species and M. zulu are the only African members of the genus
possessing a transverse ridge on the protrochanter. The aedeagal characters are diag-
nostic for each of these two species.
Megarthrus negus sp. n. (Figs 43a-h, 44a-g)
Type material. Holotype 3: Ethiopia, Balé prov., Dinshu, 3200m, 1971 (G. de Rouge-
mont) BMNH.
Paratypes (6): same data as holotype, 14, 19 in BMNH and 16 in MHNG; Balè prov..,
Sabsebe Washa NP <07°03'N: 39°39'E> 3600m, 1971 (R. O. S. Clarke) under stones near stream,
12 in MHNG; Gamo Gofa prov., Gughé highlands, Mt. Tola, c. 3000m, 10-14.xii.1948 (H.
Scott) 16 in BMNH and 1d in MHNG.
Distribution. Southern Ethiopia.
Description. Similar to M. simienensis from which it may be distinguished as
follows: Anterior frontal edge not carinate and submentum weakly convex. Antenna as
in Fig. 44f. Pronotum (Fig. 44g) with deep median groove. Elytron with apical contour
straight or convex near suture, and inner apical angle obtuse. Metasternal median ridge
absent. Ratios: EL 1.6-1.7; ET 1.6-1.7; HW 1.6-1.7; MIE 1-5:SP 275-3: 04 P ES ons
3. Protarsal segment | bearing adhesive setae. Mesofemur (Fig. 43f) as long as
metafemur. Mesotibia (Fig. 43c) with peg-like setae arranged in a double row; shorter
than metatibia (Fig. 43d). Peg-like setae arranged in a single row on mesotrochanter
(Fig. 43f) and metatibia, absent from protibia and metatrochanter. Apex of abdominal
tergite 8 as in Fig. 43e,h. Sternite 8 as in Fig. 43g. Aedeagus as in Fig. 43a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 44b, d. Sternite 8 as
in Fig. 44e. Genital segment as in Fig. 44a, c.
Comments. Megarthrus negus, M. ras, M. rougemonti and M. simienensis share
conspicuous dark colouration of the body, narrowed elytral base and well developed
projection of the female abdominal tergite 8. Megarthrus negus differs notably in
having a single row of metatibial peg-like setae.
AFROTROPICAL MEGARTHRUS 681
Megarthrus niloticus sp. n. (Figs 45a-k, 46a-g)
Type material. Holotype d: Uganda, Bugisu prov., Mt. Elgon, Sipi, 1750m, 31.v.1993
(G. Cuccodoro & D. Erne) ex moist leaf litter at foot of tree in banana plantation, MHNG.
Paratypes (7): Rwanda, Cyangugu pref., around Nyakabuye, 16.x.1984, 1d in ZMHB;
same data, but 3.x.1985, 12 in MHNG; same data, but 31.xii.1985, 19 in ZMHB; Zaire,
Ruwenzori Range (Albert NP) Kyandolire, river Mulaku, tributary of Kakalari, 1750m, 14.x.1952
(P. Vanschuytbroeck & J. Kekenbosch) ex humus, #1241-43, 16 in MHNG and 36 in MRAC.
Distribution. Uganda: Mt. Elgon; Rwanda; Zaire: Ruwenzori Range.
Description. Similar to M. apicicornis from which it may be distinguished as
follows: Length 1.5-1.6 mm; width 0.9-1.1 mm. Pronotum (Fig. 46g) and elytron brown
or dark brown; head, metasternum and abdomen darker than pronotum; sutural margin
of elytron darkened; mouth parts and legs yellowish-brown; antenna reddish-brown,
antennomeres 11 somewhat paler. Eye strongly convex. Antenna as in Fig. 46e. Apical
contour of elytron straight near suture; inner apical angle obtuse. Metasternal median
ridge fine or absent. Ratios: AL 2.0; EL 1.5; ET 1,7; TPF 5.0.
d. Protarsal segment | (Fig. 45f) bearing adhesive setae. Mesofemur (Fig. 45b)
somewhat shorter than metafemur (Fig. 45c). Mesotibia (Fig. 45g) shorter than meta-
tibia (Fig. 45h). Peg-like setae grouped in a field on mesotrochanter (Fig. 45b), meso-
tibia, metatrochanter (Fig. 45c) and metatibia, absent from protibia. Apex of abdominal
tergite 8 as in Fig. 45a, k. Sternite 8 as in Fig. 451. Aedeagus as in Fig. 45d, e.
2. Medioapical projection of abdominal tergite 8 as in Fig. 46a, b. Sternite 8 as
in Fig. 46f. Genital segment as in Fig. 46c, d.
Comments. Megarthrus niloticus differs from other species with the head
widened behind the eyes (see discussions under M. apicicornis and M. congoensis) by
the shape of the male mesotibia in combination with that of the aedeagal tip.
Megarthrus ovalis Cameron (Figs 47a-1, 48a-f)
Megarthrus ovalis CAMERON, 1950: 5.
Type material. Holotype d : Rwanda, V. Visoke (Albert NP) Bishoke, 2800-3300m, 13-
14.11.1935 (G. F. de Witte) #1125, MRAC.
Paratype (9 ): same data as holotype, BMNH.
Additional material (4): Rwanda, S V. Visoke, Virungas, 2950m, 20.11.1978 (I. M.
Redmond) ex dung of Gorilla. g. beringei, 19 in BMNH and 19 in MHNG; Zaire, Kivu (Albert
NP) Tshiaberimu sect., river Mbulikerere, tributary of river Kalivina, 2720m, 25.11.1954 (P.
Vanschuytbroeck & H. Synave) #8295, 1d in MHNG and 1d in MRAC.
Distribution. Rwanda; Zaire: Kivu.
Description. Length 1.35-1.45 mm; width 1.0-1.1 mm. Head blackish; prono-
tum, elytron and abdomen yellowish-brown; metasternum dark brown; sutural margin
of elytron darkened; mouth parts and legs reddish-brown; antenna yellowish-brown,
antennomeres 1-4 somewhat paler. Dorsal pubescence fairly uniform, denser on
humeral area and along sutural margin of elytron. Metasternal setae shorter than those
on prosternum, becoming sparser medioposteriorly and longer anteriorly. Pubescence
of abdominal sternites 4-7 uniform, except for a pair of long setae near medioapical
margins. Puncturation fine on anterior portion of hypomeron; median area of
682 GIULIO CUCCODORO & IVAN LÒBL
metasternum impunctate. Frons raised above level of vertex. Anterior frontal edge not
carinate, evenly arcuate. Entire frontal impression shallow. Eye almost hemispherical,
with highest point below level of vertex. Temple similar to that in Fig. 2g. Submentum
weakly convex. Antenna (Fig. 48d) with patches of sensilla on antennomeres 6-10;
antennomere 3 slightly asymmetrical, 4 strongly asymmetrical. Pronotal disc (Fig. 48f)
deeply depressed along entire lateral edge, flat along median groove; latter deep,
parallel-sided. Anterior prosternal margin not bordered by longitudinal ridges.
Protrochanter without transverse ridge. Lateral portion of prepectal ridge straight.
Scutellum similar to that in Fig. 2b. Elytron abruptly narrowed at base; base abruptly
inclined, overhanging. Elytral disc deeply depressed along lateral edge; latter strongly
convex in dorsal view; apical margin convex near suture; inner apical angle obtuse.
Metasternal median ridge fine, low. Abdominal tergite 3 slightly vaulted transversally.
Sternites 2 and 3 with median processes as in Fig. 21. Sternite 4 almost flat.
Ratios: AL 1.8; EL 1.6-1.7; ET 1.6; EW 1.4; EY 2.9-3.0; GT 2.0; GW 1.6-1.7;
ERWV2223ME1.62MPII 5, PT2.225P4.0-5.02TPR5.0:
d. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 47f) as long as
metafemur (Fig. 47g). Mesotibia (Fig. 47c) as long as metatibia (Fig. 47d), both with
peg-like setae grouped in a field. Metatrochanter (Fig. 47g) with 2-3 peg-like setae.
Peg-like setae arranged in a single row on mesotrochanter (Fig. 47f), absent from
protibia. Apex of abdominal tergite 8 as in Fig. 47e, i. Sternite 8 as in Fig. 47h.
Aedeagus as in Fig. 47a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 48b. Sternite 8 as in
Fig. 48e. Genital segment as in Fig. 48a, c.
Comments. Megarthrus ovalis is characterised by presence of protarsal adhesive
setae in combination with the aedeagus distinctly asymmetrical .
Megarthrus panga sp. n. (Fig, 49a-g; 50a-d)
Type material. Holotype d: Zaire, Kivu prov. (Albert NP) V. Mikeno, near Rweru,
2400m, 26-27.v11.1934 (G. F. de Witte) ex bamboo, #501, BMNH (mislabelled paratype of M.
apicicornis).
Distribution. Zaire: V. Mikeno.
Description. Similar to M. apicicornis from which it may be distinguished as
follows: Length 1.7 mm; width 1.1 mm. Elytral disc concolorous with sutural margin.
Antenna as in Fig. 49d. Pronotum as in Fig. 50d. Elytron with apical contour convex
near suture and inner apical angle obtuse. Metasternal median ridge fine, low. Ratios:
NERO IL El BNW, EX 2.6.2273H1W 2102525:
d. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 49f) as long as
metafemur (Fig. 49g). Mesotibia (Fig. 49a) as long as metatibia (Fig. 49e), with peg-
like setae arranged in a single row, and grouped in a field near apex. Peg-like setae
grouped in a field on mesotrochanter (Fig. 49f), metatrochanter (Fig. 49g) and meta-
tibia, absent from protibia. Abdominal tergite 8 (Fig. 50a, b) with subapical process
projecting ventrally. Sternite 8 as in Fig. 50c. Aedeagus as in Fig. 49b, c.
2. Unknown.
AFROTROPICAL MEGARTHRUS 683
Comments. Megarthrus panga and M. stylifer may be distinguished from other
species with the head widened behind the eyes (see discussion under M. apicicornis) by
the shape of the male abdominal tergite 8. The shape of the aedeagal tip is diagnostic
for each of these two species.
Megarthrus ras sp. n. (Fig. 5la-k)
Type material. Holotype d: Ethiopia, Gonder (= Simén) prov., Arghine, 3500m or
higher, 24.x1.1952 (H. Scott) ex roots of tufted grass in ravine, BMNH (mislabelled paratype of
M. simienensis).
Paratypes (34): same data as holotype, 2 in MHNG (mislabelled paratypes of M.
simienensis); same data, but near torrent, ex peat soil, under boulders or at roots of plants, 1 in
BMNH (mislabelled paratype of M. simienensis).
Distribution. Northern Ethiopia.
Description. Similar to M. simienensis from which it may be distinguished as
follows: Length 1.3-1.4 mm; width 0.9-1.0 mm. Body predominantly blackish, elytron
dark brown with darkened sutural margin; legs and mouth parts yellowish-brown;
antenna reddish-brown, except for paler antennomeres 1-4. Frons raised above level of
vertex. Anterior frontal edge indistinctly carinate; frontal impression shallow. Eye
strongly convex. Submentum weakly convex. Antenna as in Fig. 51c. Pronotum as in
Fig. 51h. Elytron slightly narrowed at base; apical contour convex near suture; inner
apical angle obtuse. Ratios: EL 1.6; HW 1.8; ML 1.5; SP 2.5-3.0.
3. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 51k) as long as
metafemur (Fig. 511). Mesotibia (Fig. 51d) shorter than metatibia (Fig. 5le), with peg-
like setae arranged in a double row. Peg-like setae arranged in a single row on
mesotrochanter (Fig. 51k), grouped in a field on metatrochanter (Fig. 511) and
metatibia, absent from protibia. Apex of abdominal tergite 8 as in Fig. 51f. Sternite 8 as
in Fig. 51g. Aedeagus as in Fig. Sla, b.
2. Unknown.
Comments. Megarthrus ras differs from similar species (see discussion under M.
negus) by the male metatrochanter bearing peg-like setae.
Megarthrus rougemonti sp. n. (Figs 52a-h, 53a-g)
Type material. Holotype d: Ethiopia, Balé prov., Dinshu, 3200m, 1971 (G. de Rouge-
mont) BMNH.
Paratypes (3): same data as holotype, 16, 12 in BMNH; Dinshu, 3200m, 14-28.xi1.1971
(R. O. S. Clarke) under stones in Juniper/Hagenia woodland, 15 in MHNG.
Additional material (19). Ethiopia, Ars? prov., Mt. Ch'ilalo, 3600-4000m, 21.xi.1926 (H.
Scott) Box 113, in moorland, ex decaying stem of Lobelia rhychopetalum Hems., BMNH (para-
lectotype of M. abessinus). Possibly conspecific but the specimen differs by the 8th abdominal
sternite bearing 8 long subapical setae.
Distribution. Southern Ethiopia.
Description. Similar to M. simienensis from which it may be distinguished as
follows: Length 1.45-1.55 mm; width 0.95-1.05 mm. Head, pronotum, metasternum
and abdomen blackish; elytron dark brown; sutural margin of elytron darkened; mouth
684 GIULIO CUCCODORO & IVAN LOBL
parts and legs yellowish-brown; antenna reddish-brown, antennomeres 1-4 somewhat
paler. Pronotal pubescence uniform. Anterior frontal edge not carinate; frontal
impression shallow or indistinct. Frons on level with or slightly raised above level of
vertex. Eye strongly convex. Submentum weakly convex. Antenna as in Fig. 52c.
Pronotal disc (Fig. 53e) shallowly depressed along basal portion of lateral edge; median
groove deep. Elytron slightly narrowed at base; not depressed along lateral edge; inner
apical angle obtuse. Abdominal tergite 3 almost flat. Ratios: EL 1.5-1.6; ET 1.6-1.7;
EM "62 AW ELEMENT PT 192 SPS 0 SMPES 235;
dé. Apical contour of elytron as in Fig. 53g. Protarsal segment 1 bearing
adhesive setae. Mesofemur (Fig. 52f) as long as metafemur. Metatibia (Fig. 52e) longer
than mesotibia (Fig. 52d), with peg-like setae arranged in a single row, and grouped in a
field near apex. Peg-like setae arranged in a single row on mesotrochanter (Fig. 52f),
grouped in a field on mesotibia, absent from protibia and metatrochanter. Apex of
abdominal tergite 8 as in Fig. 52g. Sternite 8 as in Fig. 52h. Aedeagus as in Fig. 52a, b.
2. Apical contour of elytron somewhat convex near suture. Medioapical pro-
jection of abdominal tergite 8 as in Fig. 53c, f. Sternite 8 as in Fig. 53b. Genital
segment as in Fig. 53a, d.
Comments. Megarthrus rougemonti differs from similar species (see discussion
under M. negus) by the short antennomere 5 in combination with the male meta-
trochanter lacking peg-like setae.
The species is dedicated to one of the collectors, Mr Guillaume de Rougemont.
Megarthrus scotti sp. n. (Fig. 54a-h)
Type material. Holotype 2: Ethiopia, Gamo Gofa prov., Gughé highlands, Dita, c.
3000m, 4.xii.1948 (H. Scott) ex humus beneath Knipholia plants, BMNH
Paratypes (29): same data as holotype, in BMNH and MHNG.
Distribution. Southern Ethiopia
Description. Similar to M. clarkei from which it may be distinguished as
follows: Length 1.55-1.65 mm; width 1.05-1.15 mm. Frontal impression deep. Antenna
as in Fig. 54a and pronotum (Fig. 54g) with median groove deep. Ratios: AL 2.0; ET
1.6; GT 2.3; PT 1.8; SP 3.3-4.0; TPF 4.0.
2. Apical contour of elytron and inner apical angle as in Fig. 54h. Medioapical
projection of abdominal tergite 8 as in Fig. 54c, e. Sternite 8 as in Fig. 54f. Genital
segment as in Fig. 54b, d.
3d. Unknown.
Comments. See discussion under M. clarkei.
The species is dedicated to the collector, Mr Hugh Scott.
Megarthrus selenitus sp. n. (Fig. 55a-g)
Type material. Holotype 9: Uganda, Ruwenzori Range, Toro prov., above Kilembe,
2000m, 4.v.1993 (G. Cuccodoro & D. Erne) ex moist vegetational debris near stream in patch of
damaged forest, MHNG.
AFROTROPICAL MEGARTHRUS 685
Distribution. Uganda: Ruwenzori Range.
Description. Similar to M. mukankundiyeorum from which it may be
distinguished as follows: Length 1.6 mm; width 1.1 mm. Antenna as in Fig. 55a.
Elytron with apical contour straight near suture and inner apical angle right-angled.
Metasternal median ridge fine, low. Ratios: EW 1.15; GT 2.2; ML 1.6; PT 1.8; TPF
abs.
2. Abdominal tergite 8 (Fig. 55c, f) lacking medioapical projection, with dark
tip. Sternite 8 (Fig. 55e) 2.2-2.3x as long as width of its median projection. Genital
segment as in Fig. 55b, d.
3. Unknown.
Comments. Among the species possessing a ridged prosternal margin (see
comments under M. africanus), M. selenitus may be distinguished easily by the parti-
cularly elongate abdominal sternite 8.
Megarthrus simienensis Fagel (Figs 56a-g, 57a-g)
Megarthrus simienensis FAGEL, 1957: 30.
Type material. Holotype d: Ethiopia, Gonder (= Simén) prov., E of Mindigabsa, over
3000m, 16.x1.1952 (H. Scott) ex dry soil in roots of tufted plants overhanging a stream, BMNH.
Paratypes (29): same data as holotype, BMNH.
Additional material (23): same data as holotype, 12 in MHNG; Ethiopia, Gonder (=
Simén) prov., Arghine, 3500m or higher, 24.x1.1952 (H. Scott) near torrent, ex peat soil, under
boulders or at roots of plants, 19 in BMNH (mislabelled as paratype); same data, but ex roots of
tufted grass in ravine, 46, 109 in BMNH (mislabelled as paratypes), 25, 19 in MHNG and 29
in MRAC; near Mindigabsa, c. 3500m, 29.xii.1952 (H. Scott) 29 in BMNH (mislabelled as
paratypes).
Distribution. Northern Ethiopia
Description. Length 1.45-1.65 mm; width 1.05-1.20 mm. Head, metasternum
and abdomen blackish; pronotum and elytron dark brown; sutural margin of elytron
darkened; mouth parts and legs yellowish-brown; antenna reddish-brown, antenno-
meres 1-4 somewhat paler. Dorsal pubescence fairly uniform, becoming somewhat
denser along pronotal median groove and on humeral area of elytron. Abdomen with
pubescence denser near posterior margin of tergite 7. Metasternal setae shorter than
those on prosternum, becoming sparser medioposteriorly and longer anteriorly.
Pubescence on abdominal sternites 4-7 uniform, except for a pair of long setae near
medioapical margins. Puncturation fine on anterior portion of hypomeron; median area
of metasternum impunctate. Frons raised above level of vertex. Anterior frontal edge
not carinate, evenly arcuate. Entire frontal impression shallow or indistnct. Eye weakly
convex, with highest point below level of vertex. Temple similar to that in Fig. 2g.
Submentum almost flat. Antenna (Fig. 57c) without patches of sensilla; antennomeres 3
and 4 slightly asymmetrical. Pronotal disc (Fig. 57f) shallowly depressed along entire
lateral edge, flat along median groove; latter shallow, parallel-sided. Anterior prosternal
margin bordered by an irregular row of fine longitudinal ridges. Protrochanter without
transverse ridge. Lateral portion of prepectal ridge straight. Scutellum similar to that in
Fig. 2a. Elytron abruptly narrowed at base; base abruptly inclined, overhanging. Elytral
686 GIULIO CUCCODORO & IVAN LOBL
disc slightly depressed along lateral edge; latter straight in dorsal view; apical margin
somewhat sinuate near suture; inner apical angle right-angled. Metasternal median
ridge fine, low. Abdominal tergite 3 slightly vaulted transversally. Sternites 2 and 3
with median processes as in Fig. 21. Sternite 4 flat at base, then slightly vaulted
transversally.
Ratios AL 1.8; EL 17-18: ET 1.8-2.0; EW 1.25 EY 2:3294AGIZ:0 EMO
IRE WE20 Mile 63 MIP 175 Pay 202 SP' 2.5 PE 33-374
d. Protarsal segment | bearing adhesive setae. Mesofemur (Fig. 56f) as long as
metafemur. Metatibia (Fig. 56e) longer than mesotibia (Fig. 56c), with peg-like setae
arranged in a double row, and grouped in a field near apex. Peg-like setae arranged in a
single row on mesotrochanter (Fig. 56f), grouped in a field on mesotibia, absent from
protibia and metatrochanter. Apex of abdominal tergite 8 as in Fig. 56d. Sternite 8 as in
Fig. 56g. Aedeagus as in Fig. 56a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 57e, g. Sternite 8 as
in Fig. 57d. Genital segment as in Fig. 57a, b.
Comments. Megarthrus simienensis differs from similar species (see discussion
under M. negus) by the pattern of the metatibial peg-like setae and by the antennomere
5 which is as long as antennomere 4.
Megarthrus spinosus sp. n. (Fig. 58a-g)
Type material. Holotype 2: Uganda, Bugisu prov., Mt. Elgon, Sipi, 1750m, 31.v.1993
(G. Cuccodoro & D. Erne) ex moist leaf litter at foot of tree in banana plantation, MHNG.
Paratypes (69): same data as holotype, 1 in BMNH, 3 in MHNG, 1 in MRAC and 1 in
ZMHB.
Distribution. Uganda: Mt. Elgon.
Description. Similar to M. mukankundiyeorum from which it may be distin-
guished as follows: Antenna as in Fig. 58d. Pronotum as in Fig. 58g. Elytron with late-
ral contour somewhat concave and apical contour convex near suture. Ratios: AL 2.2;
HW 1.6-1.7; TPF abs.
2. Abdominal tergite 8 (Fig. 58b, f) lacking medioapical projection, with tip
hyaline. Sternite 8 (Fig. 58e) about 1.8x as long as width of its median projection.
Genital segment as in Fig. 58a, c.
3. Unknown.
Comments. Among the species possessing a ridged prosternal margin (see
comments under M. africanus), M. spinosus may be distinguished easily by the
antennae which are only 2.2x as long as the pronotum.
The water loading behaviour has been recorded in this species (Megarthrus sp.
B in Cuccoporo 1995).
Megarthrus stylifer sp. n. (Figs 59a-g, 60a-k)
Type material. Holotype d: Zaire, Ruwenzori Range (Albert NP) Kyandolire, river
Mulaku, tributary of Kakalari, 1750m, 14.x.1952 (P. Vanschuytbroeck & J. Kekenbosch) ex
humus, #1241-43, MRAC.
Paratypes (5): same data as holotype, 25 in MHNG and 2d, 19 in MRAC.
AFROTROPICAL MEGARTHRUS 687
Distribution. Zaire: Ruwenzori Range.
Description. Similar to M. apicicornis from which it may be distinguished as
follows: Length 1.4-1.5 mm; width 0.9-1.1 mm. Eye strongly convex. Antenna as in
Fig. 60d. Pronotum as in Fig. 601. Metasternal median ridge fine, low. Ratios: AL 2.0;
Pip SSP 2.5; TPF 4.0.
d. Protarsal segment | bearing adhesive setae. Mesofemur (Fig. 60b) shorter
than metafemur (Fig. 60c). Mesotibia (Fig. 59b) shorter than metatibia (Fig. 59e).
Protibia lacking peg-like setae. Mesotrochanter (Fig. 60b), mesotibia, metatrochanter
(Fig. 60c) and metatibia with peg-like setae grouped in a field. Abdominal tergite 8
(Fig. 59c, f, g) with subapical process projecting ventrally. Sternite 8 as in Fig. 60a.
Aedeagus as in Fig. 59a, d.
2. Medioapical projection of abdominal tergite 8 as in Fig. 60g, k. Sternite 8 as
in Fig. 60h. Genital segment as in Fig. 60e, f.
Comments. See discussion under M. panga.
Megarthrus twa sp. n. (Figs 6la-h, 62a-g)
Type material. Holotype d: Kenya, Katamayu river (= Rift Valley prov., Gatamayu
river?) ix.1934 (A. F. J. Gedye) BMNH.
Paratypes (7): same data as holotype, 19 in BMNH and 26 in FMNH; Kenya, Central
prov., Mt. Aberdares NP, Treetops Hotel, 17.v111.1960 (D. H. & A. C. Kistner & R. Banfill) ex
sifted elephant manure, Field No. 618, 22 in BMNH and 19 in MHNG; Zaire, Kivu (Albert NP)
Tshiaberimu sect., river Mbulikerere, tributary of river Kalivina, 2720m, 25.11.1954 (P. Van-
schuytbroeck & H. Synave) #8295, 15 in MRAC.
Distribution. Kenya: Mt. Aberdares; Zaire: Kivu.
Description. Similar to M. zulu from which it may be distinguished as follows:
Length 1.1-1.4 mm; width 0.7-0.9 mm. Antenna (Fig. 62b) dark brown except for
antennomeres 1-4 paler; antennomeres 6-8 with patches of sensilla. Pronotum as Fig.
62f and protrochanter without transverse ridge. Ratios: ET 1.7; GT 2.2; GW 1.8; ML
GAS 5-70: TPE 5.0:
d. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 61f) shorter
than metafemur. Metatibia (Fig. 6le) longer than mesotibia (Fig. 61d), with peg-like
setae arranged in a double row, and grouped in a field near apex. Peg-like setae
arranged in a single row on mesotrochanter (Fig. 61f), grouped in a field on mesotibia,
absent from protibia and metatrochanter. Apex of abdominal tergite 8 as in Fig. 61c, h.
Aedeagus as in Fig. 61a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 62e, g. Sternite 8 as
in Fig. 62a. Genital segment as in Fig. 62c, d.
Comments. Megarthrus twa may be distinguished by the conspicuously dark
colouration of the body in combination with the presence of a particularly narrow pro-
jection of the abdominal sternite 8.
Megarthrus vanschuytbroecki sp. n. (Figs 63a-i, 64a-g)
Type material. Holotype d : Zaire, Ruwenzori Range (Albert NP) Kalonge, river Nyam-
wamba, tributary of Butahu, 2010m, 2-3.11.1953 (P. Vanschuytbroeck & J. Kekenbosch) #2214-
21, MRAC.
688 GIULIO CUCCODORO & IVAN LÒBL
Paratypes (14): same data as holotype, 3d, 19 in MHNG and 5d, 29 in MRAC;
Kalonge (Albert NP) river Katauleko, tributary of Butahu, 2060m, 28.vili.1952 (P. Van-
schuytbroeck & J. Kekenbosch) ex humus, #873, 16 in MRAC; Kalonge (Albert NP) river
Karambura, tributary of Katauleko, 2060m, 30.1.-21.11.1953 (P. Vanschuytbroeck & J. Keken-
bosch) #2225-59, 16 in MRAC; near Kalonge (Albert NP) Kikyo, 2180m, 30.viii.1952 (P.
Vanschuytbroeck & J. Kekenbosch) #670-71, 14 in MRAC.
Distribution. Zaire: Ruwenzori Range.
Description. Similar to M. apicicornis from which it may be distinguished as
follows: Length 1.5-1.8 mm; width 1.00-1.15 mm. Antenna as in Fig. 64c and pro-
notum as in Fig. 64g. Elytron with apical contour somewhat convex near suture and
inner apical angle obtuse. Ratios: AL 2.0-2.2; EL 1.7-1.9; ET 1.6-1.8; ML 1.4; SP 2.8-
3.4; TPF 4.0.
dé. Protarsomere 1 bearing adhesive setae. Mesofemur (Fig. 63e) shorter than
metafemur (Fig. 63a). Mesotibia (Fig. 63g) shorter than metatibia (Fig. 63h), with peg-
like setae grouped in a field. Peg-like setae arranged in a single row on mesotrochanter
(Fig. 63e), arranged in a double row on metatibia, absent from protibia and meta-
trochanter (Fig. 63a). Abdominal tergite 8 as in Fig. 63b, f. Sternite 8 as in Fig. 631.
Aedeagus as in Fig. 63c, d.
2. Medioapical projection of abdominal tergite 8 as in Fig. 64b, e. Sternite 8 as
in Fig. 64a. Genital segment as in Fig. 64d, f.
Comments. This species is easily distinguished from other African congeners by
the hook-shaped aedeagal tip. See also comments under M. apicicornis.
The species 1s dedicated to one of the collectors, Mr Paul Vanschuytbroeck.
Megarthrus watutsi sp. n. (Figs 65a-h, 66a-g)
Type material. Holotype d : Rwanda (Albert NP) at foot of V. Karisimbi, Lake n'Gando,
2400m, 8.111.1935 (G. F. de Witte) #1216, BMNH (paratype of M. apicicornis).
Paratypes (4): same data as holotype, 19 in BMNH (paratype of M. apicicornis); same
data, but 6.111.1935 (G. F. de Witte) #1243, 16 in MHNG (paratype of M. apicicornis); at foot of
V. Karisimbi (Albert NP) Ilega, 2400m, 12.11.1935, #1315, 12 in MRAC (paratype of M.
in MHNG (mislabelled paratype of M. apicicornis).
Distribution. Rwanda: Mt. Tamira and V. Karisimbi.
Description. Similar to M. apicicornis from which it may be distinguished as
follows: Length 1.5-1.6 mm; width 0.9-1.1 mm. Pronotum and elytron brown or dark
brown; head, metasternum and abdomen darker; sutural margin of elytron darkened;
mouth parts and legs yellowish-brown; antenna reddish-brown, antennomere 11 paler.
Frons raised above level of vertex. Frontal impression shallow. Antenna as in Fig. 66d.
Pronotum as in Fig. 66e. Metasternal median ridge fine or absent. Ratios: AL 2.0; EL
SEE RO STPES "0-67
d. Apical contour of elytron somewhat convex near suture; inner apical angle
obtuse. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 65g) as long as
metafemur (Fig. 65h). Mesotibia (Fig. 65e) shorter than metatibia (Fig. 65d), with peg-
like setae arranged in a double row, and grouped in a field near apex. Peg-like setae
grouped in a field on metatrochanter (Fig. 65h), mesotrochanter (Fig. 65g) and
AFROTROPICAL MEGARTHRUS 689
metatibia; absent from protibia. Apex of abdominal tergite 8 as in Fig. 65f. Sternite 8 as
in Fig. 65a. Aedeagus as in Fig. 65b, c.
2. Apical contour of elytron somewhat sinuate near suture; inner apical angle
right-angled. Medioapical projection of abdominal tergite 8 as in Fig. 66b, g. Sternite 8
as in Fig. 66f. Genital segment as in Fig. 66a, c.
Comments. Megarthrus watutsi is one of the species with head widened behind
the eyes (see discussions under M. apicicornis and M. congoensis). It is distinguished
by the male sexual characters, notably abdominal tergite 8 lacking a ventral process,
broad aedeagal tip and shape of the mesotibia.
Megarthrus wittei Cameron (Figs 67a-h, 68a-f)
Megarthrus wittei CAMERON, 1950: 5.
Type material. Holotype d: Zaire, Kivu prov. (Albert NP) Sake, 1960m, 19-22.11.1934
(G. F. de Witte) #253, MRAC.
Paratypes (2): Zaire, Kivu prov. (Albert NP) Kabasha, Kanyabayongo, 1760m, 7.xii.1934
(G. F. de Witte) #877, 16, 12 in BMNH.
Additional material (2): Cameroon, Bamenda distr., Bamenda, 25.1.1957 (V. F. Eastop)
1d in MHNG; Kenya, Rift Valley prov., Molo, 13-15.x.1954 (V. F. Eastop) 46, 52 in BMNH
and 16, 12 in MHNG; Zaire, Oriental prov., Nizi, Blukwa, 23.xii.1928 (A. Collart) 19 in
BMNH.
Distribution. Cameroon: Mt. Bambouto; Kenya: Rift Valley prov.; Zaire: Kivu.
Description. Length 1.2-1.3 mm; width 0.8-1.0 mm. Head blackish; meta-
sternum and abdomen dark brown; pronotal disc and entire elytron reddish-brown;
mouth parts and legs reddish-brown; antenna entirely yellowish. Dorsal pubescence
fairly uniform, denser on humeral area of elytron, shortened on abdomen. Metasternal
setae shorter than those on prosternum, becoming sparser medioposteriorly and longer
anteriorly. Pubescence of abdominal sternites 4-7 uniform, except for a pair of long
setae near medioapical margins. Puncturation fine on anterior portion of hypomeron;
median area of metasternum impunctate. Frons on level with vertex. Anterior frontal
edge not carinate, evenly arcuate. Frontal impression shallow in middle and indistinct
laterally. Eye almost hemispherical, with highest point below level of vertex. Temple
similar to that in Fig. 2h. Submentum weakly convex. Antenna (Fig. 68c) without
patches of sensilla; antennomeres 3 and 4 slightly asymmetrical. Pronotal disc (Fig.
68e) shallowly depressed along basal portion of lateral edge and along median groove;
latter deep, parallel-sided. Anterior prosternal margin bordered with an irregular row of
fine longitudinal ridges. Protrochanter without transverse ridge. Lateral portion of
prepectal ridge straight. Scutellum similar to that in Fig. 2b. Elytron abruptly narrowed
at base; base gradually inclined, overhanging. Elytral disc shallowly depressed along
lateral edge; latter straight in dorsal view; apical margin convex near suture; inner
apical angle obtuse. Metasternal median ridge fine, low. Abdominal tergite 3 slightly
vaulted transversally. Sternites 2 and 3 with median processes as in Fig. 21. Sternite 4
flat at base, then slightly vaulted transversally.
Ratlos Ae 210; EL ies sels eG: EW le2 END 05 0 (Gil 2103 GW 02/0220;
Misi MP STEP" SD SIP 1010)
690 GIULIO CUCCODORO & IVAN LOBL
d. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 67f) longer
than metafemur (Fig. 67h). Metatibia (Fig. 67d) as long as mesotibia (Fig. 67c), with
peg-like setae arranged in a single row, and grouped in a field near apex.
Metatrochanter (Fig. 67h) with 1-3 peg-like setae. Peg-like setae arranged in a double
row on mesotrochanter (Fig. 67f), grouped in a field on mesotibia, absent from protibia.
Apex of abdominal tergite 8 as in Fig. 67g. Sternite 8 as in Fig. 67a. Aedeagus as in
Fig. 67b, e.
2. Medioapical projection of abdominal tergite 8 as in Fig. 68f. Sternite 8 as in
Fig. 68d. Genital segment as in Fig. 68a, b.
Comments. See discussion under M. falasha.
Megarthrus zulu sp. n. (Figs 69a-h, 70a-g)
Type material. Holotype d : Republic of South Africa, Cape prov. S, Marathon <34°02'S;
23°19'E> 9.x11.1976 (S. Endròdy-Younga) ex cattle and horse dung, E-Y:1306, TMSA.
Paratypes (411): same data as holotype, 1d, 12 in MHNG and 26, 19 in TMSA;
Republic of South Africa, Cape prov. S, Harkeville Forest <34°04'S; 23°10'E> 7.xi1.1976 (S.
Endrödy-Younga) ex elephant dung, E-Y:1300, 45,42 in MHNG and 126, 49 in TMSA; same
data, but 13.xii.1976, groundtrap with meat bait, E-Y:1311, 18 in MHNG and 16 in TMSA;
Keurboomstrand <34°00'S; 23°27'E> 8.x11.1976 (S. Endrödy-Younga) groundtraps, 8 days, E-
Y:1301, 15 in TMSA; Cape prov. E, Katberg, 1300m, 1-15.1.1933 (R. E. Turner) 19 in BMNH;
Cape prov., Amatole, Isidenge For. St. <32°41'S; 27°15'E> 1.x11.1986 (S. Endrödy-Younga)
beating in forest, E-Y:2337, 16 in TMSA; same data, but 19.xi.1987, ex horse dung, E-Y:2527,
36,39 in MHNG, 16,1% in MRAC and 17d, 119 in TMSA; same data, but ex Quercus forest
litter, E-Y:2517, 16, 39 in TMSA; same data, but Isidenge For. St. Bl <32°41'S; 27°14'E>
15.xi.1987, ex Pinus bark, E-Y:2516, 16, 19 in TMSA; George, Kranshoek, 23.xii.1981 (S.
Peck) ex forest litter under carrion, FMHD #81-728, 16 in MHNG and 26, 19 in FMNH;
George, Gouna, 23.xii.1981 (S. Peck) ex forest log litter, berlese, FMHD #81-729, 29 in FMNH;
George, Saasveld, 26.x11.1981 (S. Peck) ex forest litter under carrion bait, berlese, FMHD #81-
731, 12, FMNH; Groenkop <33°57'S; 22°33'E> i.1985 (J. Koen) 16 in TMSA; Knysna,
Buffelsnek, 25.xii.1981 (S. Peck) ex forest litter under bait trap, berlese, FMHD #81-633, 2d,
22 in FMNH and 26, 19 in MHNG; same data, but 800m (S. & J. Peck) ex forest litter under
carrion baits, fynbos, #81-187b, 5d, 32 in FMNH; Knysna, Diepwalle, 450m, 12-30.x11.1981 (S.
& J. Peck) ex mixed Podocarpus forest, window-malaise trap, 26, 22 in FMNH and 16,19 in
MHNG; same data, but (S. Peck) ex forest, malaise trough, FMHD #81-711, 18, 22 in FMNH
and 16, 22 in MHNG; same data, but 12.xii.1981, ex elephant dung, berlese, FMHD #81-603,
306, 202 in FMNH, 46 , 49 in MHNG and 1g, 19 in MRAC; same data, but 23.xii.1981, ex
forest litter, berlese, FMHD #81-629, 28,22 in MHNG and 26, 39 in FMNH; same data, but
17.x11.1981, ex forest fungi litter, berlese, FMHD #81-721, 19 in FMNH; Plettenberg Bay
Natures Vly., 29.x11.1981 (S. Peck) ex forest litter under dung, berlese, FMHD #81-641, 3d, 1?
in FMNH and 24, 19 in MHNG; Stormsrivier, Goesabos, 15-30.x11.1981 (S. Peck) ex forest,
malaise trough, FMHD #81-704, 12 in FMNH; Natal prov. Middland, Doring Clark Nat. Res.
<29°34'S; 30°17'E> 11.x11.1989 (S. Endrödy-Younga & J. Klimaszewsk1) ex river bank in forest,
E-Y:2759, 16 in TMSA; Karkloof for. <29°18'S; 30°13'E> 11.x11.1989 (S. Endrödy-Younga & J.
Klimaszewski) ex horse dung, E-Y:2762, 1% in TMSA; Northington <29°28'S; 30°01'E> 1420m,
12.xii.1989 (S. Endròdy-Younga & J. Klimaszewski) ex sifted forest litter, E-Y:2764, 26,29 in
MHNG and 36, 52 in TMSA; Natal prov., 75km WSW Estcourt, Cathedral Pks. For. Sta.,
1400m, 26.xii.1979 (S. & J. Peck) ex Eucalyptus logs, frass, fungi and decaying bark, Ber 22, 19
in FMNH; same data, but berlese residue, FMHD 79-189, 16 in FMNH; same data, but 7-
31.x11.1979, 46, 172 in FMNH and 38,3% in MHNG; same data, but 2000m, 20.x11.1979 (S. &
J. Peck) ex rotted Boletus litter in pine plantation, Ber 14, 14 in CNCI, 259, 342 in FMNH and
14,19 in MHNG; same data, but B #14, FMHD #79-195, 16 in FMNH: same data, but berlese
AFROTROPICAL MEGARTHRUS 691
residue, FMHD #79-195, 74, 112 in FMNH, 24,29 in MHNG and 1d, 19 in MRAC; same
data, but 1500m, 19.xii.1979, ex rotted Boletus bait station in podocarp forest, Ber 11, 126, 129
in FMNH and 26,29 in MHNG; same data, but 31.xi1.1979, ex podocarp forest, mini-carrion
bait traps, Ber 32, 1d in FMNH; same data, but ex rotted Boletus bait, 36, 79 in SEMC; same
data, but Catchment 2, 1760m, 21-31.xii.1979, ex stream side shrubs, malaise trap, 1d, 19 in
FMNH; same data, but Rainbow Gorge, 2000m, 8-31.x11.1979, ex Podocarp forest, malaise
through, 16, 19 in FMNH; Weza Forest Station <30°36'S; 29°45'E> 22.x1.1989 (S. Endrödy-
Younga & J. Klimaszewski) ex horse dung, E-Y:2711, 14,29 in TMSA; Pietermaritzburg, Fort
Napier, 1919, #127, eig. Nr. 36,1926, 19 in FMNH; Weza, Bangeni forest <30°38'S; 29°39'E>
21.xi.1989 (S. Endrödy-Younga & J. Klimaszewski) ex sifted forest litter, E-Y:2706, 1d, in
MHNG and 2¢ in TMSA; Weza, Ingeni forest <30°32'S; 29°41'E> 18.x1.1989 (S. Endrödy-
Younga & J. Klimaszewski) ex forest floor litter, E-Y:2689, 12 in TMSA; same data, but ex
bushbuck excrement, E-Y:2690, 16 in MHNG and 1d, 19 in TMSA; same data, but 23.x1.1989,
ex sifted grass in forest, E-Y:2714, 16 in TMSA; Weza, Iower Stinkwood for. <30°34'S;
29°43'E> 17.x1.1989 (S. Endrödy-Younga & J. Klimaszewski) ex sifted forest litter, E-Y:2686,
16 in TMSA; Transkei prov., Port St. Johns, Pondoland, 10-31.vii.1923 (R. E. Turner) 1d in
BMNH; Transvaal prov. E, Berlin For. St. <25°33'S; 30°44'E> 9.x11.1986 (S. Endròdy-Younga)
ex horse dung, E-Y:2370, 16 in MHNG and 24 in TMSA; same data, but gorge <25°32'S;
30°44'E> 21.ix.1986, groundtraps, 32 days, E-Y:2281, 19 in MHNG and 14, 29 in TMSA;
same data, but 23.x.1986, groundtraps, 42 days, E-Y:2303, 1d in TMSA; same data, but gorge-
edge <25°32'S; 30°44'E> 5.11.1987, ex mule dung, E-Y:2418, 16 in TMSA; same data, but
Sinkhole <25°31'S; 30°46'E> 20.1x.1986 (S. Endrödy-Younga) groundtraps, 33 days, E-Y:2276,
14,19 in TMSA; same data, but 23.x.1986, groundtraps, 42 days, E-Y:2300, 14 in MHNG and
26, 12 in TMSA; Blyderiver Canyon <24°35'S; 30°49'E> 5.v.1981 (S. Endrödy-Younga)
groundtraps with faeces bait, 5 days, E-Y:1781, 168 in MHNG and 1d in TMSA; 30km W
Trichardtsdal, 23-30.x11.1985 (S. Peck) downs podocarp forest, carrion trap, P#85-307, FMHD
#85-874, 12 in FMNH; Pilgrims rest, 19-31.xii.1985 (S. Peck) ex relict nature forest dung trap,
P#85-293, FMHD #85-860, 16, 22 in FMNH and Id, 12 in MHNG; Transvaal prov.,
Pietersburg dist., Helmekaar river, 23-30.ix.1961 (N. Leleup) ex humus, 1% in TMSA; Uitsoek,
Grootkloof ind. forets <25°15'S; 30°33'E> 15.xii.1986 (S. Endrödy-Younga) groundtraps, 53
days, E-Y:2391, 16 in TMSA; same data, but 17.x11.1986, ex forest litter after rain, E-Y:2396,
1d in TMSA; Lesotho E, Sani Pass Valley, Drakensberg <29°39'S; 29°12'E> 8.iii.1976 (S.
Endrödy-Younga & Breytenb.) ex dung collection, E-Y:1054, 26 in MHNG and 24, 29 in
TMSA; same data, but (S. Endrödy-Younga) ex horse and cattle dung, E-Y:1054, 12 in MHNG
and 12 in TMSA; Drakensberg, Vann Reenen <28°22'S; 29°23'E> xi.1926 (R. E. Turner) 1¢ in
BMNH.
Distribution. Lesotho; Republic of South Africa.
Description. Length 0.9-1.3 mm; width 0.6-0.8 mm. Body usually entirely dark
brown or blackish, with paler legs and mouth parts; pronotum and elytral disc
sometimes paler. Dorsal pubescence fairly uniform, sparser on elytron, becoming
denser near apical margin of abdominal tergite 7. Metasternal setae uniform, shorter
than those on prosternum. Pubescence of abdominal sternites 4-7 uniform, except for a
pair of long setae near medioapical margins. Puncturation fine on anterior portion of
hypomeron; median area of metasternum impunctate. Frons slightly raised above level
of vertex. Anterior frontal edge not carinate, evenly arcuate. Entire frontal impression
Shallow or indistinct. Eye strongly convex, with highest point below level of vertex.
Temple similar to that in Fig. 2g. Submentum weakly convex. Antenna (Fig. 70a)
without patches of sensilla; antennomere 3 slightly asymmetrical, 4 strongly asym-
metrical. Pronotal disc (Fig. 70f) flat along lateral edge, deeply depressed along median
groove latter deep, parallel-sided. Anterior prosternal margin not bordered by
longitudinal ridges. Protrochanter with transverse ridge. Lateral portion of prepectal
692 GIULIO CUCCODORO & IVAN LOBL
ridge straight. Scutellum similar to that in Fig. 2b. Elytron slightly narrowed at base;
base abruptly inclined, overhanging. Elytral disc flat along lateral edge; latter straight in
dorsal view; apical margin straight or convex near suture; inner apical angle obtuse.
Metasternal median ridge fine, low. Abdominal tergite 3 slightly vaulted transversally.
Sternites 2 and 3 with median processes as in Fig. 21. Sternite 4 flat at base, then
slightly vaulted transversally.
Ratios AE 1.72 EE 1-5-1627 Et 1.55-1.65: EW 1.2: EY 2.5; GIZ4IIEGWALSIE
HW 1.6-1.7; ML 1.4; MP 1.7; PT 1.8-1.9; SPS 2.5-3.0; SP9 3.3-3.5; TPF 6.5-10.0.
d. Protarsal segment 1 bearing adhesive setae. Mesofemur (Fig. 69f) longer
than metafemur (Fig. 69d). Mesotibia (Fig. 69c) as long as metatibia (Fig. 69e), with
peg-like setae arranged in a double row, and grouped in a field near apex. Peg-like setae
arranged in a single row on mesotrochanter (Fig. 69f), metatrochanter (Fig. 69d) and
metatibia; absent from protibia. Apex of abdominal tergite 8 as in Fig. 69h. Sternite 8 as
in Fig. 69g. Aedeagus as in Fig. 69a, b.
2. Medioapical projection of abdominal tergite 8 as in Fig. 70e, g. Sternite 8 as
in Fig. 70d. Genital segment as in Fig. 70b, c.
Comments. See discussion under M. nanus.
DISCUSSION
The Afrotropical Megarthrus occur in three geographically separated areas (Fig.
1). Equatorial East Africa, especially the mountainous areas near Lake Victoria, has 27
species (71%). A second area with relatively high species diversity (9 species) is in
Ethiopia. None of the latter species has been found outside of Ethiopia and none of the
former occurs in Ethiopia. With two species from Cameroon and one species in the
Republic of South Africa, the genus is depauperate in West and South Africa. The
genus has not been found so far in Madagascar and Mascarene Islands.
The altitudinal records indicate highest diversity of the Equatorial East African
Megarthrus between 1700-2700m (19 species); two species are present only above
2900m and three species at 900m; none below 900m. The Ethiopian species appear to
be confined to higher altitudes, between 2700-4000m. The single South African spe-
cies, M. zulu, has been found between 400-2000m.
ACKNOWLEDGMENTS
The present paper is part of a joint project between the Muséum d'histoire
naturelle, Geneva and the Natural History Museum, London, funded by the Swiss
National Science Foundation (Project no. 31-32331.91, awarded to I. Löbl, D.
Burckhardt, MHNG, and P. M. Hammond, BMNH). The following colleagues have
provided specimens for the present study: H. M. André, MRAC; J. S. Ashe, SEMC; S.
Endrédy-Younga, TMSA; A. F. Newton Jr., FMNH; A. Smetana, CNCI; M. Uhlig,
ZMHB. Mrs D. Cuccodoro assisted with the inking of drawings, and D. Erne, Geneva,
made the field work in Uganda more pleasant. The field work in Uganda was supported
by a travelling grant of the "Académie suisse des sciences naturelles”.
AFROTROPICAL MEGARTHRUS 693
FIG. 3
Megarthrus abessinus, male; a, b: aedeagus, ventral and lateral; c: abdominal sternite 8; d:
protibia and protarsomere 1; e: mesotibia; f: metatibia; g: mesofemur and mesotrochanter; h:
metafemur and metatrochanter; i: apex of abdominal tergite 8, dorsal. Scale bar = 0.2 mm.
694 GIULIO CUCCODORO & IVAN LÒBL
FIG. 4
Megarthrus abessinus; a: pronotum; b, d: female, genital segment, tergites (b) and sternites (d); c:
female, abdominal sternite 8; e, f: female, apex of abdominal tergite 8, lateral and dorsal; g:
antenna. Scale bar = 0.2 mm.
AFROTROPICAL MEGARTHRUS 695
FIG. 5
Megarthrus africanus; a: pronotum; b: male, mesotibia; c: antenna (antennomere 7 deformed); d:
male, abdominal sternite 8; e, f: aedeagus, ventral and lateral; g: male, apex of abdominal tergite
8, dorsal: h: male, mesotrochanter. Scale bars = 0.2 mm.
696 GIULIO CUCCODORO & IVAN LOBL
a,b,e-g
Fic. 6
Megarthrus apicicornis; a, b: female, genital segment, sternites (a) dorsal and tergites (b); c:
antenna; d: female, abdominal sternite 8; e, g: female, apex of abdominal tergite 8, dorsal and
lateral; f: pronotum. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 697
(0)
_DROQ>
ESS
TE
i
NR, f |
LARE NI |
ify A ind
J Ge paral |S LA n \ )
] ur 11 ft u dh |
Mi PISTE
I 1p, AN À 417 /
\ A AE oe
\ an a 10 = x /
\ ~— =~ M —
1 SM /
\ / ; i
N / I
x 2 y
x A
Te =
Fic. 7
Megarthrus bantu, male; a, b: aedeagus, lateral and ventral; c: mesotibia; d: metatibia; e, i: apex
of abdominal tergite 8, lateral and dorsal; f: abdominal sternite 8; g: mesofemur and meso-
trochanter; h: metafemur and metatrochanter. Scale bars = 0.2 mm.
698 GIULIO CUCCODORO & IVAN LOBL
Fic. 8
Megarthrus bantu; a, b: female, genital segment, sternites (a) and tergites (b); c: antenna; d:
female, abdominal sternite 8; e, g: female, apex of abdominal tergite 8, lateral and dorsal; f: pro-
notum. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 699
Fic. 9
Megarthrus basilewskyi, male; a: metafemur and metatrochanter; b, d: aedeagus, ventral and late-
ral; c: mesofemur and mesotrochanter; e: metatibia; f: mesotibia; g: protibia. Scale bar = 0.2 mm.
700 GIULIO CUCCODORO & IVAN LOBL
Fic. 10
Megarthrus basilewskyi; a: pronotum; b: female, apical contour of elytra; c: antenna; d, e: male,
abdominal tergite 8, entire (e) lateral and apex (d) dorsal; f, g: female, abdominal tergite 8, dorsal
and lateral. Scale bar = 0.2 mm.
AFROTROPICAL MEGARTHRUS 701
Fic. 11
Megarthrus basilewskyi; a-c: female, genital segment, sternites (a) dorsal, (b) lateral and tergites
(c); d, e: abdominal sternite 8, female and male. Scale bars = 0.2 mm.
702 GIULIO CUCCODORO & IVAN LÒBL
Fic. 12
Megarthrus clarkei, male; a, b: aedeagus, ventral and lateral; c: mesotibia; d: metatibia; e:
mesofemur and mesotrochanter; f, h: apex of abdominal tergite 8, dorsal and lateral; g: abdominal
sternite 8. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 703
Fic. 13
Megarthrus clarkei; a: pronotum; b: antenna; c, f: female, apex of abdominal tergite 8, dorsal
and lateral; d, e: female, genital segment, sternites (d) and tergites (e); g: female, abdominal
sternite 8. Scale bars = 0.2 mm.
704 GIULIO CUCCODORO & IVAN LÒBL
Fic. 14
Megarthrus congoensis, male; a-e: apex of abdominal tergite 8, lateral and dorsal; b: abdominal
sternite 8; c, d: aedeagus, ventral and lateral; f: mesotibia; g: metatibia; h: mesofemur and
©
mesotrochanter; i: metafemur and metatrochanter. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 705
Fic. 15
Megarthrus congoensis; a: antenna; b, e: female, apex of abdominal tergite 8, lateral and dorsal;
c: pronotum; d, f: female, genital segment, sternites (d) and tergites (f); g: female, abdominal
sternite 8. Scale bars = 0.2 mm.
706 GIULIO CUCCODORO & IVAN LOBL
TR III I
Lo N
u.
j
[
Fic. 16
Megarthrus dominicae; a: antenna; b, c: aedeagus, ventral and lateral; d: male, mesotibia; e: male,
mesofemur and mesotrochanter; f, h: male, apex of abdominal tergite 8, lateral and dorsal; g:
male, metatrochanter; i: male, abdominal sternite 8. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 707
Fic. 17
Megarthrus dominicae; a: pronotum; b, c: female, apex of abdominal tergite 8, lateral and dorsal;
d, e: female, genital segment, tergites (d) and sternites (e); f: female, abdominal sternite 8. Scale
bars = 0.2 mm.
708 GIULIO CUCCODORO & IVAN LOBL
’
4
y
h
A
|
Fic. 18
Megarthrus falasha, male; a, b: aedeagus, ventral and lateral; c: mesotibia; d: metatibia; e, g:
apex of abdominal tergite 8, dorsal and lateral; f: mesofemur and mesotrochanter; h: meta-
trochanter; i: abdominal sternite 8 . Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 709
Ny
un
(O) | UE dn Neon
NN er
ar IN iR
Fic. 19
Megarthrus falasha; a: antenna; b: pronotum; c, e: female, genital segment, tergites (c) and
sternites (e); d, f: female, apex of abdominal tergite 8, lateral and dorsal; g: female, abdominal
sternite 8. Scale bars = 0.2 mm.
710 GIULIO CUCCODORO & IVAN LÒBL
Fic. 20
Megarthrus gigas, male; a: metafemur and metatrochanter; b, c: aedeagus, lateral and ventral; d:
mesofemur and mesotrochanter; e, f: apex of abdominal tergite 8, lateral and dorsal. Scale bar =
0.2 mm.
AFROTROPICAL MEGARTHRUS 711
Fic. 21
Megarthrus gigas; a: pronotum; b: female, apical contour of elytra; c: antenna; d, e: abdominal
sternite 8, male (d) and female (e). Scale bar = 0.2 mm.
712 GIULIO CUCCODORO & IVAN LÔBL
N
III, ID
oS
=
SS
=
III
SS
> —_
Ff
=
ra —
ZA —=
TWYTW_WSTS
x
LS
47) \
S II
FIG. 22
Megarthrus gigas; a-d: female, genital segment, sternites (a) dorsal, (b) lateral, (d) posterior and
tergites (c) ventral; e: male, metatibia; f: male, mesotibia; g: male, protibia and protarsomere 1;
h, i: female, apex of abdominal tergite 8, lateral and dorsal. Scale bar = 0.2 mm.
AFROTROPICAL MEGARTHRUS 713
À
samba ds
à 4
DI à D
SALI Ar y
4
Fic. 23
Megarthrus horticola, male; a, b: aedeagus, ventral and lateral; c: mesotibia; d, e: apex of
abdominal tergite 8, lateral and dorsal; f: mesofemur and mesotrochanter; g: abdominal sternite 8;
h: metatrochanter. Scale bars = 0.2 mm.
714 GIULIO CUCCODORO & IVAN LOBL
FIG. 24
Megarthrus horticola; a: antenna; b c: female, genital segment, sternites (b) and tergites (c); d: fe-
male, abdominal sternite 8; e, g: apex of abdominal tergite 8, dorsal and lateral; f: pronotum.
Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 715
Fic. 25
Megarthrus hutu, male; a, b: aedeagus, ventral and lateral; c: mesotibia; d: metatibia; e:
mesofemur and mesotrochanter; f, g: apex of abdominal tergite 8, dorsal and lateral; h:
metafemur and metatrochanter. Scale bars = 0.2 mm.
716 GIULIO CUCCODORO & IVAN LOBL
FIG. 26
Megarthrus hutu; a: antenna; b, d: female, apex of abdominal tergite 8, dorsal and lateral; c, f:
abdominal sternite 8, female (c) and male (f); e, g: female, genital segment, sternites (e)
and tergites (g); h: pronotum. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS ANG
N }
N
A
À Ih
AN
Ni
y \\ hy
IN 1)
\ VA.
| \ WAR.
Ex N Se RR, RI, \ | à
NS I PR aw) Zi\\ À
SS ye \ SEZIONI i W i
SR Ti ue ZERI = yt |
eS SCE tines ur
|
Fic. 27
Megarthrus magnicaudatus; a: male, abdominal sternite 8; b, k: male, abdominal tergite 8, entire
(b) lateral and apex (k) dorsal; c: male, mesofemur and mesotrochanter; d, e: aedeagus, lateral
and ventral (apex broken); f: male, protibia and protarsomere 1; g: male, mesotibia; h: male,
metatibia; i: pronotum. Scale bar = 0.2 mm.
718 GIULIO CUCCODORO & IVAN LOBL
FIG. 28
Megarthrus magnicaudatus; a, d: female, genital segment, tergites (a) and sternites (d); b, f:
female, abdominal tergite 8, dorsal and lateral; c: antenna; e: female, abdominal sternite 8. Scale
bar = 0.2 mm.
AFROTROPICAL MEGARTHRUS 719
Va
//
/
en x E
a
x >
—
=
SSS SO =
i — m
Fic. 29
Megarthrus mahnerti; a: male, metatibia; b: male, mesofemur and mesotrochanter; c: male, apex
of abdominal tergite 8, dorsal; d; antenna; e, f: aedeagus, lateral and ventral; g: male, mesotibia;
h: male, metatrochanter; 1: male, abdominal sternite 8. Scale bars = 0.2 mm.
720 GIULIO CUCCODORO & IVAN LOBL
Fic. 30
Megarthrus mahnerti; a, c: female, genital segment, tergites (a) and sternites (c); b: female, abdo-
minal sternite 8; d, e: female, apex of abdominal tergite 8, lateral and dorsal; f: pronotum; g:
female, apical contour of elytra. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 72
Fic. 31
Megarthrus major; a, b, d, f: female, genital segment, sternites, lateral (a), dorsal (b), posterior (f)
and tergites (d); c, g: female, apex of abdominal tergite 8, dorsal and lateral; e: antenna; h: lateral
and basal edges of pronotum; i: female, abdominal sternite 8; k: female, apical contour of
elytra. Scale bar = 0.2 mm.
GIULIO CUCCODORO & IVAN LOBL
/
SS
po Se
=>
Fic. 32
Megarthrus maniwaata, male; a, b: aedeagus, ventral and lateral; c: mesotibia; d: mesofemur and
mesotrochanter; e: metatibia; f: metafemur and metatrochanter; g: abdominal sternite 8; h: apex
of abdominal tergite 8, dorsal view. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 723)
Fic. 33
Megarthrus maniwaata; a, b: female, genital segment, sternites (a) and tergites (b); c: antenna; d:
female, abdominal sternite 8; e, g: female, apex of abdominal tergite 8, lateral and dorsal; f:
pronotum. Scale bars = 0.2 mm.
724 GIULIO CUCCODORO & IVAN LÒBL
NNO
=
e
=O —
>
Fn
sae
a
eo.
>=
e D
N
NS
SS SSE
Fic. 34
Megarthrus merabet, male; a, b: aedeagus, ventral and lateral; c: mesotibia; d: metatibia; e:
mesofemur and mesotrochanter; f: metafemur and metatrochanter; g: abdominal sternite 8; h:
apex of abdominal tergite 8. Scale bar = 0.2 mm.
AFROTROPICAL MEGARTHRUS 725
Fic. 35
Megarthrus merabet; a, b: female, genital segment, sternites (a) and tergites (b); c: antenna; d:
female, abdominal sternite 8; e, g: female, apex of abdominal tergite 8, lateral and dorsal; f: pro-
notum. Scale bars = 0.2 mm.
126 GIULIO CUCCODORO & IVAN LÔBL
S
SS
N
SI
N
/
f
N
h
È
À
Fic. 36
Megarthrus monticola, male; a, i: apex of abdominal tergite 8, dorsal and lateral; b: abdominal
sternite 8; c, d: aedeagus, ventral and lateral; e: mesotibia; f: metatibia; g: mesofemur and
mesotrochanter; h: metafemur and metatrochanter. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS OG]
Fic. 37
Megarthrus monticola; a: antenna; b: female, abdominal sternite 8; c, g: female, apex of abdo-
minal tergite 8, lateral and dorsal; d: pronotum; e: female, genital segment, tergites (e) and
sternites (f); h: female, apical contour of elytra. Scale bars = 0.2 mm.
728 GIULIO CUCCODORO & IVAN LOBL
IYSs SON
= _——r-- "="
L z Bo; A TS
je — SS
T7
ver eat ai]
Nino, pai
ZA UD
he LE 6
Vie LA
SAQVWAIWdNH
Fic. 38
Megarthrus mukankundiyeorum, male; a, b: aedeagus, ventral and lateral; c, d: apex of abdominal
tergite 8, lateral and dorsal; e: metatrochanter; f: mesotibia; g: metatibia; h: mesofemur and
mesotrochanter; i: abdominal sternite 8. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 729
Fic. 39
Megarthrus mukankundiyeorum; a, b: female, genital segment, tergites (a) and sternites (b); c, d:
female, apex of abdominal tergite 8, dorsal and lateral; e: antenna; f: female, abdominal sternite
8; g: pronotum. Scale bars = 0.2 mm.
730 GIULIO CUCCODORO & IVAN LÒBL
S
NS
SS
= SS
—
- ==
ND SKK
ET
=
3
>>
A
Fic. 40
Megarthrus mwami, male; a, b: aedeagus, ventral and lateral; c: mesotibia; d: metatibia; e, i: apex
of abdominal tergite 8, lateral and dorsal; f: abdominal sternite 8; g: mesofemur and meso-
trochanter; h: metafemur and metatrochanter. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 731
Fic. 41
Megarthrus mwami; a: antenna; b, c: female, genital segment, tergites (b) and sternites (c); d, g:
female, apex of abdominal tergite 8, dorsal and lateral; e: female, abdominal sternite 8; f: pro-
notum. Scale bars = 0.2 mm.
782 GIULIO CUCCODORO & IVAN LÒBL
b,c,e,f,h-k
d,g
Fic. 42
Megarthrus nanus; a: lateral and basal edges of pronotum; b: male, abdominal sternite 8; c: male,
apex of abdominal tergite 8, dorsal; d, g: aedeagus, lateral and ventral; e: male, mesotibia; f:
male, metatibia; h: antenna; 1: male, mesofemur and mesotrochanter; k: male, metafemur and
metatrochanter. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 733
FIG. 43
Megarthrus negus, male; a, b: aedeagus, ventral and lateral; c: mesotibia; d: metatibia; e, h: apex
of abdominal tergite 8, dorsal and lateral; f: mesofemur and mesotrochanter; g: abdominal sternite
8. Scale bar = 0.2 mm.
734 GIULIO CUCCODORO & IVAN LOBL
Fic. 44
Megarthrus negus; a, c: female, genital segment, sternites (a) and tergites (c); b, d: female, apex
of abdominal tergite 8, lateral and dorsal; e: female, abdominal sternite 8; f: antenna; g: pro-
notum. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 735
TT
NS
DX
~~
CHE
Se
Par
===
SS
90099 9% G
b,c,f-h
a,d,e,i,k
Fic. 45
Megarthrus niloticus, male; a, k: apex of abdominal tergite 8, dorsal and lateral; b: mesofemur
and mesotrochanter; c: metafemur and metatrochanter; d, e: aedeagus, ventral and lateral; f:
protarsi; g: mesotibia; h: metatibia; 1: abdominal sternite 8. Scale bars = 0.2 mm.
736 GIULIO CUCCODORO & IVAN LOBL
c,d,g ————_
a,b,e,f
Fic. 46
Megarthrus niloticus; a, b: female, apex of abdominal tergite 8, dorsal and lateral; c, d: female,
genital segment, sternites (c) and tergites (d); e: antenna; f: female, abdominal sternite 8; g:
pronotum. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 737
Fic. 47
Megarthrus ovalis, male; a, b: aedeagus, ventral and lateral; c: mesotibia; d: metatibia; e, i: apex
of abdominal tergite 8, dorsal and lateral; f: mesofemur and mesotrochanter; g: metafemur and
metatrochanter; h: abdominal sternite 8. Scale bars = 0.2 mm.
GIULIO CUCCODORO & IVAN LOBL
738
Fic. 48
Megarthrus ovalis; a, c: female, genital segment, tergites (a) and sternites (c); b: female, apex of
abdominal tergite 8, dorsal; d: antenna; e: female, abdominal sternite 8; f: pronotum. Scale bars =
0.2 mm.
AFROTROPICAL MEGARTHRUS 739
7
INN
>,
=
Fic. 49
Megarthrus panga; a; male, mesotibia; b, c: aedeagus, ventral and lateral; d: antenna; e: male,
metatibia; f: mesofemur and mesotrochanter; g: metafemur and metatrochanter. Scale bar = 0.2
mm.
740 GIULIO CUCCODORO & IVAN LÒBL
Fic. 50
Megarthrus panga; a, b: male, abdominal tergite 8, ventral and lateral; c: male, abdominal ster-
nite 8; d: pronotum. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 741
Fic. 51
Megarthrus ras; a-b: aedeagus, ventral and lateral; c: antenna; d: male, mesotibia; e: male, meta-
tibia; f: male, apex of abdominal tergite 8, dorsal; g: male, abdominal sternite 8; h: lateral and
basal edges of pronotum; i: male, metafemur and metatrochanter; k: male, mesofemur and meso-
trochanter. Scale bars = 0.2 mm.
742 GIULIO CUCCODORO & IVAN LOBL
FIG. 52
Megarthrus rougemonti; a, b: aedeagus, ventral and lateral; c: antenna; d: male, mesotibia; e:
male, metatibia; f: male, mesofemur and mesotrochanter; g: male, apex of abdominal tergite 8,
dorsal; h: male, abdominal sternite 8. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 743
®
h
Fic. 53
Megarthrus rougemonti; a, d: female, genital segment, sternites (a) and tergites (d), basal portion
broken; b: female, abdominal sternite 8; c, f: female, apex of abdominal tergite 8, dorsal and
lateral; e: pronotum; g; male, apical contour of elytra. Scale bars = 0.2 mm.
744 GIULIO CUCCODORO & IVAN LÔBL
Fic. 54
Megarthrus scotti; a: antenna; b, d: female, genital segment, tergites (b) and sternites (d); c, e:
female, apex of abdominal tergite 8, dorsal and lateral; f: female, abdominal sternite 8; g: pro-
notum; h: female, apical contour of elytra. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 745
Fic. 55
Megarthrus selenitus; a: antenna; b, d: female, genital segment, sternites (b) and tergites (d); c, f:
female, apex of abdominal tergite 8, dorsal and lateral; e: female, abdominal sternite 8; g: pro-
notum. Scale bars = 0.2 mm.
746 GIULIO CUCCODORO & IVAN LÒBL
N
(e)
SK
x
SSS.
SS
Fic. 56
Megarthrus simienensis, male; a, b: aedeagus, ventral and lateral; c: mesotibia; d: apex of abdo-
minal tergite 8, dorsal; e: metatibia; f: mesofemur and mesotrochanter; g: abdominal sternite 8.
Scale bar = 0.2 mm.
AFROTROPICAL MEGARTHRUS 747
a-e,g
Fic. 57
Megarthrus simienensis; a, b: female, genital segment, tergites (a) and sternites (b); c: antenna; d:
female, abdominal sternite 8; e, g: female, apex of abdominal tergite 8, lateral and dorsal; f: pro-
notum. Scale bars = 0.2 mm.
748 GIULIO CUCCODORO & IVAN LOBL
Fic. 58
Megarthrus spinosus; a, c: female, genital segment, sternites (a) and tergites (c); b, f: female,
apex of abdominal tergite 8, dorsal and lateral; d: antenna; e: female, abdominal sternite 8; g: pro-
notum. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 749
h
|
ti
Fic. 59
Megarthrus stylifer, male; a, d: aedeagus, ventral and lateral; b: mesotibia; c: abdominal segments
7-10, lateral; e: metatibia; f, g: abdominal tergite 8, lateral and ventral. Scale bars = 0.2 mm.
750 GIULIO CUCCODORO & IVAN LOBL
Fic. 60
Megarthrus stylifer; a, h: abdominal sternite 8, male (a) and female (h); b: male, mesofemur and
mesotrochanter; c; male, metafemur and metatrochanter; d, antenna; e, f, female, genital segment,
tergites (e) and sternites (f); g, k: female, apex of abdominal tergite 8, dorsal and lateral; 1: pro-
notum. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 751
Fic. 61
Megarthrus twa, male; a, b: aedeagus, ventral and lateral; c, h: apex of abdominal tergite 8, dorsal
and lateral; d: mesotibia; e: metatibia; f: mesofemur and mesotrochanter; g: abdominal sternite 8.
Scale bars = 0.2 mm.
752 GIULIO CUCCODORO & IVAN LOBL
FIG. 62
Megarthrus twa; a: female, abdominal sternite 8; b: antenna; c, d: female, genital segment, ter-
gites (c) and sternites (d); e, g: female, apex of abdominal tergite 8, lateral and dorsal; f: pro-
notum. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 753
ON
NAS
SAQ
Fic. 63
Megarthrus vanschuytbroecki, male; a: metafemur and metatrochanter; b, f: apex of abdominal
tergite 8, dorsal and lateral; c, d: aedeagus, ventral and lateral; e: mesofemur and mesotrochanter;
g: mesotibia; h: metatibia; 1: abdominal sternite 8. Scale bar = 0.2 mm.
754 GIULIO CUCCODORO & IVAN LÔBL
a-f
Fic. 64
Megarthrus vanschuytbroecki; a: female, abdominal sternite 8; b, e: female, apex of abdominal
tergite 8, lateral and dorsal; c: antenna; d, f: female, genital segment, sternites (d) and tergites (f);
g: pronotum. Scale bars = 0.2 mm.
155
AFROTROPICAL MEGARTHRUS
(e)
x
sii eet
> /
IN, SK
==
SS >
Fic. 65
Megarthrus watutsi, male; a: abdominal sternite 8; b, c: aedeagus, ventral and lateral; d: meta-
tibia; e: mesotibia; f: apex of abdominal tergite 8, dorsal; g: mesofemur and mesotrochanter; h:
metafemur and metatrochanter. Scale bar = 0.2 mm.
756 GIULIO CUCCODORO & IVAN LOBL
FIG. 66
Megarthrus watutsi; a, c: female, genital segment, sternites (a) and tergites (c); b, g: female, apex
of abdominal tergite 8, dorsal and lateral; d: antenna; e: pronotum; f: female, abdominal sternite
8. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 157
Ya
7
Fic. 67
Megarthrus wittei, male; a: abdominal sternite 8; b, e: aedeagus, ventral and lateral; c: mesotibia;
d: metatibia; f: mesofemur and mesotrochanter; g: apex of abdominal tergite 8, dorsal; h:
metafemur and metatrochanter. Scale bars = 0.2 mm.
758 GIULIO CUCCODORO & IVAN LÖBL
Fic. 68
Megarthrus wittei; a, b: female, genital segment, tergites (a) and sternites (b); c: antenna; d; f:
female, abdominal sternite 8; e: pronotum; f: female, apex of abdominal tergite 8, dorsal. Scale
bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 759
> 22 > —- >
Ar \ EN
yale AR À IR
ise vA eZ, Wi A ff ; | \
‘ Aya "Al \ | N \
on qe AA as =/ I 3
K | ta eae EN 1, /
LR S
SIR >
SSS SS SN
fy SÌ
Fic. 69
Megarthrus zulu, male; a, b: aedeagus, ventral and lateral; c: mesotibia; d: metafemur and
metatrochanter; e: metatibia; f: mesofemur and mesotrochanter; g: abdominal sternite 8; h: apex
of abdominal tergite 8, dorsal. Scale bar = 0.2 mm.
760 GIULIO CUCCODORO & IVAN LOBL
Fic. 70
Megarthrus zulu; a: antenna; b, c: female, genital segment, sternites (b) and tergites (c); d:
female, abdominal sternite 8; e, g: female, apex of abdominal tergite 8, dorsal and lateral; f: pro-
notum. Scale bars = 0.2 mm.
AFROTROPICAL MEGARTHRUS 761
REFERENCES
BERNHAUER, M. 1931. Entomological Expedition to Abyssinia, 1926-7: Coleoptera, Staphy-
linidae. J. Linn. Soc., Zool., 37: 559-605.
BERNHAUER, M. & R. PAULIAN. 1942. Coléoptères Staphylinides du Cameroun (Mission P.
Lepesme, R. Paulian et A. Villier, 1939). Revue Zool. Bot. afr., 35: 344-375.
CAMERON, M. 1942. New Species of Staphylinidae (Col.) collected by the Coryndon Museum
Expedition to the Chyulu Hills. Ann. Mag. nat. Hist., 53: 321-332.
CAMERON, M. 1950. Staphylinidae (Coleoptera Polyphaga). Explor. Park nat. Albert, Miss. de
Witte (1933-1935), Brussels, fasc. 59: 85pp.
COoIFFAIT, H. 1982. Contribution à la connaissance des Staphylinides de I’ Himalaya (Népal,
Ladakh, Cachemire) (Insecta: Coleoptera: Staphylinidae). Senckenbergiana biol. 62: 21-
179.
Cuccoporo, G. 1995. Two new species of Megarthrus (Coleoptera, Staphylinidae, Proteininae)
and a note on "water loading" behaviour. J. Zool., 236: 253-264.
EICHELBAUM, F. 1913. Verzeichnis der von mir in den Jahren 1903 und 1904 in Deutsch- und
Britisch-Ostafrika eingesammelten Staphylinidae. Arch. Naturgesch., Abt. A (3): 114-
168.
FAGEL, G. 1957. Contribution à l’étude de la faune entomologique du Ruanda-Urundi (Mission P.
Basilewsky 1953). CXIX.Coleoptera Staphylinidae: Piestinae, Omaliinae, Proteininae,
Osorinae, Oxytelinae. Annis Mus. r. Congo belge, Ser. 8°, Zool., 58: 26-72.
HAYASHI, Y. 1988. Studies on Staphylinidae from Japan. 2. Entomol. Rev. Japan, 45: 17-23.
NEWTON, A. F. Jr. 1985. South temperate Staphylinoidea (Coleoptera): their potential for biogeo-
grafic analysis of austral disjunctions. In: G. E. Ball (ed.), Taxonomy, Phylogeny and
Zoogeography of Beetles and Ants, pp. 179-220. Dordrecht.
NEWTON, A. F. Jr. & M. K. THAYER. 1992. Current Classification and Family-Group Names in
Staphyliniformia (Coleoptera). Fieldiana: Zool., 67: 1-92.
STEEL, W. O. 1966. A revision of the Staphylinid subfamily Proteininae (Coleoptera) I. Trans. R.
ent. Soc. Lond., 118: 285-311.
THAYER, M. K. 1987. Biology and phylogenetic relationships of Neophonus bruchi, an ano-
malous south Andean staphylinid (Coleoptera). Syst. Ent., 12: 389-404.
aren) eran
canali
AE
MORN
REVUE SUISSE DE ZOOLOGIE, 102 (3) : 763-768; septembre 1995
Eine neue Haroldius-Art aus Borneo (Coleoptera, Scarabaeidae)
Joachim SCHEUERN
Westerwaldstr. 18, D-53489 Westum, BRD.
A new Haroldius species from Borneo (Coleoptera, Scarabaeidae).-
Haroldius pauliani n. sp. is described and illustrated from Sabah. Some
ecological aspects of Haroldius are briefly discussed, differences to similar
species (H. fleutiauxii Paulian and A. sumatranus Paulian & Scheuern) are
listed. A new key of the known species of the genus is provided.
Key-words: Coleoptera - Scarabaeidae - Haroldius - Taxonomy - Ecology
- Borneo.
EINLEITUNG
BALTHASAR (1963) beschreibt die Haroldius-Arten als echte Myrmecophile
mit unvollständigem Trutztyp, schlieBt auf Synoekie und führt H. heimi (Wasm.) (in
Nestern von Pheidole latinoda) und H. philippinensis Per. (bei Diacamma vagans-
Gruppe) an. H. perroti Paulian wurde zusammen mit der Histeride Bacanius Er. in
faulendem Laub gefunden (PAULIAN, 1939). Bei Ameisen und Termiten hat JANSSENS
(1949, 1953) afrotropische Arten der ehemaligen Gattung Afroharoldius Jans. (syn. zu
Haroldius Bouc., PAULIAN, 1985) nachgewiesen.
Von Scarabaeidae werden stets nur einzelne Exemplare der Gattung Harol-
dius, dagegen häufiger Panelus-Arten aus Gesieben von faulenden Blättern, Asten,
manchmal von faulenden Früchten an Waldrändern oder in Wäldern gelesen (Löbl, in
litt.). Zwar sind in diesen Habitaten auch Ameisen anzutreffen, doch ist eine generelle
Myrmecophilie der Gattung Haroldius Bouc. nicht ableitbar.
Die neue Art Haroldius pauliani n. sp. wurde wie H. sumatranus Paulian &
Scheuern und H. thailandensis Paulian & Scheuern (1994) aus Laubgesieben gelesen.
Der Bau der Mundwerkzeuge läßt auf eine saprophage, mycetophage oder
myrmecophage Ernährung, die Fundumstände auf phyllicole und detriticole Lebens-
weise von Haroldius Bouc. schließen.
PAULIAN (1993) beschreibt Haroldius discoidalis und H. borneensis aus
Borneo. Aus derselben Region "Sabah, Poring Hot Springs" liegt nun eine dritte, für
die asiatische Region die achtzehnte Art dieser Gattung vor.
Manuskript angenommen am 12.01.1995.
764
JOACHIM SCHEUERN
BESTIMMUNGSTABELLE DER ASIATISCHEN HAROLDIUS-ARTEN
1
2
Elytrenbasis deutlich zu einer Quergrube vertieft, Scheibe erhoht.......... 2
Elytren gleichmäßig gewölbt, Basis nicht grubig vertieft................. 5
Pronotumbasis scharf gekantet, ohne Kerben, Mitte in ein breites,
längsgekieltes Dreieck ausgezogen. Scheibe sehr fein, sehr spärlich
punktertsBanse23.0:mm.Borneo. 2, 2.2. EEE EEE pauliani n. sp.
Pronotumbasis lateral mit je 6 oder 7 kurzen, breiten, punktförmigen
Kerben, Mitte einfach gekantet mit einigen längeren, zarten, nach vorn
konvergierenden Streiten. Pronotumscheibe unpunktiert. 1,7-1,9 mm.
Sumatran: NOI ORE TATE LIMO AR, sumatranus R. Paul. & Scheuern
Pronotumbasis mit zahlreichen langen, schmalen Streifen, die fast die
fein und spärlich punktierte Scheibenmitte erreichen. 2,0 mm. Thailand,
NIET AR Re ee a et ata RE Sh fleutiauxii R. Paul.
Pronotumbasis mit querer, unscharf begrenzter Impression, ohne Ker-
ben oder Streifen. Pronotumscheibe unpunktiert, Seiten stark winkelig
eRWelterte2 Smmalhnalland "ter 07 thailandensis R. Paul. & Scheuern
Pronotumbasis mit einer queren, vorn deutlich begrenzten Impression,
danebenieventuell mit kerbenden Bunkten. 2 ee 4
Probotumbasis mit einer queren, unscharf begrenzten Impression und
sehr langen, sehr dicht stehenden, die Scheibenmitte erreichenden
Streifen2:5-3:0imm* Pakistan, Indien, Sri Lanka 2 72 2 ws oe 6
Pronotumbasis ohne Impression, mit mehr oder weniger langen Streifen
oder die Basis kerbenden Punkten. Ist die Basismitte schwach vertieft,
dann’stehen-laterallkerbende Punkte. "re LI NE 7
Pronotumbasis einfach gerundet, ohne Auszeichnungen. Clypeus neben
der Ausrandune micht/eeschweilts. RG 11
Probotumbasis neben der Impression mit einer dichten Reihe kerbender
Punkte. Vorderwinkel fein und spärlich, Scheibe groß, dicht, schwach
oval, dazwischen sehr fein punktiert. Punktdurchmesser größer als die
Zwischenräume, schwarz, kahl. 3,3-3,6 mm. Nepal, Darjeeling,
Assan Ihalland:, Eonkin.. ccs. cena ee yo Se eee stevensi Arr.
Kante der Pronotumbasis ohne kerbende Punkte. Mindestens der Vor-
derkorper gelbrot, Elytren fein behaart. CE zen u 5)
Körper oval, Pronotum mit verstreuten, ziemlich groben Punkten, Ely-
tren lang und fein behaart. 1,5 mm. Singapore. ........... fairmairei Bouc.
Körper gerundet, Pronotum sehr fein, spärlich punktiert, Elytren sehr
kurz und sehr spärlich behaart. 2,0-2,4 mm. Borneo. . . ... discoidalis R. Paul.
Außenkante der Hinterschienen einfach konvex gebogen, äußere Apikal-
spitze kurz und wenig scharf. Oberseite, besonders der Vorderkörper,
schwach, aber sichtbar chagriniert. 2,5 mm. Srı Lanka..... herrenorum R. Paul.
Außenkante der Hinterschienen konkav gegen den Apex gebogen, äußere
Apikalspitze scharf und leicht ausgezogen. Vorderkörper nicht, Elytren
erloschen chagriniert. 2,5-3,0 mm. Nordost-Indien, Pakistan. . . . heimi (Wasm.)
10
11
HAROLDIUS AUS BORNEO 765
Innenkante der Hinterschienen knieförmig gebogen. Nepal... . /assallei Camb.
Innenkante’derklinterschienenfeintachikonkay are Nr 8
Pronotumbasis mit langen, dicht stehenden Streifen, die lateral die
Scheibenmitte oder in Vorderwinkelnähe fast den Vorderrand
erteichen#25 2:85mm2Sin2aporem ar ee rugatulus Bouc.
Pronotumbasis mit kurzen, kräftigen Streifen. Arten aus Borneo oder
AEREI PPS en Eu ae. SOME IAA ere 9
Die gekerbte Pronotumbasis mit groben Punkten. 2,5-3,0 mm. Arten
ausi@entral-Indient.t. we trat a. I RR CIR. 10
2,0 mm, Oberseite chagriniert, Elytren lang behaart, Streifen ausge-
[ESC BONE SEE EU rk GI SEP SRLS, MEI ee borneensis R. Paul.
2,0 mm, Oberseite glatt, kahl, Elytrenstreifen ausgelöscht. Philippinen.
eroe deri Dito E gia: de ed globosus Bouc.
3,5-4 mm, Oberseite glatt, Elytren mikroskopisch fein behaart, Streifen
an der Basis deutlich, am Apex verschwindend. Philippinen. . philippinensis Per.
Scheitel deutlich chagriniert, dicht und doppelt punktiert, Punktdurch-
messer größer als die Zwischenräume. Punktierung des Pronotums vorn
und in der Basismitte dicht und kraftig, lateral bedeutend feiner und
spärlicher. Basiskante lateral mäßig fein, bis zum Niveau des 1.
Elytrenzwischenraumes grober crenuliert. Elytren sehr kurz, spärlich,
kaumssichtbaribeborstetzrr wre annandalai (Silv.)
Scheitel glänzend, erloschen chagriniert, mit wenigen feinen und sehr
feinen Punkten. Punktierung des Pronotums doppelt, fein und spärlich.
Basismitte scharf gekantet, bis zum Niveau des 2. Elytrenstreifens
schwach dreieckig eingedrückt, von den Hinterwinkeln bis zur Mitte des
5. Elytrenzwischenraumes fein crenuliert. Haare der Elytren spärlich,
deutlich, ihre Länge beträgt ca. ein Viertel einer Zwischenraumbreite.
ORI AL cardoni Bouc.
Pronotum fein, relativ dicht punktiert, wie die Elytren kaum bemerkbar
chagimer 1-9 mm lonkane anwaney el cis ae ee perroti R. Paul.
Pronotum sehr fein, mäßig dicht punktiert, quermaschig, Elytren längs-
maschig, deutlich chagriniert. 2,0 mm. Thailand............. loebli R. Paul.
Haroldius pauliani n. sp. (Abb.: 1-8)
Holotypus d : Borneo: Sabah Mt. Kinabalu National Park, Poring Hot Springs, 510 m,
30.VIII.1988, A. Smetana leg. (B162); Muséum d'histoire naturelle, Génève.
Diagnose: Kenntlich durch die Genitalarmaturen des d, den starken Apikal-
und sehr schwachen Basalzahn am Außenrand der Vorderschienen und die ventral
fehlende subapikale Querleiste, den neben den Zähnen ausgeschweiften Clypeusrand
und die breite, querovale, tiefe Depression von Elytren- und Pronotumbasis, die in der
Mitte zu einem breiten, spitzen Dreieck ausgezogen ist. Mäßig gewölbt, Elytren stark
gerundet, mäßig glänzend, Oberseite deutlich fein netzmaschig, Pronotumdepression
und Pygidium erloschen chagriniert, sehr fein, sehr spärlich, einfach punktiert. Beine,
766 JOACHIM SCHEUERN
Kopf, Pronotumvorderrand und -seiten und die Apikalrundung der Elytren rötlich-
braun; Pronotumdepression und Elytren schwarzbraun; Nahtzwischenraum leicht
aufgehellt; Pydigium, Sternite, Mundorgane und Fiihler gelbbraun. Kahl, nur Elytren-
seiten und -absturz mit winzigen, weißgelben Borsten. Augen schmal.
d: Kopf 1,5 mal breiter als lang, Clypeus tief, parabelförmig ausgerandet,
neben den beiden kräftigen, außen gerundeten Zähnen deutlich ausgeschweift, bis zu
den stumpfen Wangenecken gerundet, nur an der fein eingeritzten Naht kurz aus-
gerandet und schwach gewinkelt.
Pronotum 2,3 mal breiter als in der Mitte lang, Pronotumvorderrand in der
Mitte schwach nach vorn gebogen, lateral in Höhe der Augen und Wangenecken
jeweils schwach winkelig in die fast geraden, zum Elytrenseitenrand schwach diver-
gierenden Seiten übergehend. Hinter den deutlichen Hinterwinkeln sehr schwach
konkav. Basis ab dem 6. bis zum 3. Elytrenstreifen konvex gerundet, dann stark
konkav und in der Mitte in ein breites, spitz zulaufendes, hochgewölbtes, längsge-
kieltes, glänzendes Dreieck ausgezogen. Eine glänzende, querovale, tiefe Depression
wird kurz hinter dem Pronotumvorderrand und auf Höhe des 6. Elytrenzwischen-
raumes durch die wulstartige Scheibe begrenzt. Seiten zu den Vorderwinkeln dach-
artig abfallend. Nur die Basis ungerandet, scharf gekantet, vor dem 5. Elytrenstreifen
mit Randungsrudiment, ohne Längslinien oder Punkte.
Elytrenstreifen deutlich, seicht, mäßig breit, unpunktiert, an der Basis kaum
vertieft, am Apex erlöschend.
Zwischenräume flach; Naht bis zur Mitte leicht gekielt, vor der Spitze mit sehr
kurzem Streifen und kleinem Grübchen. 2. Zwischenraum an der ungerandeten Basis
so breit wie der 3., Basis bis zum 5. vertieft, an der Naht dreieckig ausgeschnitten,
kurz aufgebogen, scharf gekantet. Apex breit ausgerandet, Elytren nicht gemeinsam
verrundet.
Pygidium doppelt so breit wie hoch, Scheibe kaum gewölbt, glänzend, rundum
fein, Basis winkelig gerandet.
Vorderwinkel des Prosternum mit tiefer Grube, diese durch einen Querkiel und
-wulst von einer Furche zum Einlegen der Vorderschenkel getrennt. Vorderhüften
vorn scharf gekantet, dort mit einer Reihe kurzer, apikal mit einer Gruppe langer
Borsten. Episternum hinten spärlich, am Seiten- und Hinterrand dicht und kurz, davor
dicht und lang behaart. Scheibe des Metasternum im vorderen Drittel in ganzer Breite
tief ausgehöhlt. Vorn doppelt und scharf gekantet, die Hinterkante bedeutend höher
gelegen, in der Mitte kurz unterbrochen und fast die Vorderkante erreichend. Beide
Kanten enden dicht bei den Mittelhüften. Scheibe sehr fein, kaum sichtbar punktiert,
vorn glatt, nach hinten und zu den Seiten deutlich längs chagriniert. Zwischen den
Mittel- und Hinterhüften deutlich, mäßig dicht, längs punktiert. Episterna und
Epipleuren stark längs chagriniert, mit einigen Punkten. Epipleuren in Höhe der
Hinterhüften mit runder Grube, nach hinten mit einigen flachen, kleineren Depres-
sionen. Sternite quer chagriniert, an den Seiten deutlicher punktiert, nur das letzte in
der Mitte nicht verengt und am Hinterrand mäßig dicht, deutlich punktiert und kurz
behaart.
HAROLDIUS AUS BORNEO 767
ABB. 1-8
Haroldius pauliani n. sp.: 1: Kopf und Pronotum dorsal, 2: Vorderschiene dorsal, 3: dto.
ventral, 4: Mittelschiene dorsal, 5: Hinterschiene, dorsal, 6: Aedoeagus lateral; 7: Parameren
dorsal, 8: Kopulationsspange. Maßstriche: 0,5 mm; kurzer Maßstrich: Abb. 1.
Vorderschienen zum Apex gleichmäßig verbreitert, dorsal mit scharfer, zum
Außenrand parallel laufenden Kante; daneben kräftig, einreihig, in der Mitte feiner,
am Apex unregelmäßig punktiert. Außen mit einem sehr kräftigen, spitzen Apikal-
zahn, zur Basis gerundet in ein sehr winziges, stumpfes Zähnchen auslaufend. Vor-
derrand ventral mit einem deutlichen, vorn behorsteten Querwulst, eine subapikale
Querleiste fehlt. Vorderschenkel vorn mit einer Längsdepression zum Einlegen der
Schiene, dorsal innen mit einem dichten Haarfeld.
768 JOACHIM SCHEUERN
Mittel- und Hinterschienen apikal nach außen stark gerundet erweitert, innen
gleichmäßig, schwach gebogen. Mittel- und Hinterschenkel schmal, lang, die
Epipleuren überragend, ohne Auszeichnungen.
Genitalarmaturen: Abb. 6, 7, 8.
Holotypus d : Länge; 3,0 mm, Elytrenbreite: 2,2 mm.
Derivatio nominis: Benannt zu Ehren des ausgezeichneten Scarabaeidae-
Spezialisten Prof. Dr. R. Paulian, Bordeaux.
Beziehungen: Haroldius pauliani n. sp. bildet mit H. fleutiauxii R. Paul. und
H. sumatranus R. Paul. & Scheuern eine gut definierte Artengruppe: Symmetrisch
gebaute Parameren mit stumpfwinkelig abgeknickten Spitzen, Clypeus neben der
Ausrandung ausgeschweift, Vorderschienen ventral ohne subapikale Querleiste. Die
neue Art unterscheidet sich von den beiden genannten durch die Form der
Kopulationsspangen, die querovale Depression der Pronotumbasis mit dem scharf
ausgezogenen Dreieck, die unbeborstete Elytrenscheibe, den vorn nicht eingeengten
2. Elytrenzwischenraum, die Form der Schienen und die stattliche Größe. H. suma-
tranus R. Paul. & Scheuern weist am 4. Elytrenzwischenraum eine Basalrandung auf.
DANK
Herrn Prof. Dr. R. Paulian, Bordeaux, danke ich sehr fiir wertvolle Hinweise,
Herrn Dr. I. Lôbl, Genève, und Herrn Dr. Y. Cambefort, Paris, zudem für die
Ausleihe von Typen.
LITERATUR
BALTHASAR, V. (1963). Monographie der Scarabaeidae und Aphodiidae der palaearktischen und
orientalischen Region, Band 1. Verlag der Tschechoslowakischen Akademie der Wis-
senschaften, Prag.
JANSSENS, A. (1949). Un Scarabaeinae termitophile nouveau du Congo Belge. Revue Zool. Bot.
afr., 42(2): 183-184.
JANSSENS, A. (1953). Contribution à l'étude des Coléoptères Lamellicornes d'Afrique. II.
Espèces nouvelles du genre Afroharoldius A. Janssens. Revue Zool. Bot. Afr., 47(34):
344-347.
PAULIAN, R. (1939). Quelques nouvelles espèces des Coléoptères Lamellicornes coprophages.
Bull. Soc. Ent. Fr., 44: 68-74.
PAULIAN, R. (1945). Coléoptères Scarabaeides de l'Indochine. Faune de l'Empire Frangais, 3,
Paris.
PAULIAN, R. (1985). Notes sur les Coléoptères Scarabaeidae du Muséum de Genève II. Revue
suisse Zool., 92(1): 189-203.
PAULIAN, R. (1993). Deux nouveaux Haroldius Boucomont de Bornéo (Coléoptères Scara-
baeidae). Revue suisse Zool., 100(1): 169-173.
PAULIAN, R. & SCHEUERN, J. (1994). Haroldius Boucomont nouveaux ou peu connus de la
région Orientale (Coléoptères Scarabaeidae). Revue suisse Zool., 101(2): 435-440.
REVUE SUISSE DE ZOOLOGIE, 102 (3) : 769-778; septembre 1995
Dichelyne moraveci n. sp., parasite de Pseudoplatystoma fasciatum
et notes sur les Cucullanidae du Paraguay
Annie J. PETTER
Laboratoire de Biologie Parasitaire, Protistologie, Helminthologie,
associé au CNRS, Muséum National d'Histoire Naturelle, 61, rue Buffon,
F-75231 PARIS Cedex 05, France.
Dichelyne moraveci sp. n. parasite of Pseudoplatystoma fasciatum, with
remarks on Cucullanids from Paraguay. - The following Cucullanids are
recorded from freshwater fishes of Paraguay: Dichelyne (D.) moraveci sp.
n. in Pseudoplatystoma fasciatum, Luciopimelodus pati and Megalonema
platanum; Cucullanus pinnai in Pseudoplatystoma coruscans and Megalo-
nema platanum; females of Cucullanus sp. in Pimelodus maculatus. Diche-
lyne (D.) moraveci differs from D. (D.) leporini, the other South American
congener, by its very long spicules. A key to the 10 South American
Cucullanus species is given.
Key-words: Nematoda - Cucullanidae - New species - Freshwater Fishes -
Paraguay - South America.
INTRODUCTION
La famille des Cucullanidae est fondamentalement parasite de Poissons, avec
quelques espèces chez les Chéloniens. Elle est actuellement représentée chez les
Poissons d'eau douce sud-américains par 3 genres: 1 genre endémique, Neocucullanus
avec 2 espèces, et 2 genres cosmopolites: Cucullanus avec 10 espèces et Dichelyne
avec | espèce [voir PETTER (1989) et MORAVEC & al. (1993)]. Nous décrivons ici une
nouvelle espèce du genre Dichelyne récoltée au Paraguay par l'Expédition Zoologique
du Museum d'Histoire Naturelle de Genève en 1986; nous signalons également la
présence au Paraguay d'une espèce du genre Cucullanus, C. pinnai, déja connue au
Brésil et en Argentine.
MATERIEL ET MÉTHODES
Les viscères des Poissons sont fixés au formol à 4% sur le terrain; la recherche
des parasites est effectuée au Laboratoire à Genève; les Nématodes recueillis sont
conservés dans l'alcool à 70°C et éclaircis au lactophénol pour l'étude. Le matériel est
Manuscrit accepté le 16.12.1994.
770 ANNIE J. PETTER
déposé au Laboratoire de Biologie Parasitaire du Muséum National d'Histoire
Naturelle de Paris (MNHN) et au Muséum d'histoire naturelle de Genève (MHNG).
Toutes les mensurations sont en um.
Cucullanus pinnai Travassos, Artigas & Pereira, 1928 (fig.1)
Matériel étudié: 12 d et 8 2 juvéniles n° MNHN 403 BC; hôte: Pseudoplatystoma
coruscans (Agassiz) (Pimelodidae, Siluriformes) n° de terrain Py 4971. 6 8,5 2 juvéniles et
une région antérieure n° MHNG 18166 INVE: hôte: Megalonema platanum (Günther)
(Pimelodidae) n° de terrain Py 4972 (co-parasites de Dichelyne (D.) moraveci). Rio Parana, en
face de Candelaria, prov. Itapua, 10 et 12.12.1986.
Dimensions ( Extrêmes, moyenne entre parenthèses).
Males [dans l'ordre: parasites de Pseudoplatystoma coruscans (11 spécimens),
parasites de Megalonema platanum (5 spécimens)].
Long. 3000-4000 (3490); 2450-5600 (3470). Oesophage 390-600 (509); 350-
725 (474). Extrémité antérieure - anneau nerveux 180-250 (211); 140-250 (188). Extré-
mité antérieure - deirides 350-450 (389); 320-575 (425). Spicules 250-370 (325); 115-
470 (307).
Femelles [dans l'ordre: parasites de P. coruscans (2 spécimens), parasites de
M. platanum (5 spécimens)].
Long. 3700/4100 (3900); 2400-3300 (2700). Oesophage 520/600 (560); 360-
500 (400). Extrémité antérieure - anneau nerveux 220/250 (235); 130-210 (165).
Extrémité antérieure - deirides 380/450 (415); 300-400 (339). Queue 120/340 (230);
120-150 (125).
Les spécimens correspondent par l'ensemble de leurs caractères et de leurs
dimensions a l'espèce Cucullanus pinnai décrite par TRAVASSOS & al. (1928) chez
Pimelodus clarias et Pseudoplatystoma sp.; en particulier la disposition des papilles
ad-cloacales, très constante chez tous nos spécimens, et la forme de l'extrémité distale
des spicules sont tout-à-fait semblables à celles figurées par ces auteurs. Signalons
que les spécimens juvéniles parasites de Pseudoplatystoma coruscans sont dépourvus
de ventouse pré-cloacale.
L'espèce a été signalée ultérieurement par différents auteurs chez plusieurs
espèces de Pimelodidae, Characidae et Anostomidae et redécrite par MORAVEC & al.
(1993) chez Pimelodus ornatus et Ageneiosus valenciennesi. Ces auteurs décrivent
chez Pseudoplatystoma coruscans une autre espèce, Cucullanus pseudoplatystomae et
suggèrent que les spécimens trouvés précédemment chez des espèces du genre
Pseudoplatystoma et attribués à Cucullanus pinnai pourraient appartenir en réalité à
cette dernière espèce. C. pseudoplatystomae diffère de C. pinnai par la position des
phasmides, par la longueur plus grande du corps, de l'oesophage et de la queue de la
femelle, et par une position différente de l'anneau nerveux et des deirides par rapport a
l'extrémité de l'oesophage. Si l'on excepte la position des phasmides qui sont situées
antérieurement aux deux dernières paires de papilles postcloacales comme chez C.
pseudoplatystomae, les dimensions de nos spécimens correspondent a C. pinnai
(tableau I), le genre Pseudoplatystoma peut donc héberger les deux espèces.
DICHELYNE MORAVECI WIA
Fic. 1
Cucullanus pinnai Travassos, Artigas & Pereira, 1928. A a E: spécimens parasites de Pseu-
doplatystoma coruscans. A, femelle, région antérieure, vue médiane. B, femelle, queue, vue
latérale. C, male, région postérieure, vue ventrale. D, male, extrémité postérieure, vue ventrale.
E, male, extrémité postérieure, vue latérale. F a H: spécimens parasites de Megalonema plata-
num. F, male, région antérieure, vue médiane. G, H, males, régions postérieures, vues latérales.
Echelles: A, B, J, K, L, N: 100 um; C, D, G, HI : 200 um; E, 30 pm; F, 50 um; O, 20 um.
772 ANNIE J. PETTER
TABLEAU I
Dimensions comparées de Cucullanus pseudoplatystomae et C. pinnai suivant MORAVEC & al.
(1993); comparaison avec le présent matériel.
Espèce C. pseudoplatystomae C. pinnai C. pinnai
Auteurs Moravec & al. présent matériel
= P. ornatus
Hötes Ps. coruscans A i . Ps. coruscans — M. platanum
. valenciennesi
long. totale 8570-15160 3700-9070 3000-4100 2400-5600
long. oesoph. 1090-1470 626-898 390-600 350-725
long. queue 9 299-394 150-258 120-340 120-150
extr. ant. - an. nerveux
/long. oesoph. 26-31% 35-40% 33-46% 34-46%
extr. ant. - deirides
/long. oesoph. 41-65% 74-85% 60-96% 75-100%
Cucullanus sp. (fig.2)
Matériel étudié: 2 2 juvéniles n° MNHN 300 BC, une larve n° MNHN 311 BC, une
femelle juvénile n° MNHN 395 BC. Hote: Pimelodus maculatus Lacépède (Pimelodidae) n° de
terrain Py 4964, Rio Parana, en face de Ouro Verde, prov. Alto Parana, 30.11.1986; Py 4767,
San Lorenzo, prov. Central, 13.7.1986; Py 4980, Rio Parana en face de Candelaria, prov.
Itapua, 15.12.1986.
Dimensions (dans l'ordre: 2 femelles 300 BC, une femelle 395 BC)
Long. 5300/4900/3850. Oesophage 750/775/700. Extrémité antérieure -
anneau nerveux 300/280/280. - deirides 600/650/400. - vulve 3200/2900/2350. Queue
200/220/200.
La morphologie et les dimensions correspondent a celles de C. pinnai; en
l'absence de males, nous préférons cependant ne pas faire d'identification spécifique.
Dix espèces parasites de Poissons d'eau douce sud-américains sont à présent
connues dans le genre Cucullanus. La plupart sont parasites de Siluriformes, en parti-
culier de Pimelodidae, et trois espèces ont été trouvées chez des Characiformes;
l'espèce C. pinnai a été signalée à la fois chez des Siluriformes et des Characiformes,
la spécificité parasitaire n'est donc pas très stricte. D'autre part, une méme espèce hòte
peut être parasitée par deux espèces différentes (ainsi, l'espèce Zungaro zungaro est
parasitée a la fois par Cucullanus oswaldocruzi et par C. zungaro).
Plusieurs caractères ont été utilisés pour différencier les espèces:
1. La longueur du corps et celle des spicules; ce caractère doit être utilisé avec
prudence car ces dimensions présentent de grandes variations dans une méme
espèce: ainsi, chez Cucullanus pinnai, la longueur varie de 2,4 mm a 8,5 mm
et celle des spicules de 2,8 à 14% de la longueur du corps.
2. La présence d'ailes caudales: ce caractère signalé chez certaines espèces
nécessiterait d'être vérifié car une légère saillie du corps de chaque côté du
cloaque peut étre prise pour des ailes.
DICHELYNE MORAVECI 773
FIG. 2
Cucullanus sp. A, région antérieure, vue médiane. B, queue, vue latérale.
3. La présence ou l'absence d'une ventouse pré-cloacale; ce caractère est constant
dans une espèce mais n'est pas applicable aux spécimens juvéniles (nous avons
vu plus haut que chez C. pinnai, les males juvéniles sont dépourvus de ven-
touse).
4. La forme de l'extrémité distale des spicules: chez certaines espèces, l'extrémité
distale des spicules est coudée; ce caractère est un bon caractère spécifique car
il paraît constant dans une espèce (nous l'avons observé chez tous les males de
C. pinnai examinés); cependant, il peut étre difficile 4 observer chez les spéci-
mens où les spicules ne saillent pas hors du cloaque, et a pu étre omis par
certains auteurs.
5. La disposition des papilles cloacales; ce caractère paraît également très cons-
tant dans une espèce d'après nos observations sur C. pinnai; cette disposition
est cependant mal décrite chez certaines espèces.
Nous donnons une clé des espèces, en nous efforçant pour les raisons exposées
ci-dessus d'indiquer dans la mesure du possible plusieurs caractères pour définir
chaque espèce.
Nous avons utilisé pour désigner les papilles la terminologie de CHABAUD &
PETTER (1961) et PETTER (1974) que nous rappelons brièvement: il existe générale-
ment dans le genre Cucullanus 11 paires de papilles: 8 paires subventrales numérotées
1, 2, 3, 5, 6, 7, 9, 10, 2 paires latérales numérotées 4 et 8 (fig. 3) et une paire de
774 ANNIE J. PETTER
500
FIG. 3
Cucullanus colossomi Diaz-Ungria, 1968. Male holotype, extrémité postérieure, vue latérale.
Les phasmides n'ont pas été vues.
phasmides latérales; les papilles 4, 5, 6, 7 et quelquefois 9 sont situées à proximité du
cloaque [papilles ad-cloacales de Petter (1974)].
Chez les espèces sud-américaines C. pseudoplatystomae et C. pimelodellae,
Moravec & al. (1993) signalent chez certains spécimens une paire pré-cloacale
supplémentaire entre la 3ème et la Sème paire.
CLE DES ESPECES SUD-AMERICAINES DU GENRE Cucullanus
1 Ventouserabsente. : um... Maike Ser ET I 2
Ventouse présente: ML Le in O TIE, 5
2 Spicules mesurant moins de 10% de la longueur du corps
Se 0 ARDITI ICI DIRI LU C. pimelodellae Moravec & al., 1993
= Spicules mesurant plus de 15% de la longueur du corps
SE AA Be OE FIALE ae Pe Eat al eek C. grandistomis (Ferraz & Thatcher,1988)
DICHELYNE MORAVECI 775
3) Ailes latérales présentes; spicules mesurant moins de 2% de la longueur
UICOLDS ere DONS. ae Ar C. brevispiculus Moravec & al.,1993
- Ailes latérales absentes; spicules mesurant plus de 2,5 % de la longueur
AULEOTBSK FE foro re a Ele. dere BOs 2 4
~ Spiculesicoudessaskextnemite distale teo RER EE EP PTE 5
Spicules simplement incurvés ventralement à l'extrémité distale. .......... 7
5 Longueur des mâles supérieure à 12 mm; ailes caudales présentes;
papilles n° 7 beaucoup moins grosses que les papilles n° 6 et n° 9
a a a I We C. pauliceae Vaz & Pereira, 1934
(= C. schubarti Travassos, 1948)
_ Longueur des mâles inférieure à 12 mm; ailes caudales absentes;
papilles n° 7 à peu près de même taille que les papilles n° 6etn° 9........ 6
6 Anneau nerveux légèrement antérieur au milieu de l'oesophage (situé
entre 33 et 46 % de sa longueur); deirides légèrement antérieures ou au
niveau de l'extrémité de l'oesophage (situées entre 60 et 100 % de sa
ln) SERRE ere C. pinnai Travassos et al., 1928
= Anneau nerveux nettement antérieur au milieu de l'oesophage (situé
entre 26 et 31 % de sa longueur); deirides nettement antérieures a
l'extrémité de l'oesophage (situées entre 41 et 65 % de sa longueur)
Re e C. pseudoplatystomae Moravec & al., 1993
7 Longueur des mâles supérieure a 12 mm; ailes caudales absentes.......... 8
- Longueur des mâles inférieure à 12 mm; ailes caudales présentes.......... 9
8 Papilles n° 9 deux fois plus proches des papilles n° 7 que des papilles
n° A0 sy Gi ce Se ne C. colossomi Diaz-Ungria, 1968 !
— Distance entre les papilles n°7 et n° 9 a peu près égale a la distance
entrelles;papillesn2 Yetnal 0.22.02... C. oswaldocruzi Santos & al., 1979
9 Papilles n° 5 situées latéralement par rapport aux papilles n°6; papilles
n°5 et 6 d'une part, 4, 7 et 9 d'autre part formant 2 groupes nettement
séparés; papilles n° 7 nettement post-cloacales. . . . . C. mogi Travassos, 1947
— Papilles n° 5, 6, 7 et 9 alignées en 2 rangées subventrales à peu pres
régulièrement espacées; papilles n° 7 en position ad-cloacale.
I RE) C. zungaro Vaz & Pereira, 1934
Dichelyne (Dichelyne) moraveci n. sp. (fig.4)
Matériel-type: 4 holotype, © allotype, une région postérieure 4 et deux régions anté-
rieures paratypes, n°MNHN 408 BC; un d paratype n° MHNG 18167 INVE. Hôte: Pseudo-
platystoma fasciatum (L.) (Pimelodidae, Siluriformes), n° de terrain Py 4958. Rio Parana, en
face de Puerto El Dorado, prov. Itapua, 23.11.1986.
Autre matériel: - un d juvénile et une vingtaine de larves n° MHNG 18168 INVE. Hote:
Luciopimelodus pati (Val.) (Pimelodidae) n° de terrain Py 4770. Méme localité, 25.9.1986. - un
d et une région antérieure 9 n° MHNG 18169 INVE. Hote: Megalonema platanum (Günther),
n° de terrain Py 4972. Méme localité,12.12.1986. (Co-parasites de Cucullanus pinnai).
! Nous avons examiné le d holotype de C. colossomi déposé au MNHN et pu ainsi pré-
ciser la disposition des papilles cloacales de cette espèce (fig. 3).
116 ANNIE J. PETTER
200
Fic. 4
Dichelyne (D) moraveci n. sp. A, vue apicale. B, coupe optique au niveau des piéces scléro-
tisées attachées au cadre péribuccal. C, femelle, région antérieure, vue latérale. D, mâle, région
antérieure, vue médiane. E, deiride. F, male, post-deiride. G, femelle, extrémité postérieure, vue
latérale. H, vulve et ovéjecteur. I, male, région postérieure, vue latérale (a: post-deiride gauche).
J, male juvénile, région postérieure, vue ventrale. K, male, extrémité postérieure, vue ventrale.
L, mâle, extrémité postérieure, vue latérale. M, extrémité distale du spicule. N, larve, région
antérieure, vue latérale. O, larve, extrémité antérieure, vue médiane. Echelles: A, B, G, F: 200
um; C, D, H, E: 100 pm.
DICHELYNE MORAVECI 777
Cucullanidae de petite taille, à corps robuste. Cuticule épaisse (15 à 20 um).
Collerette membraneuse denticulée entourant l'ouverture buccale comprenant une
cinquantaine de dents de chaque côté. Pièces sclérotisées attachées au cadre péri-
buccal bien visibles. Oesophage avec renflement antérieur plus marqué que le ren-
flement postérieur. Caecum intestinal dorsal présent, mesurant environ la moitié de la
longueur de l'oesophage chez les spécimens parasites de Pseudoplatystoma fasciatum,
plus court chez les spécimens parasites de Luciopimelodus pati et Megalonema
platanum. Deirides terminées par une pointe fine, situées en avant de l'extrémité de
l'oesophage. Post-deirides bien développées, arrondies a l'extrémité. Pore excréteur
situé légèrement en arriére des deirides. Queue conique, terminée par une pointe fine
dans les deux sexes.
Male: ventouse absente. 10 paires de papilles caudales: 3 paires pré-cloacales;
5 paires ad-cloacales (selon la terminologie de Petter, 1974) comprenant 3 grosses
paires subventrales (2 antérieures au cloaque et une postérieure), une paire latérale
située entre les 2 paires subventrales antérieures, et une paire de petites papilles ven-
trales pédonculées encadrant la lèvre supérieure du cloaque; 2 paires post-cloacales
situées au voisinage de l'extrémité postérieure; phasmides situées antérieurement aux
2 paires post-cloacales. Lèvre supérieure du cloaque bilobée, surmontée par une
protubérance papilliforme, mais dépourvue de terminaison nerveuse. Gubernaculum
peu sclérotisé. Spicules très longs (de 40 a 53% de la longueur du corps), ailés,
terminés en pointe fine.
Femelle: vulve à lèvres saillantes, légèrement post-équatoriale. Ovéjecteur
impair comprenant un vagin à paroi musculaire épaisse de 350 um et une trompe à
paroi mince de 400 um. Uterus opposés. Oeufs à coque mince, non embryonnés dans
l'uterus.
Larves: les larves, vraisemblablement des 4èmes stades, sont dépourvues de
collerette membraneuse denticulée; l'oesophagostome est muni de 2 dents subven-
trales proéminentes; le caecum intestinal est absent.
Dimensions: Males (dans l'ordre: holotype/paratypes)
Long. 4650/4750/1600/1750. Larg. maximale 400/350/100/350.
Oesophage 900/850/430/720.Caecum intestinal 560/500/150/260. Extrémité
antérieure - anneau nerveux 250/250/170/240. - deirides 550/600/230/605. - pore
excréteur 670/625/260/ non vu. Extrémité postérieure - post-deiride gauche
900/940/420/900. - post-deiride droite 2200/1770/ non vue . Queue 130/120/95/110.
Spicules 2050/2500/700/1930.
Femelle: Long. 5400. Larg. maximale 400. Oesophage 960. Caecum intestinal
450. Extrémité antérieure - anneau nerveux 280. - deirides 530. - pore excréteur 600. -
vulve 3100. Extrémité postérieure - post-deiride gauche 1300. Post-deiride droite non
vue. Queue 140. Oeufs 60/50.
Discussion. Par la présence d'un caecum intestinal et l'absence de ventouse
pré-cloacale, l'espèce se place dans le genre Dichelyne (sous-genre Dichelyne) suivant
la classification de PETTER (1974) et de CHABAUD (1978). Une seule espéce appar-
tenant au genre Dichelyne a été décrite chez les Poissons d'eau douce sud-américains:
778 ANNIE J. PETTER
D.(D.) leporini décrite par PETTER (1989) chez Leporinus friderici au Paraguay, et
redécrite par MORAVEC & al. (1993) chez Schizodon fasciatus au Brésil. La nouvelle
espèce s'en différencie par des spicules beaucoup plus longs (44 à 53% de la longueur
du corps contre 7 a 11% chez D. leporini, une cuticule beaucoup plus épaisse (15 a 20
um contre 4 à 5) et une disposition différente des papilles cloacales (les 2 paires
situées immédiatement en avant du cloaque sont accolées ou sur un méme niveau
transversal chez D. (D.) leporini). La nouvelle espèce se différencie de toutes les
autres espèces connues dans le sous-genre Dichelyne soit par des spicules plus longs
par rapport a la longueur du corps, soit par une pointe caudale dépourvue d'épines,
soit par une disposition différente des papilles cloacales.
BIBLIOGRAPHIE
CHABAUD, A.G. 1978. CIH keys to the nematode parasites of Vertebrates. 6. Keys to genera of
the superfamilies Cosmocercoidea, Seuratoidea, Heterakoidea and Subuluroidea.
Commonwealth Agricultural Bureaux, Farnham Royal, Bucks, England, 71 pp.
CHABAUD, A.G. & A.J. PETTER. 1961. Remarques sur l'évolution des papilles cloacales chez les
Nématodes Phasmidiens parasites de Vertébrés. Parassitologia 3: 51-70.
DIAZ-UNGRIA, C. 1968. Helmintos de Peces de Venezuela, con descripcion de un genero y tres
especies nuevas. Boln Soc. venez. Cienc. nat., 27: 537-549.
FERRAZ, E. & V.E. THATCHER. 1988. Bacudacnitis grandistomis gen. et sp. nov. (Nematoda:
Cucullanidae) parasita intestinal de um bagre, Pseudodoras niger (Valenciennes) da
Amazonia brasileira. Amazoniana 10: 249-253.
MORAVEC, F., A. KoHN & B.M.M. FERNANDES. 1993. Nematode parasites of fishes of the
Parana River, Brazil. Part 2. Seuratoidea, Ascaridoidea, Habronematoidea and Acua-
rioidea. Folia parasit. 40: 115-134.
PETTER, A.J. 1974. Essai de classification de la famille des Cucullanidae. Bull. Mus. natn. Hist.
nat., 3e sér., 255, Zool. 177: 1469-1490.
PETTER, A.J. 1989. Nématodes de Poissons du Paraguay. V. Cucullanidae. Description de deux
espèces nouvelles et redéfinition du genre Neocucullanus Travassos et al. Revue suisse
Zool. 96: 591-603.
SANTOS E., J.J. VICENTE & C.R. JARDIM. 1979. Helmintos de peixes de Rios Amazonicos da
coleçao helmintologica do Instituto Oswaldo Cruz. II. Nematoda. Atas Soc. Biol. Rio de
J. 20: 11-19.
TRAVASSOS, L. 1948. Contribuigao ao conhecimento dos helmintos dos peixes d'agua doce do
Brasil. III. Duas novas espécies do genero Cucullanus Müller, 1877. Mems Inst.
Oswaldo Cruz 45: 551-554.
TRAVASSOS, L., P. ARTIGAS & C. PEREIRA. 1928. Fauna helminthologica dos peixes de agua
doce do Brasil. Archos Inst. Biol. S. Paulo 1: 5-68.
VAZ, Z. & C. PEREIRA. 1934. Contribuiçao ao conhecimento dos nematoides de peixes fluviais
do Brasil. Archos Inst. Biol. S. Paulo 5: 89-103.
REVUE SUISSE DE ZOOLOGIE, 102 (3) : 779-846; septembre 1995
Aleocharinae della Sottoregione Africana Orientale al
Museo di Ginevra (Coleoptera, Staphylinidae) Parte II *
Roberto PACE
Via Vittorio Veneto, 13. 37032 Monteforte d'Alpone (Verona), Italia.
Aleocharinae from the Eastern African Subregion in the Geneva
Museum (Coleoptera, Staphylinidae). Part II. — Based on the study of
80 species of the tribes Tachyusini and Athetini, 54 species are described
as new. A key to the species of the genus Alomacrotona Pace is given.
Illustrations for fifteen holotypes or lectotypes of species described by
Fauvel, Bernhauer, Eichelbaum and Cameron are given. One species,
Atheta mombassana Bernhauer, 1934, is newly synonymised with Atheta
paludosa Bernhauer, 1931. Three species are transferred to other genera.
The new genus Charicera is described and illustrated, based on a new
species. It is related to Pelioptera Kraatz. The new genus Askeptoxenia,
erected for Atheta acuticollis Fauvel (= A. fauveli Fenyes) is illustrated.
Key-words: Coleoptera - Staphylinidae - Aleocharinae - Taxonomy - East
Africa.
INTRODUZIONE
La prima parte dello studio sulle Aleocharinae dell'Africa orientale raccolte nel
corso di missioni di ricerca, promosse dal Museo di Storia Naturale di Ginevra, è stata
da me pubblicata sulla presente "Revue suisse de Zoologie" nel 1994. Il presente lavoro
è la sua prosecuzione e riguarda due tribù: Tachyusini e Athetini.
La seconda di queste tribù comprende qui un maggior numero di specie, dato
che le Aleocharinae della tribù Athetini prediligono il clima temperato e il clima
temperato freddo e le ricerche del Museo di Ginevra si sono svolte prevalentemente in
zone d'alta quota dei maggiori massicci dell'Africa orientale.
Anche nella presente Parte II, come nella Parte I, pubblico e illustro i tipi delle
specie di Aleocharinae relative alle due tribù trattate e designo il lectotypus, quando
necessario. Mi sono limitato a pubblicare solo quindici tipi: la pubblicazione di tutti i
tipi di Atheta dell'Africa da me esaminati, avrebbe comportato un notevole aumento di
pagine e avrebbe un po' fuorviato il fine principale del presente lavoro, quello cioè di
* 126° Contributo alla conoscenza delle Aleocharinae.
Manoscritto accettato il 23.12.1994.
780 ROBERTO PACE
rendere noto e mettere in ordine sistematico quanto scoperto nel corso delle ricerche
promosse dal Museo di Ginevra.
Al materiale affidatomi in studio dal Museo di Ginevra ho aggiunto alcune
specie datemi in esame dal Museo di Storia Naturale di Genova, dal D.E.I. di
Eberswalde e dal Museo di Erfurt.
Gli holotypi delle nuove specie si conservano nel Museo di Storia Naturale di
Ginevra (MG), nel Museo di Storia Naturale di Genova (MGE) e nel D.E.I. di Ebers-
walde. Paratypi sono custoditi nel Museo di Ginevra, in mia collezione e nel Museo di
Erfurt.
ELENCO SISTEMATICO DEI GENERI E DELLE SPECIE E DESCRIZIONI
TACHUSINI
Amanota externa (Fauvel, 1907) (Figg. 1-2)
Tachyusa externa FAUVEL, 1907: 59.
Amanota externa, PACE, 1986: 95, 109.
Materiale esaminato: Lectotypus 2, Nakuro (Rift-Valley), externa FVL, Tachyusa,
presente designazione; 3 paralectotypi, stessa provenienza (Mus. Roy. Bruxelles).
La specie è qui illustrata per la prima volta.
Amanota interna sp. n. (Figg. 3-4)
Holotypus 9, Kenya, Embu, pr. Ishiara, 900 m, 14.X.1977 (Mahnert & Perret leg., MG).
Descrizione. Lungh. 2,1 mm. Corpo lucido e bruno-rossiccio con uriti liberi 4° e
5° bruni; antenne bruno-rossicce con antennomero basale rossiccio e 11° giallo; zampe
bruno-rossicce con base dei femori e tarsi giallo-rossicci. La punteggiatura del capo è
quasi indistinta, su un fondo non reticolato. Tubercoletti fini e ben salienti sono presenti
solo sulla metà posteriore del pronoto. La punteggiatura e la reticolazione delle elitre
sono distinte. Gli uroterghi sono privi di reticolazione e hanno una punteggiatura
superficiale. Spermateca fig. 4.
Comparazioni. E' da escludere che la nuova specie sia identica ad A. eastopi
Tottenham, 1957 del Kenya o ad A. albicornis Tottenham, 1957 dell'Urundi, perché
queste specie presentano l'11° antennomero e i due uriti liberi basali colorati di giallo
pallido e le elitre sono più lunghe del pronoto, e non un po' più corte come quelle della
nuova specie. Per il colore delle antenne e dell'addome, come per la forma della
spermateca, la nuova specie sembra sistematicamente vicina ad A. externa (Fauvel,
1907) (Figg. 1-2) della "Rift-Valley", ma la spermateca nettamente più corta, le elitre
più lunghe del pronoto e il pronoto più largo osservabili in externa, sono caratteri
sufficienti a distinguere externa da interna sp. n.
Etimologia. Il nome della specie è dato in contrapposizione alla specie A.
externa (Fauvel).
ATHETINI
Amischa kaszabi Pace, 1986
Amischa kaszabi PACE, 1986: 96.
781
ALEOCHARINAE DELL’ AFRICA ORIENTALE
Imm
ine)
01 mm
01 mm
FIGG. 1-4
Habitus e spermateca. 1-2: Amanota externa (Fauvel), lectotypus; 3-4: Amanota interna sp. n.
782 ROBERTO PACE
1 ©, Kenya, Narok près de Narok, 200 m, 3.X1.1977 (Mahnert & Perret leg.); 2 89,
Kenya, Lac Nakuru, Pare Nat., 6.XI.1974 (Mahnert & Perret leg.); 35 es., Kenya, Nakuru, Lac
Naivasha près de Mundui Estate, 1950 m, 8.X11.1977 (Mahnert & Perret leg.); 1 9, Kenya,
Nakuru, Lac Elmenteita, 1800 m, 7.XI.1977 (Mahnert & Perret leg.); 1 es., Kenya, L. Baringo
env. Loruk, 1100 m (Mahnert & Perret leg.).
Specie già nota della Tanzania e di Kibosho.
Aloconota asymmetrica sp. n. (Figg. 5-6)
Holotypus 9, Rhodésie, Umtali, 11.1969 (R. Mussard leg., MG).
Descrizione. Lungh. 2,5 mm. Corpo lucido e bruno; margine posteriore degli
uroterghi rossiccio; antenne brune con antennomero basale giallo-bruno; zampe gialle.
La punteggiatura del capo e del pronoto è fine e distinta. Le elitre e l'addome sono
coperti di tubercoletti distinti. Spermateca Fig. 6.
Comparazioni. Per avere le elitre lunghe quanto il pronoto, la nuova specie è
chiaramente distinta da A. praticola (Fauvel, 1907) (Figg. 17-19) del Kilimangiaro, che
presenta elitre più lunghe del pronoto. Inoltre il 5° antennomero della nuova specie è
nettamente trasverso, mentre in praticola è lungo quanto largo; il pronoto di praticola è
nettamente reticolato, mentre quello della nuova specie è privo di reticolazione. La
nuova specie è pure distinta da A. africana (Bernhauer, 1934) (Figg. 7-10) per la forma
della spermateca, per le elitre più corte e per gli antennomeri 5° e 6° nettamente
trasversi.
Etimologia. La nuova specie prende nome da un carattere della spermateca:
l'asimmetria del bulbo distale.
Aloconota africana (Bernhauer, 1934), comb. n. (Figg. 7-10)
Pelioptera africana Bernhauer, 1934: 214.
Materiale esaminato: Lectotypus 4, W Riwenzori, Kalonge, myrmecophil avec Dorylus,
8.V111.1932, Burgeon, africana BRNH, Typus, Pelioptera (Mus Chicago).
1d e 2 9 9, Kenya, Mt. Aberdares, Parc National, 2300 m, 25.X1.1974 (Mahnert &
Perret leg.); 1 6, Kenya, Embu, Irangi for., 12.X.1977 (Mahnert & Perret leg.); 2 d d, Kenya,
Tambach, E Eldoret, 2000 m, 17.X1.1974 (Mahnert & Perret leg.); 3 es., Kenya, Nyandarua, 10
km SE Njabini, 2550 m, 9.X1.1977 (Mahnert & Perret leg.); 4 es., Kenya, Nyandarua, South
Kingango, 2550 m, 9.XI.1977 (Mahnert & Perret leg.); 2 es., Kenya, Nakuru, Mau Esc. près
Erangiperi, 2700 m, 6.X.1977 (Mahnert & Perret leg.); 1 es., Kenya, Nakuru, Elmenteita, 1800
m, 7.XI.1977 (Mahnert & Perret leg.); 3 es., Kenya, Embu, Irangi For. 2000 m, 11.X.1977
(Mahnert & Perret leg.); 3 es., Rwanda, Kayove, 2100 m, 29.1V.1973 (Werner leg.); 24 es.,
Rwanda, Rangiro, 1800 m, 1.1.1976, 26.11.1976, 14.IX.1976 (Werner leg.); 3 es., Rhodésie,
Umtali, 11.1969 (R. Mussard leg.).
Discussione. Il transferimento di questa specie dal genere Pelioptera Kraatz,
1857 al genere Aloconota Thomson, 1858, si è reso necessario allorché ho rinvenuto
due femmine tra il materiale del Museo di Ginevra. La spermateca non ha la forma
consueta al genere Pelioptera, cioè bulbo distale molto sviluppato e parte prossimale
della spermateca stessa fortemente ricurva al lato sinistro e molto assottigliata, cioè al
contrario di quanto è osservabile sulla spermateca di africana (fig.10). Effettivamente
le mesocoxe in africana sono un po' scostate tra loro, carattere questo riconducibile a
Pelioptera e che ha probabimente indotto Bernhauer all'attribuzione a Pelioptera.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 783
FIGG. 5-10
Habitus, spermateca ed edeago in visione laterale e ventrale. 5-6: Aloconota asymmetrica sp. n.;
7-10: Aloconota africana (Bernhauer), lectotypus d.
784 ROBERTO PACE
Tuttavia non sempre questo carattere è costante nell'ambito di un genere, come ho
constatato anche in altri generi di Athetini.
Al genere Aloconota appartengono specie per lo più ripicole. La raccolta di A.
africana con formiche è da ritenersi occasionale. Infatti al Museo di Ginevra sono
conservati numerosi esemplari di questa specie raccolti al di fuori dell'ambiente mirme-
cofilo.
Specie finora nota solo del Ruwenzori occidentale e ritenuta mirmecofila perché
raccolta con Dorylus sp.
Aloconota mussardi sp. n. (Figg. 11-16)
Holotypus d, Rhodésie, Umtali, II.1969, Melsetter, 1700 m (R. Mussard Leg., MG).
Paratypi: 2 9 9, stessa provenienza (MG, CPA).
Descrizione. Lungh. 2,6 mm. Corpo lucido e bruno; margine posteriore degli
uroterghi rossiccio; antenne brune; zampe giallo-rossicce. La reticolazione del capo è
molto svanita, quella del pronoto è assente e quella delle elitre e dell'addome è distinta.
La punteggiatura del capo è svanita, quella del pronoto fine. Le elitre sono coperte di
tubercoletti distinti, l'addome di tubercoletti superficiali. Edeago figg. 12-13, sperma-
teca figg. 14-15 e 6° urotergo libero del maschio Fig. 16.
Comparizioni. Specie simile ad A. africana (Bernhauer, 1934) (Figg. 7-10) del
Kenya e del Ruwenzori, a motivo delle elitre più lunghe del pronoto e della forma
dell'edeago. Tuttavia la nuova specie ha antenna più corte, edeago più ricurvo al lato
ventrale e pezzi copulatori del sacco interno dell'edeago, nettamente più robusti.
Aloconota praticola (Fauvel, 1907), comb. n. (Figg. 17-19)
Atheta praticola FAUVEL, 1907: 56.
Atheta (Microdota) praticola, Bernhauer & Scheerpeltz, 1926: 630.
Pelioptera praticola; Pace, 1986: 109.
Materiale esaminato: Holotypus d, Afrique or. all., Kilimanjaro, zone des prairies, 3200
altit., Ch. Alluaud, 1904, praticola FVL, type (Mus. Bruxelles).
Discussione. Il rinvenimento nel materiale del Museo di Ginevra di una specie
assai affine a praticola (A. mauensis sp. n.), che presenta spermateca tipica del genere
Aloconota Thomson, 1858 e non spermateca simile alle specie del genere Pelioptera
Kraatz, 1857, induce alla nuova attribuzione generica.
Aloconata mauensis sp. n. (Figg. 20-23)
Holotypus dé, Kenya, Nakuru Mau Escarpment, S East Mau, 2850 m, 6.X1.1977
(Mahnert & Perret leg., MG).
Paratypus, 1 9, stessa provenienza.
Descrizione. Lungh. 2,9 mm. Corpo lucido e bruno con elitre e addome rossicci;
antenne brune con antennomero basale bruno-rossiccio; rampe giallo-rossicce. La
punteggiatura del capo è distinta, su un fondo non reticolato. Sul pronoto sono presenti
tubercoletti salienti, sulle elitre tubercoletti molto salienti e sull'addome tubercoletti
distinti. La reticolazione del pronoto è distinta. Edeago figg. 21-22, spermateca Fig. 23.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 785
15
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Figc. 11-19
Habitus, edeago in visione laterale e ventrale, spermateca e 6° urotergo libero del maschio.
11-16: Aloconota mussardi sp. n.; 17-19: Aloconota praticola (Fauvel), Holotypus ¢.
756 ROBERTO PACE
Comparazioni. Specie affine ad A. praticola (Fauvel, 1907) (Figg. 17-19), del
Kilimangiaro, a motivo della forma dell'edeago. Tuttavia la nuova specie presenta elitre
più corte, pronoto debolmente reticolato (nettamente reticolato in praticola); tuber-
coletti delle elitre del maschio molto salienti presso la sutura (poco salienti e fini in
praticola) e 5° urotergo libero del maschio con una carena mediana (senza carena in
praticola). L'edeago delle due specie presenta il seguente carattere differenziale più
evidente: la piastra basale interna del sacco interno dell'edeago della nuova specie ha
uno stiletto corto, mentre quello corrispondente di praticola è lunghissimo.
Charicera gen. n. (Figg. 24-29)
Typus generis. Charicera ruandensis sp. n.
Diagnosi. Il nuovo genere si colloca tra i generi Aloconota Thomson, 1858 e
Pelioptera Kraatz, 1857, a motivo della presenza di una ligula divisa in due lembi a
base larga, ma questi lembi sono nettamente più stretti e molto distanziati tra loro (tra
loro contigui in Aloconota e Pelioptera). Inoltre le antenne sono cortissime (general-
mente lunghe in Aloconota) e la spermateca è di tipo diverso.
Descrizione. Habitus di Atheta Thomson, 1858, con antenne molto corte. Capo
più stretto del pronoto, con tempie marginate; palpi mascellari (Figg. 28-29) di 4
articoli; lobo interno delle "maxillae" con lunghe e fitte spine al margine apicale
interno; lobo esterno pubescente all'estremità; palpi labiali di 3 articoli (fig.26); ligula
divisa in due lembi tra loro largamente distanziati; paraglosse nulle; mento (fig.27)
trapezoidale con margine anteriore arcuato verso l'interno; processo mesosternale acuto,
insinuato fino a metà delle mesocoxe che tra loro sono contigue; formula tarsale 4-5-5;
primo tarsomero posteriore corto. Spermateca fig. 25.
Etimologia. Il nome del nuovo genere significa "Antenne graziose".
Charicera ruandensis sp. n. (Figg. 24-29)
Holotypus ©, Rwanda, Kayove, 2100 m, 29.1V.1973 (Werner leg., MG).
Descrizione. Lungh. 2,6 mm. Corpo lucido e giallo-rossiccio sporco; antenne
bruno-rossicce con i due antennomeri basali giallo-rossicci; zampe giallo-rossicce. La
reticolazione del capo e del pronoto è svanita, quella delle elitre assente e quella
dell'addome estremamente svanita. Il capo presenta punteggiatura fine e netta, il resto
del corpo è coperto di tubercoletti fini e distinti. Spermateca fig. 25.
Parapodoxya jeanneliana (Bernhauer, 1944)
Oxypoda (Podoxya) jeanneliana BERNHAUER, 1944: 20.
Oxypoda (Parapodoxya) jeanneliana, JEANNEL & PAULIAN, 1945: 96.
Parapodoxya jeanneliana, PACE, 1986: 109.
23 es., Kenya, Mt. Elgon, env. 2650 m, 14.X1.1974 (Mahnert & Perret leg.).
Specie gia nota del Mt. Elgon (localita tipica) (JEANNEL & PAULIAN, 1944) e del
Mt. Meru (PACE, 1986). Specie infeudata nei nidi di Tachyorictes splendens.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 787
û
01 mm
Fiss. 20-29
Habitus, edeago in visione laterale e ventrale, spermateca, labio con palpo labiale, maxilla, palpo
mascellare e mento. 20-23: Aloconota mauensis sp. n.; 24-29: Charicera ruandensis gen. n., sp. n.
788 ROBERTO PACE
Pelioptera aberdarensis sp. n. (Figg. 30-32)
Holotypus d, Kenya, Mt. Aberdares, Parc National, 2300 m, 25.X1.1974 (Mahnert &
Perret leg., MG).
Descrizione. Lungh. 2,6 mm. Corpo lucido e bruno, con uriti liberi 1°, 2° e 6°
giallo-rossicci; antenne brune con 1 due antennomeri basali giallo-bruni; zampe gialle.
La reticolazione del capo e del pronoto è netta, quella delle elitre quasi vigorosa e
quella dell'addome assente. L'avancorpo è coperto di tubercoletti poco distinti o
superficiali, l'addome presenta invece tubercoletti distinti. Edeago figg. 31-32.
Comparazioni. Le specie del genere Pelioptera Kraatz, 1857, della sottoregione
africana orientale sono: P. polita (Eichelbaum, 1913), P. abyssinorum Pace, 1986, P.
quaesita Pace, 1986 e P. consors Pace, 1986. E' da escludere l'affinità della nuova
specie con polita, poiché questa specie ha pubescenza dell'avancorpo molto rada. La
restanti specie hanno pubescenza fitta come della nuova specie, ma esse presentano
edeago molto largo nella regione subapicale, in visione ventrale, carattere, questo,
sufficiente a distinguere la nuova specie dalle altre dell'Africa orientale.
Pelioptera ruandensis sp. n. (Figg. 33-35)
Holotypus d, Rwanda, Kayove, 2100 m, 12.VIII.1973 (Werner leg., MG).
Descrizione. Lungh. 2,7 mm. Corpo lucido e bruno; antenne brune con antenno-
mero basale e apice dell'11° giallo-bruni; zampe giallo-rossicce. Il capo presenta
un'ampia concavità discale, punteggiatura assai distinta e reticolazione evidente solo sul
disco: al di fuori di esso è assente. Sul pronoto è visibile un appiattimento mediano
posteriore diviso sulla linea mediana da una carena superficiale, punteggiatura distinta e
reticolazione assai svanita. Le elitre presentano tubercoletti distinti, reticolazione evi-
dente e una carena alla sutura, presso l'angolo posteriore interno. Edeago figg. 33-34.
Comparazioni. La presenza di una carena mediana sul 5° urotergo libero del
maschio e la forma dell'edeago, permettono di riconoscere l'affinità della nuova specie
con P. consors Pace, 1986, della Tanzania. Tuttavia la nuova specie ha la regione
presso la "crista apicalis" dell'adeago, molto saliente e le elitre del maschio hanno una
plica all'angolo posteriore interno (assente in consors, che ha invece tubercoletti molto
salienti sulla regione suturale delle elitre).
Pelioptera dimidiata sp. n. (Figg. 36-38)
Holotypus ©, Rwanda, Rangiro, 1800 m, 6.VIII.1973 (Werner leg., MG).
Descrizione. Lungh. 2,1 mm. Corpo lucido e giallo-rossiccio con elitre e
addome bruno-rossicci e uriti liberi 4° e 5° bruni; antenne bruno-rossicce con i due
antennomeri basali e l'11° giallo-rossicci; zampe gialle. Il capo e il pronoto sono lucidi,
cioè senza reticolazione e hanno punteggiatura finissima. Le elitre e l'addome pre-
sentano tubercoletti distinti e non sono reticolati. Edeago figg. 37-38.
Comparazioni. La forma dell'edeago della nuova specie appare un po' simile a
quella di P. abyssinorum Pace, 1986. Infatti entrambe le specie presentano una plica
presso la "crista apicalis" dell'edeago e l'apice dell'edeago termina a punta di lancia. La
più vistosa differenza tra le due specie consiste nella presenza di un flagello lungamente
ALEOCHARINAE DELL’ AFRICA ORIENTALE 789
Ficc. 30-35
Habitus ed edeago in visione laterale e ventrale. 30-32: Pelioptera aberdarensis sp. n.; 33-35:
Pelioptera ruandensis sp. n.
790 ROBERTO PACE
E
E
5
Fiss. 36-40
Habitus, edeago in visione laterale e ventrale e maxilla con palpo mascellare. 36-38: Pelioptera
dimidiata sp. n.; 39-40: Askeptoxenia acuticollis (Fauvel), gen. n., holotypus d.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 791
sporgente dall'orifizio apicale dell'edeago della nuova specie, mentre in abyssinorum
non è presente e l'edeago di abyssinorum è molto più sviluppato di quello della nuova
specie.
Askeptoxenia gen. n. (Figg. 39-44)
Typus generis: Askeptoxenia acuticollis (Fauvel, 1907).
Diagnosi. Per la forma della ligula, per le antenne allungate e per il primo
tarsomero posteriore lungo quanto i due seguenti tarsomeri considerati insieme, il
nuovo genere può essere collocato accanto al genere Aloconota Thomson, 1858. Tutta-
via il pronoto sinuato davanti agli angoli posteriori, la presenza di un solo solco
trasverso basale dell'addome (e non due o tre), la presenza di scultura squamiforme
sugli uroterghi basali e la profonda flessione al lato ventrale dell'edeago, che ha apice
diviso in due lembi, sono caratteri che distinguono il nuovo genere da Aloconota, come
da altri generi affini: Pelioptera Kraatz, 1857, Geostiba Thomson, 1859, ecc.
Descrizione. Capo più stretto del pronoto, con tempie robustamente marginate;
mandibole molto allungate, con dente interno alla base della terebra, assai poco
sporgente; terebra acuta all'apice; lobo interno delle "maxillae" (fig. 40) più corto e più
stretto dell'esterno, con una fila interna di robuste spine e di fini setole; lobo esterno con
estremità pubescente; palpi mascellari composti di 4 articoli: il terzo è stretto e molto
lungo; mento (fig. 44) trasverso, con margine anteriore quasi rettilineo; palpi labiali di 3
articoli (fig. 43): il primo articolo è assai lungo e il secondo è brevissimo; ligula a base
molto larga, divisa in due lembi che si restringono verso l'apice; paraglosse non
sporgenti in avanti; antenne di 11 articoli; con undicesimo antennomero lungo quanto 1
tre antennomeri precedenti compresi insieme, nel maschio.
Elitre molto più lunghe e più larghe del pronoto. Addome lievemente ristretto
all'indietro; 1 tre uroterghi liberi basali presentano microscultura squamiforme; il primo
urotergo libero è trasversalmente solcato alla base. Formula tarsale 4-5-5; tarsomero
basale posteriore lungo quanto i due tarsomeri contigui considerati insieme. Processo
mesosternale acuto: esso raggiunge la metà delle mesocoxe, che tra loro sono contigue.
Edeago figg. 41-42. Femmina sconosciuta.
Etimologia. Il nome del nuovo genere significa "Ospite non riconosciuto".
Infatti la specie acuticollis è ospite di Termiti e il nuovo genere non è stato riconosciuto
da Fauvel quando descrisse la specie, dato che non ha esaminato ligula ed edeago,
avendo disponible un solo esemplare che non ha osato dissezionare.
Askeptoxenia acuticollis (Fauvel, 1907), comb. n. (Figg. 39-44)
Atheta (Dimetrota) acuticollis FAUVEL, 1907: 52.
Atheta (Dimetrota) fauveli FENYES, 1920: 205 ("nomen novum" non necessario);
BERNHAUER & SCHEERPELTZ, 1926: 663.
Materiale esaminato: Holotypus d, Kilimanjaro, Kibonoto, 12, Termites, acuticollis
FVL, Atheta, (Mus. Bruxelles).
Geopora aequinoctialis (Fauvel, 1900) (Figg. 45-48)
Atheta aequinoctialis, Fauvel, 1900: 73.
Atheta (s. str.) aequinoctialis, BERNHAUFR & SCHEERPELTZ, 1926: 638.
792 ROBERTO PACE
Fic. 41-44
Edeago in visione laterale, ventrale (solo l'apice) e dorsale, labio con palpo labiale e mento.
41-44: Askeptoxenia acuticollis (Fauvel), gen. n., holotypus d.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 793
Geopora aequinoctialis, PACE, 1986: 109.
Materiale esaminato: Lectotypus 4, Congo, Kinchassa, 11, Atheta aequinoctialis FVL,
type, presente designazione (Mus. Bruxelles); paralectotypi: 1 d e 2 2 9, stessa provenienza
(Mus. Bruxelles).
La specie è qui illustrata per la prima volta.
Geopora umtaliensis sp. n. (Figg. 49-50)
Holotypus 9, Rhodésie, Umtali, II.1969 (R. Mussard leg., MG).
Descrizione. Lungh. 2,0 mm. Corpo lucido e giallo con 4° urite libero giallo-
rossiccio; antenne giallo-rossicce; zampe gialle. Il capo e il pronoto hanno punteggia-
tura confusa nell'evidente reticolazione. Le elitre presentano punteggiatura poco evi-
dente e reticolazione svanita. Reticolazione netta e tubercoletti fini coprono l'addome.
Spermateca fig. 50.
Comparazioni. Il corpo molto stretto della nuova specie e la forma inconsueta
della spermateca (con gibbosità preapicale), non permettono di avvicinare sistemati-
camente la nuova specie ad altre specie note.
Leptoporodota tambachensis sp. n. (Figg. 51-52)
Holotypus ©, Kenya, Tambach, E Eldoret, 2000 m, 17.X1.1974 (Mahnert & Perret leg.,
MG).
Descrizione. Lungh. 2,1 mm. Corpo lucido e giallo rossiccio con elitre e uriti
liberi 4° e base del 5° rossicci; antenne giallo-rossicce con i due antennomeri basali
gialli; zampe gialle. La reticolazione del capo e del pronoto è distinta, quella delle elitre
e dell'addome è netta. La punteggiatura del capo è estremanente svanita. Tubercoletti
molto svaniti coprono le elitre. Spermateca fig. 51.
Comparazioni. Per la forma della spermateca (a S rovesciata), la nuova specie
può essere avvicinata sistematicamente a L. kiboensis Pace, 1985, del Kilimangiaro, ma
questa specie non presenta una profonda introflessione apicale del bulbo distale della
spermateca, nè il bulbo prossimale dilatato come nella nuova specie. Esternamente la
nuova specie ha elitre più lunghe del pronoto e occhi poco ridotti, mentre in kiboensis le
elitre sono più corte del pronoto e gli occhi sono ridottissimi.
Atheta (Microdota) tapina sp. n. (Figg. 53-57)
Holotypus d, Kenya, Narok, Loita Hills, sous Morijo, 2050 m, 5.X1.1977 (Mahnert &
Perret leg., MG).
Paratypus: 1 d, stessa provenienza.
Descrizione. Lungh. 1,9 mm. Corpo lucido e bruno, comprese le antenne; zampe
gialle. La reticolazione del capo e del pronoto è distinta, quella delle elitre è netta e
quella dell'addome superficiale. Il capo presenta punteggiatura svanita. Il pronoto e le
elitre sono coperti di tubercoletti distinti. Edeago figg. 54-55, maxille e palpo
mascellare fig. 56, labio con palpo labiale fig. 57.
Comparazioni. L'edeago della nuova specie è simile a quello di A. eichelbaumi
Bernhauer, 1915, della Tanzania, per i suoi lati paralleli nella regione subapicale, in
visione ventrale, ma l'edeago della nuova specie è più sviluppato di quello di
794 ROBERTO PACE
Figc. 45-52
Habitus, edeago in visione laterale e ventrale e spermateca. 45-48: Geopora aequinoctialis
(Fauvel), lectotypus d e paralectotypus 9; 49-50: Geopora umtaliensis sp. n.; 51-52: Lepto-
porodota tambachensis sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 795
eichelbaumi e ha un distinto pezzo copulatore arcuato del sacco interno, assente in
eichelbaumi.
Atheta (Microdota) kisumuensis sp. n. (Figg. 58-62)
Holotypus d, Kenya, Miss. Kaimosi, NE Kisumu, 1650 m, 11.XI.1974 (Mahnert &
Perret leg., MG).
Paratypi: 5 es., stessa provenienza.
Descrizione. Lungh. 1,7 mm. Corpo debolmente lucido e nero, comprese le
antenne; zampe giallo-rossicce. Solo il capo e l'addome mostrano reticolazione estrema-
mente svanita: il resto del corpo è privo di reticolazione. I tubercoletti che coprono la
superficie del capo sono superficiali, quelli sul resto del corpo sono distinti. Edeago
figg. 59-60, spermateca fig. 61 e 6° urite libero del maschio fig. 62.
Comparazioni. Per la forma simile dell'habitus e della spermateca, la nuova
specie si colloca sistematicamente vicina ad A. kiboshoana Bernhauer, 1915, della
Tanzania (nota solo sull'holotypus femmina). Le due specie sono separate dai seguenti
più evidenti caratteri: introflessione apicale del bulbo distale della spermateca molto
profonda nella nuova specie e poco profonda in kiboshoana; pronoto meno trasverso
nella nuova specie e più trasverso in kiboshoana.
Atheta (Microdota) mussardi sp. n. (Figg. 63-66)
Holotypus d, Rhodésie, Umtali, 11.1969 (R. Mussard leg., MG).
Paratypi: 4 d d, stessa provenienza.
Descrizione. Lungh. 1,7 mm. Corpo lucido e bruno, comprese le antenne; zampe
giallo-rossicce. La reticolazione del capo e dell'addome è distinta, quella del pronoto e
delle elitre è svanita. Tutto il corpo è coperto di tubercoletti salienti. Edeago figg.
64-65, 6° urotergo libero del maschio fig. 66.
Comparazioni. La nuova specie è differente da A. eichelbaumi Bernhauer, 1915,
della Tanzania, per l'edeago più sviluppato e per i lati dell'edeago (in visione ventrale)
convergenti verso l'apice e non paralleli come in eichelbaumi.
Atheta (Microdota) acusifera sp. n. (Figg. 67-70)
Holotypus d, Rhodésie, Umtali, IT.1969 (R. Mussard leg., MG).
Descrizione. Lungh. 2,1 mm. Corpo lucido e bruno con uriti liberi 3°, 4° e 5°
nero-bruni; antenne brune; zampe giallo-rossicce. Il capo e il pronoto presentano pun-
teggiatura superficiale e reticolazione distinta. Il capo ha una larga concavità discale. La
reticolazione delle elitre è superficiale, quella degli uroterghi svanita e a maglie tras-
verse. Tubercoletti svaniti coprono le elitre. Edeago figg. 68-69, 6° urotergo libero del
maschio fig. 70.
Comparazioni. L'edeago della nuova specie presenta alcuni caratteri riscontrabili
nell'edeago di A. dilaticollis Pace, 1986, della Tanzania. Tuttavia l'appendice aghiforme
ventrale dell'edeago, rende la specie finora unica.
Etimologia. Il nome della nuova specie significa "Portatrice d'ago"; infatti
l'edeago porta un'appendice aghiforme ventrale.
796 ROBERTO PACE
Le | \
In
EN
ML
N
01 mm
FIGG. 53-62
Habitus, edeago in visione laterale e ventrale, maxilla con palpo mascellare, labio con palpo
labiale, spermateca e 6° urotergo libero del maschio. 53-57: Atheta (Microdota) tapina sp. n.;
58-62: Atheta (Microdota) kisumuensis sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 797
01 mm
0,1 mm
Figc. 63-70
Habitus, edeago in visione laterale e ventrale, 6° urotergo libero del maschio. 63-66: Atheta
(Microdota) mussardi sp. n.; 67-70: Atheta (Microdota) acusifera sp. n.
798 ROBERTO PACE
Atheta (Microdota) kipsigia sp. n. (Figg. 71-74)
Holotypus d, Kenya, Mt. Elgon, env. 2650 m, 24.X1.1974 (Mahnert & Perret leg., MG).
Paratypi: 5 es., stessa provenienza, ma 2700 m e 15.X1.1974.
Descrizione. Corpo lucido e bruno con elitre bruno-rossicce e addome nero;
antenne interamente brune; zampe giallo-brune. L'avancorpo è coperto di tubercoletti e
di reticolazione netti. L'addome presenta reticolazione molto svanita e tubercoletti
distinti. Edeago figg. 72-73, spermateca fig. 74.
Comparazioni. In base alla forma della spermateca, simile a quella di A.
kiboshoana Bernhauer, 1915, della Tanzania, la nuova specie è attribuita al sottogenere
Microdota Mulsant & Rey, 1873, nonostante la taglia del corpo maggiore e le antenne
allungate. L'introflessione apicale del bulbo distale della spermateca della nuova specie,
è profonda e triangolare e non breve e a semicerchio come in kiboshoana. Inoltre la
dimensione della spermateca della nuova specie è nettamente maggiore rispetto quella
di kiboshoana. Il 4° antennomero della nuova specie è più lungo che largo, mentre
quello corrispondente di kiboshoana è nettamente trasverso.
Etimologia. La nuova specie prende nome dalla tribù nilotica dei Kipsigi,
stanziata a SE del M. Elgon, località tipica.
Atheta (Traumoecia) reptabunda Pace, 1986
Atheta (Traumoecia) reptabunda PACE, 1986: 101.
12 es., Kenya, Mt. Aberdares, Parc National, 2300 m, 25.XI.1974 (Mahnert & Perret
leg.); 1 9, Kenya, Taita Hills, 1400 m, env. Wundanyi (Mahnert & Perret leg.).
Specie finora nota solo dell'Africa orientale.
Atheta (Traumoecia) irangicola sp. n. (Figg. 75-78)
Holotypus d, Kenya, Embu, [rangi forest St., 2100 m, 12.X.1977 (Mahnert & Perret leg.,
MG).
Paratypus: | 2, stessa provenienza.
Descrizione. Lungh. 2,0 mm. Corpo debolmente lucido e giallo-bruno con 4°
urite libero bruno; antenne brune con i due antennomeri basali giallo-rossicci; zampe
giallo-rossicce. La reticolazione del capo e delle elitre è netta, quella del pronoto è dis-
tinta. Tubercoletti nettamente salienti coprono l'avancorpo. Sull'addome i tubercoletti
sono svaniti, posti su un fondo a reticolazione distinta e a maglie appena trasverse.
Edeago figg. 76-77, sesto urotergo libero del maschio fig. 78. Spermateca simile a
quella di A. Kipsigia sp. n. (fig. 74), ma più breve, con una dilatazione presso il bulbo
prossimale.
Comparazioni. L'edeago della nuova specie è simile per grandezza e per aspetto
generale a quello di A. reptabunda Pace, 1986, della Tanzania, tuttavia nel sacco
interno dell'edeago della nuova specie è assente una larga spina ricurva, presente invece
in reptabunda e le "bretelle" dell'edeago sono strettissime nella nuova specie e
larghissime in reptabunda. Inoltre la nuova specie ha elitre nettamente più lunghe del
pronoto e occhi ben sviluppati, mentre in reptabunda le elitre sono lunghe quanto il
pronoto e gli occhi sono molto ridotti.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 799
Fico. 71-78
Habitus, edeago in visione laterale e ventrale, spermateca e 6° urotergo libero del maschio.
71-74: Atheta (Microdota) kipsigia sp. n.; 75-78: Atheta (Traumoecia) irangicola sp. n.
800 ROBERTO PACE
Atheta (Tropatheta) shimbaensis sp. n. (Figg. 79-82)
Holotypus d, Kenya, Shimba Hills, Madakara For., 30.XI.1974 (Mahnert & Perret leg.,
MG).
i Paratypi: 7 es., stessa provenienza.
Descrizione. Lungh. 2,3 mm. Corpo lucido e bruno; antenne bruno-rossicce con
i tre antennomeri basali giallo-rossicci; zampe rossicce. La reticolazione del capo e del
pronoto è vigorosa, quella delle elitre è svanita. La punteggiatura del capo è ombelicata
e netta, quella del pronoto è superficiale e quella delle elitre è altrettanto netta. Il
pronoto ha uno stretto e superficiale solco mediano. Edeago figg. 80-81, spermateca
fig. 82.
Comparazioni. L'appendice stiliforme ventrale dell'edeago della nuova specie è
carattere unico e ben distinto che, insieme ad altri numerosi caratteri differenziali, sia
dello stesso edeago, che della spermateca, permette una netta distinzione da A. gestroi
Bernhauer, 1915, dell'Eritrea.
Atheta (Tropatheta) savanicola sp. n. (Figg. 83-85)
Holotypus d, Usambara, coll. Kraatz (D.E.I.).
Descrizione. Lungh. 2,9 mm. Corpo lucido e bruno; antenne giallo-brune con i
quattro antennomeri basali giallo-rossicci; zampe giallo-rossicce con femori rossicci. La
reticolazione del capo e del pronoto è netta, quella delle elitre è distinta e quella
dell'addome è assente. La punteggiatura del capo è svanita, quella delle elitre è distinta.
Tubercoletti poco salienti coprono il pronoto. Edeago figg. 84-85.
Comparazioni. L'edeago della nuova specie, rispetto quello di A. gestroi
Bernhauer, 1915, dell'Eritrea, è nettamente meno sviluppato (1/3 di meno) e più
profondamente ricurvo al lato ventrale. E' assente, nel sacco interno dell'edeago della
nuova specie, un lungo flagello, presente invece nel sacco interno dell'edeago di
gestroi. I caratteri dell'esoscheletro sono invece simili.
Atheta (Tropatheta) loitensis sp. n. (Figg. 86-89)
Holotypus d, Kenya, Narok Morijo, Loita Hills, 2300 m, 4.X1.1977 (Mahnert & Perret
leg., MG).
Paratypi: 71 es., stessa provenienza, ma anche in data 5.X1.1977 e altitudine 2050-2200 m.
Descrizione. Lungh. 2,2 mm. Corpo lucido e bruno-rossiccio con pronoto giallo-
rossiccio; antenne bruno-rossicce con i due antennomeri basali giallo-rossicci; zampe
giallo-rossicce. La reticolazione dell'avancorpo è svanita, quella degli uroterghi è
estremamente svanita. La punteggiatura del capo i tubercoletti che coprono il pronoto e
le elitre sono superficiali. Edeago figg. 87-88, spermateca fig. 89.
Comparizioni. Le elitre lunghe quanto il pronoto e la forma dell'edeago e della
spermateca, sono caratteri sufficienti a distinguere la nuova specie da A. gestroi
Bernhauer, dell'Eritrea.
Atheta (Tropatheta) embuicola sp. n. (Figg. 90-91)
Holotypus 2, Kenya, Embu, Kirimiri Forest, O de Runyenje, 1550 m, 13.X.1977
(Mahnert & Perret leg., MG).
ALEOCHARINAE DELL’AFRICA ORIENTALE 801
Figc. 79-85
Habitus, edeago in visione laterale e ventrale e spermateca. 79-82: Atheta (Tropatheta) shimba-
ensis sp. n.; 83-85: Atheta (Tropatheta) savanicola sp. n.
802 ROBERTO PACE
FIGG. 86-91
Habitus, edeago in visione laterale e ventrale e spermateca. 86-89: Atheta (Tropatheta) loitensis
sp. n.; 90-91: Atheta (Tropatheta) embuicola sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 803
Descrizione. Lungh. 2,9 mm. Corpo lucido e bruno-rossiccio con uriti liberi 3°,
4° e 5° bruni; antenne bruno-rossicce con i due antennomeri basali giallo-rossicci;
zampe giallo-rossicce. La reticolazione dell'avancorpo è distinta, quella dell'addome
assente. Una fina e distinta punteggiatura è sul capo. Pronoto ed elitre sono coperti di
tubercoletti svaniti. Distinti sono invece i tubercoletti addominali. Spermateca fig. 91.
Comparazioni. L'habitus della nuova specie è molto simile a quello di A. gestroi
Bernhauer, 1915, dell'Eritrea, ma la spermateca è molto differente, per avere bulbo
distale non sferico come in gestroi e parte prossimale della stessa spermateca molto
allungata.
Atheta (Tropatheta) nanyukensis sp. n. Figg. 92-95)
Holotypus d, Kenya, env. Nanyuki, 1900 m, 22.X1.1974 (Mahnert & Perret leg., MG).
Paratypi: 6 es., stessa provenienza; 25 es., Kenya, Mau Escarpment, S-East Mau, 2850 m;
6.X1.1977 (Mahnert & Perret leg.); 7 es., Kenya, Taita Hills, env. Wundanyi, 1400 m, 2.XII.1974
(Mahnert & Perret leg.).
Descrizione. Lungh. 2,6 mm. Corpo lucido e bruno, con estremità addominale
bruno-rossiccia; antenne bruno-rossicce con i due antennomeri basali giallo-rossicci;
zampe giallo-brune con tarsi giallo-rossicci. La reticolazione del capo è vigorosa, quella
del pronoto è distinta, quella delle elitre è estremanente svanita e quella dell'addome è
assente. La punteggiatura del capo e delle elitre è distinta, quella del pronoto è
superficiale. Edeago figg. 93-94, spermateca fig. 95.
Comparazioni. La forma dell'edeago della nuova specie ricorda molto da vicino
quella dell'edeago di A. gestroi Bernhauer, 1915, dell'Eritrea, sia per dimensione che
per aspetto generale. Tuttavia la nuova specie presenta elitre più corte del pronoto e non
più lunghe del pronoto come in gestroi e l'edeago, in visione ventrale, ha apice arcuato,
mentre quello di gestroi è acuto.
Atheta (Tropatheta) mashonarum sp. n. (Figg. 96-99)
Holotypus d, Rhodésie, Umtali, II.1969 (R. Mussard leg., MG).
Paratypi: 12 es., stessa provenienza.
Descrizione: Lungh. 2,4 mm. Capo e pronoto opachi, resto del corpo lucido.
Corpo bruno-rossiccio con 4° urite libero bruno; antenne rossicce con i due antenno-
meri basali e la base del 3° giallo-rossicci; zampe giallo-rossicce. La reticolazione del
capo e del pronoto è vigorosa, quella delle elitre e dell'addome è distinta, trasversa
sull'addome. La punteggiatura del capo e del pronoto è assai superficiale. Tubercoletti
svaniti coprono le elitre. Edeago figg. 97-98, spermateca fig. 99.
Comparazioni. Le elitre lunghe quanto il pronoto, gli antennomeri 4° a 8° più
lunghi che larghi, l'edeago di dimensione molto ridotta e il bulbo distale della sperma-
teca di ridotta grandezza, distinguono la nuova specie da A. gestroi Bernhauer, 1915,
dell'Eritrea, che presenta elitre più lunghe del pronoto, antennomeri 7° e 8° trasversi,
l'edeago molto sviluppato e il bulbo distale della spermateca ipertrofico.
Etimologia. La nuova specie prende nome dai Mashona, popolazione stanziata
ad oriente dello Zimbabwe.
804 ROBERTO PACE
Fig. 92-99
Habitus, edeago in visione laterale e ventrale e spermateca. 92-95: Atheta (Tropatheta) nanyu-
kensis sp. n.; 96-99: Atheta (Tropatheta) mashonarum sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 805
Atheta (Taxicera) suprema Bernhauer, 1934 (Figg. 100-103 bis)
Atheta (Plataraea) suprema BERNHAUER, 1934: 244.
Materiale esaminato: Lectotypes d, Ruwenzori, 4200 m, VII.1932, Burgeon, Atheta
(Bessobia) suprema BRNH, typus, suprema BRNH, typus (Plataraea (Mus. Chicago), presente
designazione; paralectotypi: 5 es., stessa provenienza (Mus. Chicago).
1d el 9, Ruwenzori, Bujuko Hut, 1350 ft., 1.1984 (L. Lengeler leg.).
Nota. La presenza di una lunga setola su ciascun lembo della ligula (fig. 103
bis), permette di attribuire la specie preferibilmente a Taxicera Mulsant & Rey, 1873,
sottogenere che include specie presentanti due lunghe setole sulla ligula, mentre nel
sottogenere Plataraea Thomson, 1858, le specie sono prive di dette setole sulla ligula.
Specie già nota del Ruwenzori (località tipica) e della "Nyamgazani Valley"
(inedito del "Field Museum" di Chicago).
Rinvenuta su Senecio erioneuron (annotazione inedita del "Field Museum" di
Chicago).
Atheta (Taxicera) lacrymalis Fauvel, 1907
Atheta lacrymalis FAUVEL, 1907: 54.
Atheta (Acrotona) lacrymalis, BERNHAUER & SCHEERPELTZ, 1926: 675.
1 ©, Mt. Kenya, 3050 m, St. Lodge, 25.X1.1974 (Mahnert & Perret leg.).
Specie finora nota solo sull'holotypus 9 dell'Escarpment (Wa-Kikouyou), rac-
colto nell'agosto del 1904.
Esemplare comparato con l'holotypus. Grazie all'esame della ligula la specie va
attribuita al sottogenere Taxicera Mulsant & Rey, 1973. Infatti su ciascum lembo della
ligula è presente una lunga setola, carattere questo tipico del sottogenere Taxicera.
Atheta (Dimetrota) malevestita sp. n. (Figg. 104-107)
Holotypus 6, Rwanda, Kayove, 2100 m, 29.1V.1973 (Werner leg., MG).
Paratypi: 6 es., stessa provenienza.
Descrizione. Lungh. 2,1 mm. Corpo lucido e bruno-rossiccio con pronoto ed
estremità addominale rossicci; antenne brune con i due antennomeri basali giallo-
rossicci; zampe giallo-rossicce. La reticolazione è distinta, tranne sul pronoto dove è
svanita. La punteggiatura del capo è distinta. I tubercoletti che coprono il pronoto sono
superficiali, quelli delle elitre sono ben salienti. Edeago figg. 105-1067, spermateca fig.
107.
Comparazioni. Per la forma della spermateca e dell'edeago, la nuova specie è
ben distinta da A. convexula Eichelbaum, 1913, della Tanzania (figg. 118-121).
Atheta (Dimetrota) mandibularis sp. n. (Figg. 108-111)
Holotypus d, Rwanda, Kayove, 2100 m, 29.1V.1973 (Werner leg., MG).
Descrizione. Lungh. 2,0 mm. Corpo lucido e bruno con estremità addominale
bruno-rossiccia; antenne bruno-rossicce; zampe gialle. La reticolazione del capo è
assente, quella del pronoto è molto svanita, quella delle elitre e dell'addome è netta. Il
capo presenta punteggiatura svanita. Il pronoto e le elitre non mostrano distinti
tubercoletti. Edeago figg. 109-110, mandibola fig. 111.
806 ROBERTO PACE
Figc. 100-106
Habitus, edeago in visione laterale e ventrale, ligula e spermateca. 100-103: Atheta (Taxicera)
suprema Bernhauer, lectotypus d e paralectotypus 2; 104-106: Atheta (Dimetrota) malevestita
sp. n.
ALEOCHARINAE DELL’AFRICA ORIENTALE 807
Comparazioni. Specie che per la forma dell'edeago appare simile ad A. male-
vestita sp. n. sopra descritta. Si distingue per avere mandibole molto lunghe (fig. 111) e
acute (tozze in malevestita), per la netta reticolazione delle elitre, per l'11° antennomero
più lungo e per l'edeago più sviluppato.
Atheta (Datomicra) rudicollis (Bernhauer, 1915) (Figg. 112-115)
Oxypoda rudicollis BERNHAUER, 1915: 188.
Atheta (Datomicra) rudicollis, PACE, 1986: 109.
Materiale esaminato: Holotypus 2, Arusha, Ju, 1905.XII, Africa or., Katona, Oxypoda
rudicollis BRNH, typus (Mus. Budapest).
22 es., Kenya, Narok, Loita Hills, sous Morija, 5.XI.1977 (Mahnert & Perret leg.).
L'holotypus è qui illustrato per la prima volta.
Specie già nota della Tanzania e Zaire. Nuova per il Kenya.
Atheta (Datomicra) kiboensis Pace, 1985
Atheta (Datomicra) kiboensis Pace, 1985: 140.
1 ©, Kenya, Narok, Loita Hills, sous Morijo, 5.X1.1977 (Mahnert & Perret leg.).
Specie finora nota solo del Kilimangiaro.
Atheta (Datomicra) ferrugata Fauvel, 1907 (Figg. 116-117)
Atheta ferrugata FAUVEL, 1907: 53.
Atheta (Acrotona) ferrugata, BERNHAUER & SCHEERPELTZ, 1926: 673.
Atheta (Datomicra) ferrugata, PACE, 1986: 86.
Materiale esaminato: Holotypus ©, Afrique allemande, ferrugata FVL (Mus. Bruxelles).
La specie è qui illustrata per la prima volta.
Atheta (Dimetrota) convexula Eichelbaum, 1913 (Figg. 118-121)
Atheta (Coprothassa) convexula EICHELBAUM, 1913: 146.
Atheta (Acrotona) convexula, PACE, 1986: 109.
Materiale esaminato: Lectotypus é, Amani, Afheta convexula Eichelbaum, type, pre-
sente designazione (D.E.L.).
Atheta (Datomicra) ferrugatoides sp. n. (Figg. 122-123)
Holotypus 9, Kenya, Mau For., Mau Saumit Kedowa, 7.X1.1974 (Mahnert & Perret leg.,
MG).
i Paratypus: 1 2, Kenya, env. Endebess pr. Kitale, 14.X1.1974 (Mahnert & Perret leg.).
Descrizione. Lungh. 2,7 mm. Corpo lucido e bruno-rossiccio; antenne brune con
i due antennomeri basali gialli; zampe giallo-rossicce. I tubercoletti della superficie del
capo e delle elitre sono poco salienti, quelli del pronoto e dell'addome sono molto
salienti. Non vi è traccia di reticolazione sulla superficie del corpo. Spermateca fig. 123.
Comparazioni. Specie simile ad A. ferrugata Fauvel, 1907, della Tanzania (figg.
116-117). Se ne distingue soprattutto per i caratteri della spermateca. L'introflessione
apicale del bulbo distale della spermateca della nuova specie è corta, mentre in ferru-
gata è profonda e il bulbo prossimale della spermateca della nuova specie è nettamente
più largo di quello di ferrugata.
808 ROBERTO PACE
Fico. 107-115
Spermateca, habitus, edeago in visione laterale e ventrale e mandibola. 107: Atheta (Dimetrota)
malevestita sp. n.; 108-111: Atheta (Dimetrota) mandibularis sp. n.; 112-115: Atheta (Dato-
micra) rudicollis (Bernhauer), holotypus 2.
809
ALEOCHARINAE DELL’ AFRICA ORIENTALE
We
01 mm
120
Mo.
01 mm
Ficc. 116-121
Habitus, spermateca ed edeago in visione laterale e ventrale. 116-117: Atheta (Datomicra) ferru-
gata Fauvel, holotypus 2; 118-121: Atheta (Dimetrota) convexula Eichelbaum, lectotypus d .
810 ROBERTO PACE
Imm
FicG. 122-126
Habitus, spermateca ed edeago in visione laterale e ventrale. 122-123: Atheta (Datomicra)
ferrugatoides sp. n.; 124-126: Atheta (Acrotona) semifura Cameron, esemplare della Rodesia.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 811
Atheta (Phanerosphaena) kenyensis Pace, 1985
Atheta kenyensis PACE, 1985: 149.
2346 e4 99, Kenya, Narok, Loita Hills, 4.X1.1977, Morijo, 2200 m (Mahnert & Perret
leg.); 1 d el 2, Kenya, Embu, pr. Ishiara, 900 m, 14.X.1977 (Mahnert & Perret leg.).
Specie gia nota solo dei dintorni di Nairobi.
Atheta (Acrotona) parasita Bernhauer, 1945
Atheta (Acrotona) parasita BERNHAUER, 1945: 19; Pace, 1986: 86.
16 el ©, Kenya, Mt. Elgon, 2800 m, 14.X1.1974 (Mahnert & Perret leg.).
Specie già nota del M. Meru e dell'Elgon.
Atheta (Acrotona) nigricola Pace, 1986
Atheta (Acrotona) nigricola PACE, 1986: 100.
1 4, Kenya, Lac Nakuru, Parc Nat., 6.XI.1974 (Mahnert & Perret leg.); 1 4, Rhodésie,
Umtali, 11.1969 (R. Mussard leg.).
Specie nota dell'Abissinia, Tanzania, Natal e Citta del Capo. Nuova per il
Kenya.
Atheta (Acrotona) armentorum Pace, 1985
Atheta (Acrotona) armentorum PACE, 1985: 142.
20 es., Kenya, Narok, Loita Hills, sous Morijo, 5.X1.1977 (Mahnert & Perret leg.); 12 es.,
Kenya, Narok, près de Narok, 2000 m, 3.XI.1977 (Mahnert & Perret leg.); 1 2, Kenya, Lac
Nakuru, 6.X1.1974 (Mahnert & Perret leg.); 14 es., Kenya, Kikuyu Escar., 2000 m, 3.X1.1977
(Mahnert & Perret leg.); 1 ©, Kenya, Tambach, E Eldoret, 2000 m, 17.X1.1974 (Mahnert &
Perret leg.); 2 2 2, Rwanda, Kayove, 2100 m, 15.V.1973 (Werner leg.).
La specie finora era nota solo del M. Meru.
Atheta (Acrotona) semirufa Cameron, 1950 (Figg. 127-129)
Atheta (Acrotona) semirufa Cameron, 1950: 64.
Materiale esaminato: Paratypus d, Congo Belge, P.N.A., Kanyabayondo (Kabasha),
1760 m, 7.XII.1934, G. F. de Witte: 878, A. rufonigra Cam. (cancellato), A. (Acrotona) semirufa
n. n. Cam. (British Museum).
3 d d, Rhodésie, Umtali, 11.1969 (R. Mussard leg.)
Specie finora nota solo dello Zaire. Esemplari comparati con un syntypo.
Edeago e habitus qui illustrati per la prima volta.
Atheta (Acrotona) tugen sp. n. (Figg. 130-132)
Holotypus d, Kenya, Mt. Elgon, 2400 m, Mt. Elgon For. Res., 14.X1.1974, Mahnert &
Perret leg., MG).
Descrizione. Lungh. 2,4 mm. Corpo lucido e rossiccio scuro; antenne bruno-
rossicce con i due antennomeri basali giallo-rossicci; zampe giallo-rossicce. La
superficie del corpo non è reticolata ed è coperta di tubercoletti fitti e poco salienti.
Edeago figg. 131-132.
Comparazioni. Specie simile ad A. basipennis Fauvel, 1907, della Tanzania, a
motivo della forma dell’edeago e dell’habitus. Se ne distingue tra l’altro per avere
l’apice dell’edeago diviso (in basipennis apice intero).
Etimologia. La nuuova specie prende nome dai Tugen, gruppo etnico del Kenya.
812 ROBERTO PACE
Olmm
128 120)
01 mm
face
132 ne
131
Ficc. 127-132
Habitus ed edeago in visione laterale e ventrale. 127-129: Atheta (Acrotona) semirufa Cameron,
paratypus d; 130-132: Atheta (Acrotona) tugen sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 813
Atheta (Acrotona) creticornis sp. n. (Figg. 133-136)
Holotypus 4 , Rwanda, Kayove, 2100 m, 15.V.1973, (Werner leg., MG).
Paratypi: 3 d d, stessa provenienza; 2 dd e 1 2, Rwanda, Rangiro, 1800 m, 6. VIII.1973,
(Werner leg.).
Descrizione. Lungh. 2,5 mm. Corpo lucido e bruno-rossiccio; antenne bruno-
rossicce con base del primo antennomero gialla; zampe gialle. La reticolazione del capo
e del pronoto è distinta, quella delle elitre e dell'addome è assente. I tubercoletti che
coprono la superficie del capo e delle elitre sono poco distinti, quelli del pronoto sono
salienti. Edeago figg. 134-135, spermateca fig. 136.
Comparazioni. In base alla forma della spermateca, la nuova specie sembra
affine ad A. kibatiana Cameron, 1950, dello Zaire, ma la nuova specie ha la spermateca
nettamente più sviluppata e l’edeago ha apice a foggia di spatola, in visione ventrale
(stretto in kibatiana).
Etimologia. Il nome della nuova specie significa «Antenne cresciute».
Atheta (Acrotona) promissionum sp. n. (Figg. 137-138)
Holotypus ©, Kenya, env. Nanyuki, 1900 m, 22.X1.1974, (Mahnert & Perret leg., MG).
Descrizione. Lungh. 2,5 m. Corpo lucido e bruno, comprese le antenne; zampe
giallo-rossicce. La reticolazione del capo è svanita, quella del pronoto assente e quella
delle elitre è distinta. Tubercoletti nettamente salienti coprono l’avancorpo. Spermateca
fig. 138.
Comparazioni. Specie affine alla precedente A. creticornis sp. n. in base alla
forma della spermateca. Ne differisce perché la parte prossimale delle spermateca
descrive una sola spira (una spira e mezza in creticornis), le elitre sono coperte di
reticolazone distinta nella nuova specie, mentre in creticornis le elitre sono prive di
reticolazione e il 4° antennomero della nuova specie è più lungo che largo, mentre
quello corrispondente di creticornis è trasverso.
Atheta (Acrotona) zimbabwensis sp. n. (Figg. 139-142)
Holotypus d, Rhodésie, Umtali, I1.1969, (R. Mussard leg. MG).
Paratypi: 13 es., stessa provenienza.
Descrizione. Lungh. 2,4 mm. Corpo lucido e giallo-rossiccio con capo, elitre e
uroterghi rossicci; margine posteriore degli uroterghi giallo-rossiccio; antenne bruno-
rossicce con i due antennomeri basali giallo-rossicci; zampe giallo-rossicce. La
reticolazione del capo è distinta, ma assente sul disco; assente pure sul pronoto e sulle
elitre; la punteggiatura del capo è distinta e fitta, ma assente sul disco. Il pronoto e le
elitre sono coperti di tubercoletti salienti. Edeago figg. 140-141, spermateca fig. 142.
Comparazioni. Per la forma dell’edeago e per l’habitus, la nuova specie può
essere sistematicamente avvicinata ad A. parasita Bernhauer, 1845, del Kenya, ma
l’ultimo antennomero del maschio della nuova specie è chiaramente più corto del
corrispondente di parasita; l’edeago è più largo e ha lati convergenti verso l’apice nella
nuova specie, mentre in parasita 1 lati sono tra loro paralleli e la spermateca è priva di
introflessione apicale del bulbo distale (robusta e profonda in parasita).
814 ROBERTO PACE
|
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PAR
FiGG. 133-138
Habitus, edeago in visione laterale e ventrale e spermateca. 133-136: Atheta (Acrotona) creti-
cornis sp. n.; 137-138: Atheta (Acrotona) promissionum sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 815
142
01 mm
144
Ficc. 139-145
Habitus, edeago in visione laterale e ventrale e spermateca. 139-142: Atheta (Acrotona) zimbab-
wensis sp. n.; 143-145: Atheta (Acrotona) umtaliensis sp. n.
816 ROBERTO PACE
Atheta (Acrotona) umtaliensis sp. n. (Figg. 143-145)
Holotypus d, Rhodésie, Umtali, II. 1969, (R. Mussard leg., MG).
Descrizione. Lungh. 2,3 mm. Corpo lucido e bruno, comprese le antenne; zampe
di un giallo sporco. La reticolazione del capo è molto svanita, quella del pronoto è
distinta e quella delle elitre e dell'addome è netta. La punteggiatura del capo è fine, rada
e superficiale. Il pronoto presenta tubercoletti distinti e le elitre hanno tubercoletti netti.
Edeago figg. 144-145.
Comparazioni. Specie con habitus simile a quelle di A. parasita Bernhauer,
1945, del Kenya, ma la taglia è lievemente minore (2,3 mm invece di 2,5 mm) et 111°
antennomero del maschio non è lunghissimo come quello corrispondente in parasita.
L’edeago ha caratteri differenziali più accentuati: tra l’altro la regione preapicale
dell’edeago della nuova specie è fortemente ristretta, in visione ventrale, mentre la
corrispondente porzione dell’edeago di parasita è molto larga e a lati paralleli.
Atheta (Acrotona) nyeriensis sp. n. (Figg. 146-149)
Holotypus 4, Kenya, Mt. Aberdares, Parc National, 2300 m, 25.X1.1974, (Mahnert &
Perret leg., MG).
Paratypi: 2 6 d el 9, stessa provenienza.
Descrizione. Lungh. 2,7 mm. Corpo lucido e bruno; antenne brune con i due
antennomeri basali gialli; zampe giallo-rossicce. La reticolazione è distinta sul capo e
sugli uroterghi, è molto svanita sulle elitre e assente sul pronoto. L’intero corpo è
coperto di tubercoletti fitti e distinti. Edeago figg. 147-148, spermateca fig. 149.
Comparazioni. Per la forma della spermateca, è possibile che la nuova specie sia
affine ad A. parasita Bernhauer, 1945, tuttavia l’habitus della nuova specie è chiara-
mente più snello di quello di parasita e l’edeago ha parte apicale cortissima, mentre
quella di parasita è lunga.
Etimologia. La nuova specie prende nome dalla città di Nyeri, la più vicina al
M. Aberdare, località tipica.
Atheta (Acrotona) kawaensis Cameron, 1932 (Figg. 150-151)
Atheta (Acrotona) kawaensis CAMERON, 1932: 144.
Materiale esaminato: Paratypus 9, Forêt de Kawa, 23.1V.1929, A. Collart, Atheta ka-
waensis.
Habitus e spermateca qui illustrati per la prıma volta.
Atheta (Acrotona) embuorum sp. n. (Figg. 152-153)
Holotypus 2, Kenya, Embu, près Kogari, 800 m, 15.X.1977. (Mahnert & Perret leg.,
MG).
Descrizione. Lungh. 2,5 mm. Corpo lucido e bruno-rossiccio con uriti liberi 3°,
4° e 5° bruni; antenne bruno-rossicce con i due antennomeri basali giallo-rossicci;
zampe giallo-rossicce con femori bruno-rossicci. La reticolazione del capo e del pro-
noto è netta e fine, quella delle elitre è distinta, quella dell’addome assente. La pun-
teggiatura del capo è superficiale e assente sul disco, quella del pronoto è svanita e
quella delle elitre è distinta. Spermateca fig. 153.
ALEOCHARINAE DELL’ AFRICA ORIENTALE
817
01 mm
o
a
3
©)
Ficc. 146-151
Habitus, edeago in visione laterale e ventrale e spermateca. 146-149: Atheta (Acrotona) nyerien-
sis sp. n.; 150-151: Atheta (Acrotona) kawaensis Cameron, paratypus 9.
818 ROBERTO PACE
Comparazioni. In base alla forma della spermateca, la nuova specie potrebbe
essere affine ad A. kawaensis Cameron, 1932, (figg. 150-151) dello Zaire, ma l’habitus
non è simile, dato che l'addome è divergente all’indietro, per cui l’attribuzione sotto-
generica è dubbia.
Etimologia. Il nome della nuova specie deriva dagli Embu, gruppo etnico
famoso per le danze.
Atheta (Xenota) interrogativa Pace, 1985
Atheta (Xenota) interrogativa PACE, 1985: 144.
2 9 9, Kenya, Narok,, Loita Hills, 2050 m, 5.X1.1977, (Mahnert & Perret leg.); 9 es.,
Kenya, Nairobi, 2-3.XI.1974, (Mahnert & Perret leg.); 1 9, Kenya, Embu, Kirimiri For.,
13.X.1977, (Mahnert & Perret leg.); 25 es., Nenya, Miss. Kaimosi, NE Kisumu, 11.XI.1974,
(Mahnert & Perret leg.); 1 6, Kenya, Tambach, E Eldoret, 2000 m, 17.X1.1974, (Mahnert &
Perret leg.).
Specie già nota del Kenya meridionale: Kajıdo.
Atheta (Xenota) riftensis Fauvel, 1907 (Figg. 154-156)
Atheta riftensis FAUVEL, 1907: 54.
Atheta (Xenota) riftensis, PACE, 1986: 87.
1 d, Kenya, Tambach, E. Eldoret, 2000 m, 17.X1.1974, 2000 m, (Mahnert & Perret leg.);
3 es.; Kenya, Mt. Aberdares, 2300 m, 25.XI.1974, (Mahnert & Perret leg.); 14 es., Kenya,
Nairobi, 2-3.X1.1974, (Mahnert & Perret leg.); 38 es., Kenya, Embu, Kirimiri For., 1550 m,
13.X.1977, (Mahnert & Perret leg.); 1 es., Kenya, Embu, Irangi For., 2000 m, 11.X.1977,
(Mahnert & Perret leg.); 1 ¢, Kenya, Embu, 20 Km 0 Ishiara, 900 m, 14.X.1977, (Mahnert &
Perret leg.); 22 es., Kenya, Kiambu, Kikuyu Escarp., 2000 m, 3.X1.1977, (Mahnert & Perret leg.);
1 ©, Kenya, Lac Nakuru, pres dee Mundu Estate, 1950 m, 8.X11.1977, (Mahnert & Perret leg.); 6
es., Kenya, env. Endebess, pr. Kitale, 14.X1.1974, (M. Perret leg.); 3 es., Rhodésie, Umtali,
11.1969, (R. Mussard leg.).
Specie largamente diffusa nella Tanzania e nel Kenya. Nuova per la Rhodesia.
Atheta (Xenota) coriaria (Kraatz, 1858)
Homalota coriaria KRAATZ, 1858: 282.
Atheta (Xenota) coriaria, PACE, 1984: 263.
1 2, Kenya, Nairobi, 3.XI.1974, (Mahnert & Perret leg.).
Specie cosmopolita.
Atheta (Xenota) kitalensis sp. n. (Figg. 157-160)
Holotypus 4 , Kenya, Mt. Elgon, 2700-2800 m, 15.X1.1974, (Mahnert & Perret leg., MG).
Paratypi: 8 es., stessa provenienza, ma anche 14.X1.1974.
Descrizione. Lungh. 3,8 mm. Corpo lucido e bruno con margini laterali del pro-
noto bruno-rossicci e margine posteriore degli uroterghi liberi 1°, 2. et 3° ed estremità
addominale giallo-rossicci; antenne brune con i due antennomeri basali giallo-rossicci;
zampe giallo-rossicce. Tutto il corpo è coperto di reticolazione netta. I tubercoletti della
superficie del capo e del pronoto sono distinti, quelli delle elitre sono svaniti. Edeago
figg. 158-159, spermateca fig. 160.
Comparazioni. Per la forma dell’edeago, la nuova specie sembra più affine ad A.
scotti Bernhauer, 1931 che ad A. euphorbiae Bernhauer, 1931, entrambe dell’ Abissinia.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 819
Of mm
Fiss. 152-156
Habitus, spermateca ed edeago in visione laterale e ventrale. 152-153: Atheta (Acrotona) embu-
orum sp. n.; 154-156: Atheta (Xenota) riftensis Fauvel.
820
ROBERTO PACE
01mm
01 mm
sa
Vi
e
Imm
RANA x NE
h D
| | S À
OBS LEA
CRU PAM NUE SS ;
162
Ficc. 157—162
Habitus, edeago in visione laterale e ventrale e spermateca. 157-160: Atheta (Xenota) kitalensis
sp. n.: 161-162: Atheta (Xenota) cincta sp. n.
OALEOCHARINAE DELL’AFRICA ORIENTALE 821
La nuova specie differisce da A. scotti perché ha l’apice dell’edeago smussato e più
largo (apice acuto in scotti), la «crista apicalis» poco sviluppata (molto sviluppata in
scotti), i pezzi copulatori del sacco interno molto sviluppati e robusti (poco sviluppati
ed esili in scotti), la parte prossimale della spermateca esile e non descrivente regolari
spire come in scotti e il penultimo antennomero più lungo che largo (trasverso in
scotti).
Etimologia. La nuova specie prende nome da Kitale, capoluogo del distretto
agricolo di Trans-Nzoie situato presso il M. Elgon e punto di partenza per una visita al
parco nazionale del M. Elgon.
Atheta (Xenota) cincta sp. n. (Figg. 161-162)
Holotypus 9, Rhodésie, Umtali, II.1969, (R. Mussard leg., MG).
Descrizione. Lungh. 2,2 mm. Corpo lucido e giallo-rossiccio con 4° urotergo
libero bruno; antenne bruno-rossicce con 1 tre antennomeri basali e 1’11° giallo-
rossicci; zampe gialle. La reticolazione del capo, del pronoto e dell’addome è molto
svanita, quella delle elitre è assente. Tutto il corpo è coperto di tubercoletti più o meno
salienti. Spermateca fig. 162.
Comparazioni. Solo un’altra specie del sottogenere Xenota Mulsant & Rey,
1874, presenta elitre molto accorciate e pronoto molto trasverso come nella nuova
specie: A. humidula Pace, 1985, delle Tanzania. La nuova specie si distingue da essa
per avere spermateca esile e più allungata rispetto quella di humidula e 1°11°
antennomero della femmina nettamente pit lungo nella nuova specie che in humidula.
Atheta (Xenota) tanensis sp. n. (Figg. 163-164)
Holotypus 9, Kenya, Tana River, Galole Hola, 60 m, 20.X.1977, (Mahnert & Perret leg.,
MG).
Descrizione. Lungh. 2,1 mm. Corpo lucido e bruno con elitre giallo-brune aventi
margini laterali, posteriore e suturale bruni; antenne brune con i due antennomeri basali
gialli; zampe gialle. La reticolazione del capo e del pronoto è distinta, quella delle elitre
è netta, quella dell'addome a maglie molto trasverse e superficiali. I tubercoletti della
superficie del capo sono distinti, quelli del pronoto sono fini e salienti, quelli delle elitre
sono svaniti. Spermateca fig. 164.
Comparazioni. Per la struttura della spermateca, la nuova specie sembra affine
ad A. riftensis Fauvel, 1907, della Rift-Valley, ma la parte prossimale della spermateca
non è sottile, nè descrive un’ampia spira come in riftensis. Inoltre il pronoto della nuova
specie è molto più trasverso di quello di riftensis e le elitre più larghe e più lunghe.
Atheta (Oxypodera) densiventris Fauvel, 1907 (Figg. 165-168)
Atheta densiventris FAUVEL, 1907: 54.
Atheta (s. str.) densiventris, BERNHAUER & SCHEERPELTZ, 1926: 64.
Materiale esaminato: Holotypus d, Afrique or. allemande, densiventris FVL, type (Mus.
Bruxelles).
1 d, Kenya, Nairobi, 3.X1.1974, Mahnert & Perret leg.).
822 ROBERTO PACE
FIGG. 163-168
Habitus, spermateca, edeago in visione laterale e ventrale e 6° urotergo libero del maschio.
163-164: Atheta (Xenota) tanensis sp. n.; 165-168: Atheta (Oxypodera) densiventris Fauvel,
holotypus.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 823
Habitus, edeago e quinto urotergo libero del maschio qui per la prima volta
illustrati.
Specie finora nota su un maschio di località imprecisata dell’ Africa orientale.
Atheta (Oxypodera) andreinii Bernhauer, 1927 (Figg. 169-172)
Atheta (Acrotona) andreinii Bernhauer, 1927: 80.
Materiale esaminato: Lectotypus 4, Eritrea, Adi-Caie, 10.1V.1902, Dr. Andreini, 243,
Atheta andreinii BRNH, Typ., presente designazione, (Mus. La Specola, Firenze, N° 7789);
paralectotypi: 35 es., stessa provenienza e Museo.
Nota. La spermateca e l’edeago di A. andreinii indicano senza dubbio l’appar-
tenenza della specie al sottogenere Oxypodera Bernhauer, 1915 e non al sottogenere
Acrotona Thomson, 1859.
La specie è qui illustrata per la prima volta. Essa è estremamente affine ad A.
densiventris Fauvel, 1907, sopra illustrata. I caratteri differenziali sono esposti nella
seguente chiave:
1 Occhi sporgenti; 11° antennomero del maschio lungo come 1 due pre-
cedenti antennomeri considerati insieme; reticolazione del pronoto sva-
nita; elitre più larghe; 6° urotergo libero del maschio dentellato al
margine posteriore (fig. 168); apice dell’edeago pit stretto, in visione
ventrales Luneha2,2 mm. Aimca onentale "7 densiventris Fauvel
— Occhi meno sporgenti; 11° antennomero del maschio lungo come i tre
precedenti antennomeri considerati insieme; reticolazione del pronoto
estremamente svanita; elitre piuttosto strette: 6° urotergo libero del
maschio con margine posteriore non dentellato; apice dell’edeago più
largo, in visione ventrale. Lungh. 2,7 mm. Eritrea. ....... andreinii Bernhauer
Atheta (Oxypodera) chyuluensis Cameron, 1942 (Figg. 173-176)
Atheta (Acrotona) chyuluensis CAMERON, 1942: 331.
Atheta (Xenota) chyuluensis, PACE, 1986: 87.
Materiale esaminato: Lectotypus 3, Coryndon Museum Expedit., Chyulu Hills, june
38, alt. 5660, Atheta chyuluensis Cam., presente dseignazione (Brit. Mus.); paralectotypi: 2 4 d
e 2 2 2, stessa provenienza, (Brit. Mus.).
1 2, Kenya, Nairobi, 3.X1.1974, (Mahnert & Perret leg.); 1 4, Kenya, Narok, Loita Hills,
4.XI.1977, (Mahnert & Perret leg.); 3 dd, Kenya, Tambach, E Eldoret, 2000 m, 17.X1.1974
(Mahnert & Perret leg.).
Specie diffusa dal Kenya alla Tanzania, la specie € qui illustrata per la prima
volta.
Atheta (Oxypodera) nairobianorum Pace, 1985
Atheta (Oxypodera) nairobianorum Pace, 1985: 144.
1 2, Kenya, Embu, Kirimiri Forest, 1550 m, 13.X.1977, (Mahnert & Perret leg.); 1 ©,
Kenya, Nakuru, Mau Escarpment, 2700 m, 6.XI.1977, (Mahnert & Perret leg.).
Specie gia nota del Kenya.
ROBERTO PACE
824
Fig. 169-176
Habitus, edeago in visione laterale e ventrale e spermateca. 169-172: Atheta (Oxypodera)
andreinii Bernhauer, lectotypus 6 e paralectotypus 2; 173-176: Atheta (Oxypodera) chyuluensis
Cameron, lectotypus ¢ e paralectotypus °.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 825
Atheta (Oxypodera) fugata Pace, 1985
Atheta (Oxypodera) fugata PACE, 1985: 146.
10 es., Mt. Kenya, 3250 m, S Met St. Lodge, 23.X1.1974, (Mahnert & Perret leg.).
Specie finora nota solo del M. Meru.
Atheta (Oxypodera) mombasana Bernhauer, 1934 (Figg. 177-179)
Atheta (Oxypodera) mombasana BERNHAUER, 1934: 246.
Atheta (Oxypodera) mombasana, PACE, 1986: 109, sp. bona.
Materiale esaminato: Lectotypus 4, Mombasa, 36 Km Sud Lubero, fin VII.1932, L.
Burgeon, mombasana BRNH, Type, mombasana BRNH, Type, presente designazione (Mus.
Chicago).
Specie qui illustrata per la prima volta.
Nota. Per un mio errore di trascrizione, nel 1986 ho posto in sinonimia di Atheta
paludosa Bernhauer, 1931, A. mombasana Bernhauer, 1934, in luogo di A. mom-
bassana Bernhauer, 1932. Pertanto stabilisco la corretta sinonima come segue:
Atheta (Oxypodera) paludosa Bernhauer, 1931
Atheta (Coprothassa) paludosa BERNHAUER, 1931: 602.
Atheta (Coprothassa) mombassana BERNHAUER, 1934: 214, syn. n. (nec Atheta mom-
basana BERNHAUER, 1934: 246).
Atheta (Oxypodera) pseudomombasana sp. n. (Figg. 180-182 e 198-194)
Holotypus d, Kenya, Nairobi, 3.X1.1974, (Mahnert & Perret leg., MG).
Paratypus: 1 2, stessa provenienza.
Descrizione. Lungh. 3,0 mm. Corpo lucido e bruno-rossiccio; antenne brune con
i due antennomeri basali giallo-rossicci; zampe giallo-rossicce. La reticolazione del
capo, delle elitre e dell'addome è distinta, quella del pronoto è assente. I tubercoletti
sono distinti o salienti su tutto il corpo. Edeago figg. 181-182.
Comparazioni. Specie affine ad A. mombasana Bernhauer, 1934, per la forma
dell’edeago, ma quello della nuova specie ha taglia minore (figg. 178-179 e 181-182),
non è profondamente arcuato al lato ventrale come quello di mombasana. Inoltre gli
occhi della nuova specie sono più sporgenti e i denti del margine posteriore del 6°
urotergo libero del maschio sono più distanziati tra loro nella nuova specie che in
mombasana.
Atheta (Oxypodera) kayovensis sp. n. (Figg. 183-186)
Holotypus d, Rwanda, Kayove, 2100 m, 15.V.1973, (Werner leg., MG).
Paratypi: 2 9 ©, stessa provenienza; | d e 1 2, Rwanda, Rangiro, 1800 m, 20.VII.1976,
(Werner leg.).
Descrizione. Lungh. 3,0 mm. Corpo lucido e giallo rossiccio con capo e uriti
liberi 3°, 4° e 5° bruni ed elitre rossicce; antenne bruno-rossicce, con i due antennomeri
basali giallo-rossicci; zampe gialle. La reticolazione del corpo è distinta, tranne sulle
elitre dove è molto svanita. La punteggiatura del capo è distinta. I tubercoletti sul pro-
noto sono netti, quelli sulle elitre sono superficiali. Edeago figg. 184-185, spermateca
fig. 186.
826 ROBERTO PACE
Fico. 177-182
Habitus ed edeago in visione laterale e ventrale. 177-179: Atheta (Oxypodera) mombasana
Bernhauer, lectotypus d ; 180-182: Atheta (Oxypodera) pseudomombasana sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 827
Comparazioni. Alla grande taglia del corpo di questa specie non corrisponde un
edeago proporzionalmente grande, come è osservabile in tutte le specie del sottogenere
Oxypodera Bernhauer, 1915. Una sola specie possiede edeago di taglia uguale a quello
della nuova specie: A. haramayana Bernhauer, 1931, dell’ Abissinia, ma haramayana
ha corpo di lunghezza minore (2,1 mm). Inoltre nella nuova specie è assente la “crista
apicalis”, mentre in haramayana è presente e il sacco interno dell’edeago presenta una
formazione chitinosa rivestita di setole nella nuova specie, mai osservata in altre specie
del sottogener Oxypodera.
Atheta (Oxypodera) nakuruensis sp. n. (Figg. 187-189)
Holotypus 4 , Kenya, lac Nakuru, Parc Nat., (Mahnert & Perret leg., MG).
Descrizione. Lungh. 2,7 mm. Corpo lucido e rossiccio con capo, elitre e uriti
liberi 1°, 2°, 3° e 4° bruno-rossicci; antenne rossicce con i due antennomeri basali e
1°11° giallo-rossicci; zampe giallo-rossicce. La reticolazione del capo è superficiale,
quella del pronoto e dell'addome è assente, quella delle elitre è netta. Tutto il corpo è
coperto di tubercoletti distinti. Edeago figg. 188-189.
Comparazioni. La nuova specie presenta elitre più corte del pronoto. Specie del
sottogenere Oxypodera Bernhauer, 1915, con questo stesso carattere, sono da citare A.
kenyamontis Pace, 1986, A. montanella Bernhauer, 1934, del Kivu, A. burgeoniana
Bernhauer, 1934, del Ruwenzori e A. complicans Pace, 1985, del Kilimangiaro.
Nessuna di esse ha edeago identico a quello della nuova specie. Prendendo in esame il
solo apice dell’edeago, in visione ventrale, A. kenyamontis lo ha tronco, A. burgeoniana
e A. complicans lo hanno acuto. La sola specie che presenta l’apice dell’edeago smus-
sato come quello della nuova specie, è A. montanella, ma questa specie ha edeago
molto più sviluppato, con pezzi copulatori del sacco interno che sembrano molto
differenti (purtroppo l’edeago dell’holotypus di montanella ha il sacco interno eva-
ginato). Inoltre habitus di montanella è stretto, le antenne sono proporzionalmente più
lunghe e gli occhi sono sporgenti.
Atheta (Oxypodera) kaimosensis sp. n. (Figg. 190-193)
Holotypus d, Kenya, Miss. Kaimosi, NE Kisumu, 1650 m, 11.X1.1974, (Mahnert &
Perret leg., MG).
Paratypi: 5 es., stessa provenienza.
Descrizione. Lungh. 2,3 mm. Corpo lucido e rossiccio con pronoto giallo-
rossiccio e uriti liberi 2°, 3°, 4° e 5° bruni; Antenne brune con i due antennomeri basali
giallo-rossicci; zampe giallo-rossicce. La reticolazione del capo è distinta, quella del
pronoto e dell’addome assente, quella delle elitre è svanita. La punteggiatura del capo è
distinta. Tubercoletti fini e distinti coprono il resto del corpo. Edeago figg. 191-192,
spermateca fig. 193.
Comparazioni. La nuova specie per molti caratteri dell’edeago e della sper-
mateca si presenta sistematicamente vicina ad. A. chyuluensis Cameron, 1942 (figg.
173-176), del Kenya e della Tanzania. Infatti il sacco interno dell’edeago delle due
specie mostra due lame chitinose ricurve e a punta acuta, ma mentre in chyuluensis una
828 ROBERTO PACE
01 mm
184 185
à 7 Bere
Imm
Fico. 183-189
Habitus, edeago in visione laterale e ventrale e spermateca. 183-186: Atheta (Oxypodera) kayo-
vensis sp. n.; 187-189: Atheta (Oxypodera) nakuruensis sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 829
è lunghissima e l’altra cortissima e tozza, nella nuova specie dette lame chitinose sono
simmetriche, con base molto larga. La spermateca ha bulbo distale molto stretto in
chyuluensis, largo nella nuova specie.
Atheta (Oxypodera) rangirensis sp. n. (Figg. 194-197)
Holotypus d, Rwanda, Rangiro, 1800 m, 6.VIIL.1973, (Werner leg., MG).
Descrizione. Lungh. 2,7 mm. Corpo lucido e bruno con elitre giallo-brune;
antenne bruno-rossicce con i due antennomeri basali giallo-rossicci; zampe giallo-
rossicce. Su tutto il corpo la reticolazione è distinta. La punteggiatura del capo è
evidente. I tubercoletti che coprono pronoto ed elitre sono svaniti. Edeago figg. 196-
197, 6° urotergo libero del maschio fig. 194.
Comparazioni. L’edeago delle specie del sottogenere Oxypodera Bernhauer,
1915, ha la regione della “crista apicalis” per lo più poco sporgente al lato ventrale. Fa
eccezione l’edeago della nuova specie che ha la regione della “crista apicalis” visto-
samente sporgente. Per questo carattere e per altri, come l’apice dell’edeago in visione
ventrale con lati preapicali paralleli e il 3° e il 4° degli antennomeri compressi lateral-
mente, la nuova specie è chiaramente distinta dalle altre note del sottogenere.
Atheta (Oxypodera) implicatoides sp. n. (Figg. 200-201)
Holotypus 9, Rhodésie, Umtali, II. 1969, (R. Mussard leg., MG).
Descrizione. Lungh. 2,9 mm. Corpo lucido e bruno-rossiccio con uriti liberi 4° e
5° bruni; antenne brune con i tre antennomeri basali giallo-rossicci; zampe giallo-
rossicce. La reticolazione del capo è distinta, quella del pronoto e dell’addome è sva-
nita, quella delle elitre è vigorosa. La punteggiatura del capo è distinta, quella del
pronoto è svanita e quella delle elitre è netta. Spermateca fig. 201.
Comparazioni. Specie simile ad A. implicata Pace, 1985, del Kilimangiaro, ma
la spermateca ha parte prossimale molto sottile (parte prossimale larga in implicata). La
nuova specie è pure distinta da A. pseudomombasana sp. n. sopra descritta perché la
parte prossimale della spermateca non è foggiata a spirale.
Atheta (Oxypodera) mahnerti sp. n. (Figg. 202-205)
Holotypus 4, Kenya, Narok, Loita Hills, SO Morijo, 2050 m, 4-5.X1.1977, (Mahnert &
Perret leg., MG).
Paratypi: 14 es., stessa provenienza; 3 es., Kenya, Thomson’s Falls, 2350 m, 20.X1.1974,
(Mahnert & Perret leg.); 1 d e 2 9 ©, Kenya, Nakuru, Mau Escarp. près Enangiperi, 2700 m,
6.X1.1977, (Mahnert & Perret leg.).
Descrizione. Lungh. 2,9 mm. Corpo lucido e bruno-rossiccio con capo e uriti
liberi 3°, 4° e 5° bruni; antenne bruno-rossicce con antennomero basale giallo-rossiccio;
zampe giallo-rossicce. La reticolazione del capo è svanita, quella del pronoto molto
superficiale e quella delle elitre e dell’addome é netta. La punteggiatura del capo é
svanita. I tubercoletti che coprono il pronoto sono distinti e quelli delle elitre poco
salienti. Edeago figg. 204-205, spermateca fig. 203.
Comparazioni. La nuova specie, per la forma dell’edeago e della spermateca, è
sistematicamente vicina ad A. paludosa Bernhauer, 1931, dell’ Abissinia, ma un robus-
830 ROBERTO PACE
Fico. 190-197
Habitus, edeago in visione laterale e ventrale, spermateca e 6° urotergo libero del maschio.
190-193: Atheta (Oxypodera) kaimosensis sp. n.; 194-197: Atheta (Oxypodera) rangirensis sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 831
Figc. 198-203
Habitus e spermateca. 198-199: Atheta (Oxypodera) pseudomombasana sp. n.; 200-201: Atheta
(Oxypodera) implicatoides sp. n.; 202-203: Atheta (Oxypodera) mahnerti sp. n.
832 ROBERTO PACE
tissimo pezzo copulatore del sacco interno dell’edeago della nuova specie e la parte
apicale dell’edeago della nuova specie chiaramente stretto, sono alcuni caratteri dif-
ferenziali più evidenti. La spermateca della nuova specie è maggiore, con introflessione
apicale del bulbo distale largo, robusto e fortemente slerificato (non sclerificato in
paludosa).
Atheta (Oxypodera) atopotheca sp. n. (Figg. 206-209)
Holotypus d, Kenya, Kiambu, près de Limuru, 2300 m, 2.X1.1977, (Mahnert & Perret
leg., MG).
Paratypi: 2 dd el 9, stessa provenienza.
Descrizione. Lungh. 3,2 mm. Corpo lucido e bruno-rossiccio con capo e uriti
liberi 3°, 4° e 5° bruni; antenne brune con i due antennomeri basali giallo- rossicci;
zampe giallo-rossicce. La reticolazione del capo e dell’addome è distinta, quella del
pronoto è svanita e quella delle elitre è netta. La punteggiatura del capo è distinta. I
tubercoletti del pronoto sono molto salienti, quelli delle elitre sono distinti. Edeago
figg. 207-208, spermateca fig. 209.
Comparazioni. La nuova specie risulta straordinaria e unica nel numero delle
specie del sottogenere Oxypodera Bernhauer, 1915, a motivo dell’eccezionale forma
allungata del bulbo distale della spermateca. Anche l’allungatissimo 11° antennomero
distingue la nuova specie dalle altre specie del sottogenere.
Etimologia. Il nome della nuova specie significa “Spermateca assurda”.
Alomacrotona ruandensis sp. n. (Figg. 210-213)
Holotypus ©, Rwanda, Kayove, 2100 m, 15.V.1973, (Werner leg., MG).
Paratypi: 1 ©, Burundi, Kigwena Res., 11.11.1992, (Arndt leg., Mus. Erfurt); 1 ©, Kenya,
env. Endebess, pr. Kitale, 14.X1.1974, (Mahnert & Perret leg.).
Descrizione. Lungh. 3,4 mm. Corpo lucido e giallo-rossiccio sporco; antenne
bruno-rossicce con 1 tre antennomeri basali e 1°11° rossicci; zampe gialle. La retico-
lazione del capo e delle elitre è netta, quella del pronoto è svanita e quella degli
uroterghi è assente. La punteggiatura del capo è distinta e quella delle elitre è svanita.
Tubercoletti salienti coprono la superficie del pronoto.
Comparazioni. Si veda sotto la chiave delle specie.
Alomacrotona curticornis sp. n. (Figg. 214-215)
Holotypus ©, Rwanda, Kayove, 2100 m, 15.V.1973, (Werner leg., MG).
Descrizione. Lungh. 2,2 mm. Corpo lucido e giallo rossiccio con capo ed elitre
di un giallo-rossiccio sporco; antenne rossicce; zampe gialle. Il capo è privo di
reticolazione. La reticolazione del pronoto e delle elitre è molto svanita. Gli uroterghi
liberi 1° a 4° presentano una scultura squamiforma netta. La punteggiatura del capo è
svanita. Tubercoletti distinti stanno sul pronoto e sulle elitre. Spermateca fig. 215.
Comparazioni. Si veda sotto la chiave delle specie.
Alomacrotona kenyensis sp. n. (Figg. 216-217)
Holotypus 9, Kenya, Miss. Kaimosi, NE Kisumu, 1650 m, 10.X1.1974, (Mahnert &
Perret leg., MG).
ALEOCHARINAE DELL’ AFRICA ORIENTALE 833
204 205
01 mm
0, mm
Ficc. 204-209
Edeago in visione laterale e ventrale, habitus e spermateca. 204-205: Atheta (Oxypodera)
mahnerti sp. n.; 206-209: Atheta (Oxypodera) atopotheca sp. n.
834 ROBERTO PACE
CAN
215
Fico. 210-215
Habitus e spermateca. 210-211: Alomacrotona ruandensis sp. n. del Rwanda; 212-213:
Alomacrotona ruandensis sp. n., esemplare del Kenya; 214-215: Alomacrotona curticornis sp. n.
ALEOCHARINAE DELL’AFRICA ORIENTALE 835
Descrizione. Lungh. 3,2 mm. Corpo lucido e bruno con estremità addominale
rossiccia; antenne brune con antennomero basale giallo rossiccio e secondo antenno-
mero basale rossiccio; zampe giallo-rossicce. La reticolazione del capo è svanita, quella
del pronoto e dell’addome è assente e quella delle elitre è distinta. La punteggiatura del
capo è superficiale. I tubercoletti del pronoto e delle elitre sono salienti. Spermateca fig.
DAE
Comparazioni. Si veda sotto la chiave delle specie.
Alomacrotona tambachensis sp. n. (Figg. 218-219)
Holotypus 2, Kenya, Tambach, E Eldoret, 2000 m, 17.X1.1974, (Mahnert & Perret leg.,
MG).
Descrizione. Lungh. 3,8 mm. Corpo lucido e bruno-rossiccio con addome bruno;
antenne brune con antennomero basale giallo e 2° antennomero rossiccio; zampe giallo-
rossicce. Il capo e il pronoto sono privi di reticolazione. La reticolazione delle elitre è
svanita. La punteggiatura del capo è distinta. I tubercoletti che coprono il pronoto sono
salienti, quelli delle elitre sono superficiali. Spermateca fig. 219.
Alomacrotona remota Pace, 1986 (Figg. 220-221)
Alomacrotona remota PACE, 1986: 103.
29 2, Kenya, Tana River, Lac Shakababo, près de Ngao, 28.X.1977, (Mahnert & Perret
leg.); 1 2 Kenya, Galole Hola, 60 m, 21.X.1977, (Mahnert & Perret leg.).
Specie finora nota solo di Shibati, Africa orientale. Si rinviene anche tra foglie
morte di papiro.
CHIAVE DELLE SPECIE DEL GENERE Alomacrotona PACE, 1986
1 Antenne molto lunghe: il 10° antennomero è più lungo che largo;
addome fortemente ristretto all’indietro. Lungh. 3,9 mm. Madagaskar.
5555 ODE ine EA IR ONE termitophila (Bernhauer, 1901)
= Antenne moderatamente lunghe o corte; il 10° antennomero debolmente
o fortemente trasverso; addome debolmente ristretto all’indietro. Africa
(OO) CRE ae LIL ai: a, 2
2 Terzo antennomero fortemente o debolmente trasverso e 10° antenno-
IMCLOMOMEMEMEMTASVEISO RINO RE 3
= Terzo antennomero più lungo che largo e 10° antennomero più o meno
debolmenterttasversorse.e ALTER O OA Se 5
3 Bulbo prossimale della spermateca indistinto perché largo quanto la parte
mediana della stessa spermateca. Lungh. 3,1 mm. Nairobi.
RIINA invicta (Fauvel, 1907), comb. n. (olim Atheta invicta FAUVEL, 1907: 54)
= Bulbo prossimale della spermateca distinto, più largo della parte mediana
della;stessa:spermateca:. 1; 0 lA a Re ee RESO 4
- Addome coperto di microscultura squamiforme; spermateca grande quasi
il doppio di quella di remota; bulbo distale della spermateca privo di
introflessione apicale (fig. 215). Lungh. 2,2 mm. Rwanda. . ... curticornis sp. n.
836 ROBERTO PACE
- Addome coperto di tubercoletti salienti, così da dare un aspetto rugoso
alla superficie; spermateca quasi della metà rispetto quella di curticornis;
bulbo distale della spermateca con introflessione apicale (fig. 221).
Lungh. 2,1-2,3 mm. Kenya, Tanzania, Namibia. .......... remota Pace, 1986
5 10° antennomero debolmente trasverso; introflessione apicale del bulbo
distale della spermateca, robustissima, cioè a parete spessa. ............... 6
— 10° antennomero appena trasverso; introflessione apicale del bulbo
distalevdella:spermateca;esiles is 2... 2.2. 22 aoe 2 7
6 Pronoto meno trasverso; spermateca esile, con introflessione apicale del
bulbo disale, subconica. Lungh. 3,0 mm. Tanzania.
CE ER oe i I CI grandicollis (Bernhauer, 1915)
— Pronoto più trasverso; spermateca robustissima e tozza con introflessione
apicale del bulbo distale emisferica (fig. 219). Lungh. 3,8 mm. Kenya.
CE IR Rls ics tambachensis sp. n.
U Occhi molto sviluppati, più lunghi delle tempie; parte prossimale della
spermateca più corta della distale (fig. 217). Lungh. 3,2 mm. Kenya.
ae es e Rene COBE o O e er kenyensis sp. n.
- Occhi ridotti, più brevi delle tempie; parte prossimale della spermateca
più lunga della parte distale. Lungh. 3,4 mm............... ruandensis sp. n.
Brachysipalia imbellis Pace, 1985
Brachysipalia imbellis PACE, 1985: 130.
1 ©, Kenya, Embu, près de Kogari, 800 m, 6.X1.1974, (Mahnert & Perret leg.).
Specie finora nota solo del M. Aberdare.
Brachysipalia crassa Pace, 1985
Brachysipalia crassa PACE, 1985: 125.
5 es., Kenya, Mt. Kenya, 3050 m, Met St. Lodge, 23.X.1974, (Mahnert & Perret leg.).
Specie finora nota solo del M. Aberdare.
Brachysipalia aberdarensis Pace, 1985
Brachysipalia aberdarensis PACE, 1985: 127.
1 d, Kenya, Mt. Aberdares, Parc National, 25.X1.1975, (Mahnert & Perret leg.); 8 es.
Kenya, Mt. Kenya, 3200 m, Met St. Lodge, 23.X.1974, (Mahnert & Perret leg.).
Specie finora nota solo del M. Aberdare.
Brachysipalia meruensis Pace, 1985
Brachysipalia meruensis PACE, 1985: 130.
42 es., Kenya, Mt. Kenya, 3050 m, Met St. Lodge, 23.X.1974, (Mahnert & Perret leg.).
Specie finora nota solo del M. Meru.
Brachysipalia irangensis sp. n. (Figg. 222-225)
Holotypus d, Kenya, Embu, Irangi Forest St., 2000 m, 11.X.1977, (Mahnert & Perret
leg., MG).
Paratypi: 9 es., stessa provenienza.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 837
219
FIGG. 216-221
Habitus e spermateca. 216-217: Alomacrotona kenyensis sp. n.; 218-219: Alomacrotona tam-
bachensis sp. n.; 220-221: Alomacrotona remota Pace.
838 ROBERTO PACE
Descrizione. Lungh, 3,4 mm. Capo e pronoto opachi, resto del corpo lucido.
Corpo rossiccio; antenne giallo-rossicce con i due antennomeri basali gialli; zampe
giallo-rossicce. La reticolazione del capo e del pronoto è vigorosa, quella delle elitre e
dell'addome è trasversa e distinta. La punteggiatura del capo e del pronoto è assai
svanita. Tubercoletti salienti stanno sulle elitre. Edeago figg. 223-224, spermateca fig.
DIS
Comparazioni. In base alla forma della spermateca, la nuova specie sembra
sistematicamente vicina a B. meruensis Pace, 1985, del M. Meru, ma la spermateca
presenta una debole introflessione apicale del bulbo distale (profondissima in
meruensis) e il bulbo distale della stessa spermateca, nettamente meno trasverso.
L’edeago della nuova specie è molto più sviluppato, meno profondamente ricurvo al
lato ventrale e molto meno appuntito all’apice, in visione ventrale.
Brachysipalia elgonensis sp. n. (Figg. 226-229)
Holotypus d, Kenya, Mt. Elgon, 2700 m, Nat. Park, 15.X1.1974 (Mahnert & Perret leg.,
MG).
Paratypi: 20 es., stessa provenienza, ma anche 14.X1.1974.
Descrizione. Lungh. 2,7 mm. Capo e pronoto debolmente opachi, resto del corpo
lucido. Corpo giallo-rossiccio con uriti liberi 3° e 4° bruno-rossicci; antenne e zampe
interamente rossicce. Il capo e il pronoto presentano reticolazione vigorosa e pun-
teggiatura svanita. La reticolazione delle elitre è distinta, quella dell'addome è trasversa
e netta. Le elitre sono coperte di tubercoletti distinti. Edeago figg. 227-228, spermateca
Ne 220)
Comparazioni. Specie simile a B. kenyamontis Pace, 1985, se si compara la
forma dell'edeago e della spermateca. La nuova specie differisce da kenyamontis perché
l'edeago è più esile in visione ventrale, meno profondamente ricurvo al lato ventrale e
per la presenza di un robustissimo pezzo copulatore del sacco interno tronco all'apice
(terminante in un'appendice filiforme in kenyamontis). L'introflessione apicale del
bulbo distale della spermateca è chiaramente più profonda nella nuova specie.
Brachysipalia elgonicola sp. n. (Figg. 230-233)
Holotypus d, Kenya, Mt. Elgon, 2700 m, 15.XI. 1974 (Mahnert & Perret leg., MG).
Paratypi: 27 es., stessa provenienza.
Descrizione. Lungh. 2,2 mm. Corpo lucido e giallo-rossiccio; antenne giallo-
brune con i due antennomeri basali gialli; zampe gialle. La reticolazione del capo e del
pronoto è molto svanita, quella delle elitre e dell'addome netta. La punteggiatura del
capo e del pronoto è molto superficiale. Tubercoletti svaniti coprono le elitre. Edeago
figg. 231-232, spermateca fig. 233.
Comparazioni. L'edeago e la spermateca della nuova specie hanno forma cosi
differente rispetto quella delle altre specie del genere, che non è possibile avvicinare
sistematicamente la nuova specie ad altre simili.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 839
FIGG. 222-229
Habitus, edeago in visione laterale e ventrale e spermateca. 222-225: Brachysipalia irangensis
sp. n.; 226-229: Brachysipalia elgonensis sp. n.
840 ROBERTO PACE
232
295
01 mm
Fico. 230-235
Habitus, edeago in visione laterale e ventrale e spermateca. 230-233: Brachysipalia elgonicola
sp. n.; 234-235: Brachysipalia ruwenzorensis Sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 841
Brachysipalia ruwenzorensis sp. n. (Figg. 234-235)
Holotypus 9, Ruwenzori, Kitandara Hut, 13200 ft, I. 1984 (C. Lengeler leg., MG).
Descrizione. Lungh. 3,4 mm. Corpo lucido e rossiccio scuro con uriti liberi 2°,
3°, 4° e metà basale del 5° bruni; antenne rossicce con i tre antennomeri basali giallo-
rossicci; zampe rossicce. La reticolazione del pronoto è vigorosa, quella sul resto del
corpo è netta. Il capo è coperto di tubercoletti distinti. Il pronoto presenta tubercoletti
salienti, ma sono assenti sulla linea mediana. Le elitre hanno tubercoletti poco salienti.
spermateca fig. 235.
Comparazioni. La forma della spermateca indica anche per questa specie come
la precedente, distanza sistematica rispetto le altre specie del genere, per cui non è
comparabile con nessuna.
Brachysipalia sanfilippoi sp. n. (Figg. 236-239)
Holotypus d, M. te Kenya, Met. Station, 3050 m, 22.IX.1976 (N. Sanfilippo leg., Mus.
Genova).
Paratypi: 9 es., stessa provenienza.
Descrizione. Lungh. 2,1 mm. Corpo lucido e giallo, comprese antenne e zampe.
La reticolazione del corpo è netta sul disco e svanita al lati e nella zona occipitale. Sul
resto del corpo la reticolazione è netta. La punteggiatura ombelicata del capo è distinta.
Il pronoto è coperto di punteggiatura finissima: vi sono 4 punti più sviluppati. Le elitre
sono coperte di tubercoletti nettamente salienti.
Comparazioni. La nuova specie presenta la spermateca con bulbo prossimale
indistinto, mentre in tutte le restanti specie note è da distinto a molto sviluppato.
L'edeago mostra reminiscenze con quello di B. kenyamontis Pace, 1985, ma è netta-
mente meno svilupatto e proporzionalmente più largo in visione ventrale.
Etimologia. Specie dedicata al suo raccoglitore: il noto entomologo genovese
Nino Sanfilippo, studioso di Hydrophilidae e per tanti anni segretario della Società
Entomologica Italiana.
Brachysipalia viatica sp. n. (Figg. 240-243)
Holotypus d, Kenya, Mt. Elgon, 2800 m, 14.X1.1974 (Mahnert & Perret leg., MG).
Paratypi: 5 es., stessa provenienza.
Descrizione. Lungh. 3,4 mm. Corpo lucido e giallo-rossiccio con uriti liberi 2°,
3°, 4° e 5° bruni; antenne giallo-rossicce con i tre antennomeri basali e l'11° gialli;
zampe giallo-rossicce. La reticolazione del capo è distinta, quella del pronoto è svanita
e quella delle elitre e dell'addome è netta. La punteggiatura del capo é superficiale,
quella del pronoto è indistinta. Tubercoletti nettamente salienti stanno sulle elitre.
Edeago figg. 241-242, spermateca fig. 243.
Comparazioni. Specie simile a B. crassa Pace, 1985, del M. Kenya e a B. kenya-
montis Pace, 1985. Se ne distingue da entrambe per avere l'edeago ampiamente ricurvo
al lato ventrale, con "crista apicalis" e "crista proximalis" nettamente più svilupatte. La
spermateca ha bulbo prossimale non così ipertrofico come quello delle due specie citate
sopra.
842 ROBERTO PACE
Olmm
243
Ficc. 236-243
Habitus, edeago in visione laterale e ventrale e spermateca. 236-239: Brachysipalia sanfilippoi
sp. n.; 240-243: Brachysipalia viatica sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 843
Brachysipalia embuensis sp. n. (Figg. 244-249)
Holotypus 4, Kenya, Embu, Irangi Forest, 2100 m, 12.X.1977 (Mahnert & Perret leg.,
MG).
Paratypus: 1 2, stessa provenienza.
Descrizione. Lungh. 2,2 mm. Corpo lucido e rossiccio con 4° urite libero bruno;
antenne brune con i due antennomeri basali e l'11° bruno-rossicci; zampe gialle. La
reticolazione del capo è assente, quella del pronoto è distinta. I tubercoletti delle elitre
sono molto salienti verso la sutura. Addome non reticolato. Edeago figg. 245-246,
spermateca fig. 247.
Comparazioni. La nuova specie ha edeago simile a quello di B. franzi Pace,
1985, del Kilimangiaro, ma ha dimensione ridotta e pezzi copulatori del sacco interno
di forma diversa: pezzo a uncino assente in franzi. Inoltre la spermateca è meno
sviluppata nella nuova specie, sebbene di forma simile a quella di franzi, ha però
l'introflessione apicale del bulbo distale, stretta, mentre in franzi è larghissima.
Brachysipalia volans sp. n. (Figg. 248-249)
Holotypus 9, Kenya, Mt. Elgon, 2700 m, 15.X1.1974 (Mahnert & Perret leg., MG).
Paratypi: 2 © ®, stessa provenienza, ma 2650 m e 14.X1.1974.
Descrizione. Lungh. 2,2 mm. Corpo lucido e rossiccio scuro; antenne bruno-
rossicce con antennomero basale rossiccio; zampe giallo-rossicce. Il capo, il pronoto e
l'addome sono privi di reticolazione. La reticolazione delle elitre è svanita. La pun-
teggiatura del capo e del pronoto è molto superficiale. Netti tubercoletti stanno sulle
elitre.
Comparazioni. La spermateca della nuova specie ha forma e grandezza simili
alla spermateca di B. franzi Pace, 1985, del Kilimangiaro, ma l'introflessione apicale del
bulbo distale della spermateca stessa della nuova specie, è stretta e profonda, mentre
quella di franzi è larghissima e meno profonda. Inoltre le elitre della nuova specie sono
appena più corte del pronoto, mentre in franzi sono molto più corte del pronoto. Il
pronoto della nuova specie presenta 4 punti discali, assenti in franzi.
844 ROBERTO PACE
FicG. 244-249
Habitus, edeago in visione laterale e ventrale e spermateca. 244-247: Brachysipalia embuensis
sp. n.; 248-249: Brachysipalia volans sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 845
RINGRAZIAMENTI
Ringrazio molto cordialmente il Dr I. Löbl e il Dr C. Besuchet del Museo di
Storia Naturale di Ginevra per avermi affidato in studio il materiale pubblicato nel
presente lavoro. Ringrazio per lo stesso motivo il Dr R. Poggi e il Sig. N. Sanfilippo del
Museo di Storia Naturale di Genova e il Dr L. Zerche del D.E.I. di Eberswalde. Per il
prestito di tipi rigrazio vivamente il Dr L. Baert dell'Institut Royal des Sciences
Naturelles de Belgique di Bruxelles, il Dr A.F. Newton del "Field Museum of Natural
History" di Chicago e il Dr L. Bartolozzi del Museo Zoologico dell'Università di
Firenze.
BIBLIOGRAFIA
BERNHAUER, M. 1901. Ein neuer Termitengast aus Madagaskar. Deutsche ent. Z. 1901: 252.
BERNHAUER, M. 1915. Zur Staphylinidenfauna des tropischen Afrika. Annls. hist.-nat. Mus. natn.
hung. 13: 95-189.
BERNHAUER, M. 1927. Materiali per lo studio della fauna Eritrea raccolti nel 1901-1903 dal Dr A.
Andreini, Staphylinidae. Boll. Soc. ent. Ital. 59: 79-81.
BERNHAUER, M. 1931. Entomological Expedition to Abyssinia, 1926-1927: Coleoptera, Staphy-
linidae. Systematic report by Dr Max Bernhauer. J. Linn. Soc. Lond. Zool. 37: 559-605.
BERNHAUER, M. 1934. Neue Kurzfliiger von Ruwenzori-Zivu-Gebiet. XXXIV. Beitrag zur
afrikanischen Staphylinidenfauna. Rev. Zool. Bot. afr. 25: 206-217.
BERNHAUER, M. 1944. Phleophile Staphyliniden aus Ostafrika (Coleoptera). Rev. franc. Ent. 11:
17-21.
BERNHAUER, M. & SCHEERPELTZ, O. 1926. Coleopterorum Catalogus. Staphylinidae 4: 499-988,
Berlin.
CAMERON, M. 1932. New species of Staphylinidae from the Belgian Congo. Bull. Ann. Soc. Ent.
Belg. 72: 131-146.
CAMERON, M. 1939. The fauna of British India. Coleoptera Staphylinidae 4: 691 pp., London.
CAMERON, M. 1942. New species of Staphylinidae (Col.) collected by the Coryndon Museum
Expedition to the Chyulu Hills. Ann. Mag. Nat. Hist. 9: 321-332.
CAMERON, M. 1950. Staphylinidae (Coleoptera Polyphaga). Explor. Parc natn. Albert Miss. G. F.
de Witte 59: 1-85.
EICHELBAUM, F. 1913. Verzeichnis der von mir in den Jahren 1903 und 1904 in Deutsch- und
British-Ostafrika eingesammelten Staphylinidae. Arch. Naturg. 79: 114-168.
FAUVEL, A. 1900. Staphylinidae nouveaux de Kinchassa (Congo). Rev. Ent. 19: 66-74.
FAUVEL, A. 1907. Voyage de M. Ch. Alluaud dans l'Afrique Orientale. Rev. Ent. 26: 10-70.
FENYES, A. 1920. Coleoptera: Fam. Staphylinidae, subfam. Aleocharinae. Genera Insectorum
173: 111414.
JEANNEL, R. & PAULIAN, R. 1945. Mission scientifique de l'Omo. Faune des terriers des rats-
taupes, IV: Coléoptères. Mem. Mus. Nat. Hist. 19: 51-147.
KRAATZ, G. 1857. Beiträge zur Kenntniss der Termitophilen. Linnaea Ent. 11: 44-56.
MULSANT, M.E. & REY, C. 1873. Description de divers coléoptéres brévipennes nouveaux ou peu
connus. Opusc. Ent. 15: 147-189.
PACE, R. 1984. Aleocharinae delle Mascarene. Parte II. Rev. suisse Zool. 91: 249-280.
PACE, R. 1985. Aleocharinae raccolte dal Prof. Franz sul Kenya, Kilimangiaro e Monti Aberdare.
Fragm. Ent. 18: 115-159.
846 ROBERTO PACE
PACE, R. 1986. Aleocharinae dell'Africa Orientale (Coleoptera, Staphylinidae). Ann. hist.-nat.
Mus. Natn. hung. 78: 83-143.
PACE, R. 1994. Aleocharinae della Sottoregione Africana Orientale al Museo di Ginevra (Cole-
optera, Staphylinidae) Parte I. Rev. suisse Zool. 100: 117-193.
SCHEERPELTZ, O. 1968. Coleoptera Staphylinidae. Cat. faun. Austriae 15. Wien Akad. Wiss.:
1-124.
THOMSON, C.G. 1858. Försök till uppställning af Sveriges Staphyliner. Öfv. Kön. Vet.-Akad.
Förh. 15: 27-40.
TOTTENHAM, C.E. 1957. Coleoptera Staphylinidae: Tachyporinae, Pygosteninae (cont.) and Aleo-
charinae (part.). Ann. Mus. Congo Tervuren 58: 73-135.
REVUE SUISSE DE ZOOLOGIE, 102 (3) : 847-850; septembre 1995
Chaerilus sabinae, nouvelle espèce de Scorpion anophthalme
des grottes de Matampa en Inde (Scorpiones, Chaerilidae)
Wilson R. LOURENCO
Société de Biogéographie, 57, rue Cuvier, F-75005 Paris, France.
Chaerilus sabinae, a new species of anophthalmous scorpion from the
caves of Matampa in India (Scorpiones Chaerilidae).- A new species of
anophthalmous, troglobite scorpion, Chaerilus sabinae is described from
the Matampa caves, in the region of Gua Atas in India. This species is the
second genuine troglobite scorpion known for the genus Chaerilus and the
number of known troglobite scorpions in now raised to 14.
INTRODUCTION
Le genre Chaerilus Simon, 1877, appartient a la famille monotypique des
Chaerilidae Pocock, 1900. Le nombre total des espèces n'est pas exactement connu à
l'heure actuelle car plusieurs d'entre elles ont été pauvrement caractérisées et leurs
statuts d'espèce voire de sous-espèce demeurent obscurs. Un nombre approximatif
peut étre néanmoins estimé entre 15 et 23. La répartition connue de ce genre recouvre
l'Inde, le Sri Lanka, l'Asie du sud-ouest, les Philippines, Bornéo, les Célèbes, Java,
Sumatra et plusieurs autres îles indonésiennes (SISSOM, 1990).
Les descriptions des espèces appartenant au genre Chaerilus sont anciennes et
datent des années entre 1877 et 1913, exceptée celle de Chaerilus chapmani, par
VACHON & LOURENÇO (1985), à partir des spécimens collectés dans les grottes du
Parc National Gunong Mulu au Sarawak (Bornéo).
Chaerilus sabinae n. sp. (Figs 1 a9)
Holotype: d immature, provenant des grottes de Matampa dans la région de Gua Atas
(22°12' N/85°23' E), Inde, leg. Mards, 12/VIII/1985 (Muséum d'histoire naturelle, Genève).
Etymologie: Le nom spécifique est attribué en hommage au Dr Sabine Jourdan,
Paris, France.
Description. Coloration. La couleur de base de l'ensemble du corps, pattes et
pédipalpes est jaunâtre; aucune tache n'est présente. Vésicule jaune-clair; aiguillon
légèrement rougeatre. Sternites jaunatres. Peignes et opercule génital jaune très clair,
Manuscrit accepté le 30.01.1995.
848 WILSON R. LOURENCO
Figs 1 et 2
Chaerilus sabinae, holotype d, vues dorsale et ventrale.
presque blanc; sternum, hanches et processus maxillaires jaunatres. Pattes, pédipalpes
et chélicères jaunatres.
Morphologie. Prosoma: plaque prosomienne a bord frontal légèrement échancré,
divisé par un sillon longitudinal très discret; région antérieure aplatie et lisse; deux
sillons présents dans les régions latéro-postérieures; tubercule oculaire, yeux médians
et yeux latéraux absents. Mesosoma, dorsalement chagriné; tergites sans carènes
apparentes; face ventrale: sternum pentagonal nettement plus haut que large; opercule
génital formé de deux plaques plus ou moins ovales; peignes avec 4-4 dents; fulcres
bien distincts; stigmates petits et arrondis. Metasoma: anneaux I à V avec des carènes
très discrètes, pratiquement absentes. Vésicule aplatie et allongée, chagrinée; ai-
guillon long, à peine courbé. Chélicères avec la dentition caractéristique de la famille.
Pédipalpes avec des carènes très faiblement marquées; tranchant des doigts des pinces
avec quelques séries de granules (4 ou 5) suivies de séries peu distinctes à la base des
doigts: présence de quelques granules accessoires. Trichobothriotaxie du type B
(VACHON, 1973), numériquement constante dans toute la famille des Chaerilidae.
Tarses pourvus de soies disposées en deux séries latérales avec, ventralement, une
série de petites épines.
NOUVELLE ESPECE DE SCORPION ANOPHTHALME 849
Fics 3 À 9
Chaerilus sabinae, holotype d . 3. Plaque prosomienne. 4. Sternum, opercule génital et peignes.
5. Cinquième anneau du metasoma et telson. 6. Pince, vue externe. 7. Pince, vue ventrale. 8.
Tibia, vue dorsale. 9. Fémur, vue dorsale.
Mensurations (en mm) de l'exemplaire décrit. Plaque prosomienne longueur, 1,1.
Mesosoma longueur, 2,4. Metasoma + telson longueur, 3,9. Fémur longueur, 0,9.
Tibia longueur, 1,2. Pince longueur, 2,3. Doigt mobile longueur, 1,3.
Ainsi que Chaerilus chapmani, Ch. sabinae fait partie du groupe des Chaerilus
ayant les pinces des pédipalpes longues et effilées, ayant des doigts au moins deux
fois la largeur de la pince. Cette espèce peut être distinguée des autres espèces du
genre Chaerilus par l'absence totale des yeux médians et latéraux. Chez Ch. chapmani
850 WILSON R. LOURENCO
subsiste un ceil médian réduit et un ceil latéral de chaque còté. Ces deux espèces sont
de toute évidence des troglobies véritables au sein des Chaerilidae. (VACHON &
LOURENGO, 1985). Le nombre de scorpions troglobies s'élève ainsi à 14 (LOURENÇO &
FRANCKE, 1985).
REFERENCES
LOURENÇO W.R. & O.F. FRANCKE. 1985. Révision des connaissances sur les Scorpions
cavernicoles (troglobies) (Arachnida, Scorpiones). Mém. Biospéol., 12: 3-7.
SISSOM, W.D. 1990. Systematics, Biogeography, and Paleontology. In: The Biology of
Scorpions. Polis, G.A. (ed.). Stanford Univ. Press, Stanford, pp. 64-160.
VACHON, M. 1973. Etude des caractères utilisés pour classer les familles et les genres de Scor-
pions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles trichobothriaux et
types de trichobothriotaxie chez les Scorpions. Bull. Mus. natn. Hist. Nat., Paris, 3e
sér., n° 140, Zool., 104: 857-958.
VACHON, M. & W.R. LOURENÇO. 1985. Scorpions cavernicoles du Sarawak (Bornéo). Chaerilus
chapmani n. sp. (Chaerilidae) et Lychas hosei (Pocock, 1890) (Buthidae). Mém. Bio-
spéol., 12: 9-18.
REVUE SUISSE DE ZOOLOGIE
Tome 102 — Fascicule 3
Papp, Lazlo. Oriental species of the genus Biroina Richards (Diptera:
Sphaetocenidae)y sy Werte Kr ao an E doti.
ROWELL, C. Hugh F. New and little known taxa of Rhytidochrotinae (Acri-
didacsS©rthoptera);Mrom{CostARICA ZERI
Zıcsı, Andras. Regenwürmer aus Bolivien (Oligochaeta)..............
CALDARA, Roberto. Studio dei tipi di Tychiini di Henri Tournier custoditi
presso il Museo di Storia Naturale di Ginevra (Coleoptera, Curcu-
ior hae) rere ee pee N rer OR IRE ee
Bassi, Graziano. Contributi allo studio delle Crambinae (Lepidoptera:
Crambidae). IX. Note sui generi Sebrus Bleszynski e Alphacrambus
NOVICONICESCHZIONE(dIMmUOVE Specie jee | ans ae see
Gomy, Yves. Un Anapleus Horn nouveau de |’Himalaya (Coleoptera,
Biisten do). Re anta Gir. Me UAE eee a
BEN SLIMANE, Badreddine & Marie-Claude DURETTE-DESSET. Oswaldo-
cruzia (Nematoda, Trichostrongylina, Molineoidea) parasites d’Am-
phibiens du Brésil et de l’Equateur, avec redéfinition de l’espèce-type
O. subauricularis (Rudolphi, 1819) et d’O. mazzai Travassos, 1935
Cuccoporo, Giulio & Ivan LOBL. Revision of the Afrotropical rove-beetles
of the genus Megarthrus (Coleoptera, Staphylinidae, Proteininae). . . .
SCHEUERN, Joachim. Eine neue Haroldius-Art aus Borneo (Coleoptera,
SCaraWacidac) se arts Meee ee ET See sae rr Or RI TARE
PETTER, Annie J. Dichelyne moraveci n. sp., parasite de Pseudoplastystoma
fasciatum et notes sur les Cucullanidae du Paraguay..............
PACE, Roberto. Aleocharinae della Sottoregione Africana Orientale al
Museo di Ginevra (Coleoptera, Staphylinidae). Parte II. ............
LOURENÇO, Wilson R. Chaerilus sabinae, nouvelle espèce de Scorpion
anophthalme des grottes de Matampa en Inde (Scorpiones, Chaeri-
NAS) Feet eee ARR E SFONDO OSE CARE Si SRE rio i
Pages
539-552
553-583
585-608
609-613
615-627
629-634
635-653
655-761
763-768
769-778
779-846
847-850
REVUE SUISSE DE ZOOLOGIE
Volume 102 — Number 3
Papp, Läzlö. Oriental species of the genus Biroina Richards (Diptera:
Sphaeroeenidae): fonti. aa ati SIL RODE
ROWELL, C. Hugh F. New and little known taxa of Rhytidochrotinae
(Acnididace@rthoptera) from CostaRicass sass ee eee
Zıcsı, Andras. Earthworms from Bolivia (Oligochaeta)................
CALDARA, Roberto. Study of the Tychiini types of Henri Tournier preser-
Bassi, Graziano. Contribution to the study of the Crambinae (Lepidoptera,
Crambidae). IX. Notes on genera Sebrus Blezynski and A/phacram-
bus:noys wath description. of newispeciesr. ii IT
Gomy, Yves. A new species of Anapleus Horn from the Himalaya
(GColeoptera-=Misterida o). al eo ORAL
BEN SLIMANE, Badreddine & Marie-Claude DURETTE-DESSET. Oswaldo-
cruzia (Nematoda, Trichostrongylina, Molineoidea) parasitic in
Brazilian and Ecuadorian Amphibians, with redefinition of the type
species O. subauricularis (Rudolphi, 1819) and O. mazzai Travassos,
OB SERIAL. PN N al
Cuccoporo, Giulio & Ivan LOBL. Revision of the Afrotropical rove-
beetles of the genus Megarthrus (Coleoptera, Staphylinidae, Pro-
(emma) iena ZO
SCHEUERN, Joachim. A new Haroldius species from Borneo (Coleoptera,
Scarabaeidae) es tt Sir li ee
PETTER, Annie J. Dichelyne moraveci sp. n. parasite of Pseudoplatystoma
fasciatum, with remarks on Cucullanids from Paraguay............
PACE, Roberto. Aleocharinae from the Eastern African Subregion in the
Geneva Museum (Coleoptera, Staphylinidae). Part II. ............
LOURENÇO, Wilson R. Chaerilus sabinae, a new species of anophthalmous
scorpion from the caves of Matampa in India (Scorpiones Chaeri-
IEEE ROTTI EIA
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TOME 102 — FASCICULE 4
Publication subventionnée par l'Académie suisse des Sciences naturelles
et la Société suisse de Zoologie
VOLKER MAHNERT
Directeur du Muséum d'histoire naturelle de Genève
FRANCOIS BAUD
Conservateur au Muséum d'histoire naturelle de Genève
DANIEL BURCKHARDT
Chargé de recherche au Muséum d'histoire naturelle de Genève
Comité de lecture
Le président de la Société Suisse de Zoologie
Le directeur du Muséum de Genève: Volker MAHNERT — Systématique des
vertébrés — Muséum de Genève
Le président du comité: Ivan LOBL — Systématique des Insectes — Muséum de
Genève
Patrick GUERIN — Physiologie et éthologie des arthropodes — Institut de Zoologie,
Neuchatel
Willy MATTHEY — Ecologie, entomologie — Institut de Zoologie, Neuchatel
Claude MERMOD — Ethologie et écologie des vertébrés — Université de Neuchatel
Paul SCHMID-HEMPEL — Ecoéthologie, biologie des populations — Institut f.
Terrestrische Okologie, ETH Ziirich, Schlieren
Steve STEARNS — Biologie de l'évolution — Institut f. Zoologie, Basel
Beat TSCHANZ — Ethologie des Vertébrés — Zoologisches Institut, Bern
Claude VAUCHER — Systématique des Invertébrés — Muséum de Genève
La préférence sera donnée aux travaux concernant les domaines suivants: Biogéographie,
systématique, écologie, éthologie, morphologie, et anatomie comparée, physiologie.
Administration
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REVUE SUISSE DE ZOOLOGIE, 102 (4) : 853-867; décembre 1995
ZOOLOGIA 95
Zürich, 15-17 March 1995
(Annual Conference of the Swiss Zoological Society)
Interactions between genetics and ecology
MAIN CONFERENCES
A. Meyer (Dept. of Ecology and Evolution, State University of New York,
Stony Brook, NY 11794-5245, USA): Examples of the use of DNA sequences in
population biology and phylogenetics.
Morphology and molecules both should provide congruent information about the
evolutionary history of organisms. In terms of ease of analyses, and data collection, molecular
data have some advantages for phylogeny reconstruction over morphological data. Some of
those advantages but also potential pitfalls of molecular data for the purpose molecular syste-
matics are discussed. Our understanding of the phylogenetic relationships among vertebrates
and the evolution of DNA, particularly mitochondrial DNA, has increased rapidly since the
recent invention of the polymerase chain reaction (PCR). This cloning technique together with
direct sequencing of PCR-amplified DNA has simplified and dramatically accelerated the
accumulation of DNA sequence information for phylogenetic work. Methods of data collection
and data analysis for phylogenetic studies on vertebrates with particular emphasis on fish and
mitochondrial DNA are outlined. General aspects of the biology of mitochondrial DNA that
pertain to phylogenetic reconstruction are reviewed and advantages of DNA sequences over
alternative DNA-based genetic markers are highlighted. Examples of phylogenetic work based
on mitochondrial and nuclear DNA sequences are used to illustrate the methods, advantages
and potential problems with techniques, choice of genes and phylogenetic analyses. These
examples dealt with (1) The evolution of cichlid fish species flocks in the East African lakes,
(2) the systematics and evolutionary history of cetaceans based on DNA sequences, (3) the evo-
lution of a sexually selected trait, the sword, in swordtail fishes (genus Xiphophorus) as infer-
red from a DNA-based phylogeny of the genus.
In each of these examples, DNA phylogenies resulted in different phylogenetic hypo-
theses than had traditionally been believed in. Potential reasons for the differences in the DNA-
based and the traditional morphology-based phylogenies might have to do with morphological
convergence and non-independence of morphological traits that had been used to construct the
phylogenetic hypotheses. Those are potential phylogenetic pitfalls that are more easily dealt
with in molecular than in morphological data sets. In the first case study the DNA data
suggested, contrary to traditional assumptions, that the Lake Victoria and Lake Malawi cichlid
species flocks are both monophyletic and hence morphological similarities found in both have
independently evolved. In the phylogenetic study of the evolutionary history of whales it was
concluded that baleen whales are the sistergroup to one group of toothed whales, the sperm
whales. Based on the DNA-phylogeny we concluded that baleen whales are evolutionarily
younger and all whales had a different evolutionary history than previously assumed. The
sword of males in swordtail fishes, based on the DNA-phylogeny is likely to be as old as the
preference of females to mate with males with swords. During the evolution of the species in
this genus, the sword was probably lost and re-evolved twice independently. This interpretation
is in conflict with the hypothesis that the sword evolved after the females “pre-exisiting bias”.
854 ZOOLOGIA 95
W. Watt (Stanford). Allozymes in evolutionary biology: beyond the twin
pitfalls of “neutralism” and “selectionism”.
A. Pomiankowski (London). Continuous change in mate preferences.
H. Hofer (Seewiesen). Genes, behaviour and anatomy: female penises are an
adaption for... what?
ABSTRACTS
SHORT PAPERS
A. Aebischer (Fribourg): Territory choice, mate choice and breeding success
in the Savi's Warbler Locustella luscinioides.
We investigated how territory quality, settlement date and morphometry (size, weight,
condition) affected the several components of yearly breeding success (mating success, clutch
sizes and numbers, fledging success) in male and female Savi's Warblers (Locustella lusci-
nioides) from a Swiss population.
Territories occupied by males differed from unoccupied sites of similar size and
localisation by higher and denser reeds, an extensive straw litter and a thick cover of dead sedge
leaves. Availability of suitable nesting sites seemed an important choice criterion. Territories that
most closely matched this ideal were also the first chosen by males upon arrival from migration.
These males however did not differ from those that arrived later on the basis of morphometry.
Male breeding success decreased with the date of territory occupancy, due to a decrease
in mating success. Female breeding success decreased with the date of first-clutch laying,
mainly due to a decrease in clutch numbers. Breeding success did not correlate with morpho-
metry. Its positive correlation with territory quality was indirect, and mediated through their
common dependence on occupancy date (males) and first-clutch laying date (females).
Our results thus provide clear support for territory choice by males, but not for mate
choice, and show the crucial role played by settlement date on many important parameters of the
breeding cycle of both sexes. We propose a parsimonious model to account for our observations,
and discuss it with respect to the breeding biology of better-known warbler species.
P. Agnew (ETH Ziirich): Sex ratio distortion in the mosquito Aedes aegypti
caused by a microparasite.
B. Anholt (Zürich) & R. Trembath (Toronto): Predator induced phenotypic
plasticity in tadpoles of Hyla versicolor.
Organisms often display variation in phenotype in response to variation in their
environment. A classic example of this phenotypic plasticity is predator induced defenses.
Larvae of anurans that breed in temporary ponds experience highly variable predation regimes
and also display differences in phenotype that are correlated with predator presence. To test this
possibility we raised larvae of Hyla versicolor in the presence and absence of caged dragonfly
larvae. In the non-lethal presence of a predator, tadpoles developed brightly coloured as
opposed to clear tails and the tails were deeper with thicker tail musculature. Observations of
activity showed that animals were also less active in the presence of the predators. Measure-
ments of burst speed could not distinguish between animals raised with and without predators.
In predation trials, animals with the predator-induced phenotype had a higher probability of
survival than the non-induced phenotype. It is not yet clear what disadvantages are associated
with the predator-induced phenotype. Two possibilities are reduced competitive ability, and
susceptibility to other, perhaps avian, predators.
ZOOLOGIA 95 855
C. Bender (Johann Wolfgang Goethe-Universität, Zoologisches Institut, Sies-
mayerstr. 70, D-60054 Frankfurt a.M.): Demographic and genetic studies in isolated
populations of wall lizards.
The wall lizard, Podarcis muralis, inhabits locations which are considerably influenced
by human activities. During the past decades most populations became reduced or isolated in
Central Europe, where the species is considered endangered. In fact, population reductions
followed by habitat destruction may lead to loss of genetic variation, with possible negative
consequences on the demography and survival of these populations.
In the present study, wall lizard populations sizes were estimated. Several populations
of Podarcis muralis were examined for genetic variability, using PCR based RAPD finger-
printing. Based on data from one population of approximately 100 specimens, a computer
model has been developed, predicting an average survival time of ca. 36 years. Because dif-
ferences in population size may have consequences for genetic structure of the populations,
future management efforts should incorporate demographic, as well as genetic information.
L.F. Bersier, G. Sugihara & A. Hobday (UC San Diego): A correspondence
between two classical notions of community structure: species abundances and den-
drograms of niche similarities.
We examined a possible relationship between two different descriptions of community
structure. The first is the niche overlap dendrogram that describes the ecological similarities of
species and groups or guilds of species. The second is the pattern of the relative abundances of
those species. Specifically, we shall examine the particular way in which this relationship
follows from the niche hierarchy model. This model can be viewed either as sequential
invasions of species or, more generally, to represent hierarchical organization of the kind
represented in the dendrogram, we shall focus here on this latter aspect of community structure.
Viewing the model in this way, the organization of the community is reckoned to be self-
similar, with individual species partitioning resources in the same way that large sets of species
partition resources. Moreover the organization is hierarchical in the exact sense depicted by the
niche overlap dendrogram. The physical metaphor for this type of self-similar decomposition is
a sequential, binary, breakage process applied directly to abundances, or population biomasses.
Thus one can imagine the total biomass of species in a community that is sequentially divided
so that break points correspond to the branching structure in niche similarities. We tested two
predictions of this model using studies that provide both dendrograms and relative abundances.
First, we showed that the relative abundances at the break points differ significantly from what
one would predict if the actual abundances were allocated at random to the known dendrogram.
Second, we showed that species residing at terminal branches of the dendrogram that are highly
split (more highly subdivided lines) are on average less abundant than species from branches
that are less highly subdivided.
U. Bircher, H. Jungen, R. Burch & E. Hauschteck-Jungen (Ziirich):
Multiple morphs of sperm were required for the evolution of the Sex Ratio trait in
Drosophila.
In Sex Ratio males of D. subobscura Y sperm degenerate. Nevertheless, Sex Ratio
males can produce as many offspring as non-Sex Ratio males. In order to find an explanation
for the high number of offspring of Sex Ratio males we analysed spermatogenesis and sperm
storage in females. Because D. subobscura has two morphs of sperm, short and long ones, we
connected the sperm heteromorphism with the Sex Ratio trait in this species.
Spermatids of both lengths developed simultanously in the testes. The first mature
sperm produced by a young male were nearly all short sperm. This result indicates that short
cysts need less time to mature than long cysts. Thus in a given time more short than long cysts
856 ZOOLOGIA 95
can mature. In the testes of Sex Ratio males more short cysts on the cost of long cysts were
formed than in testes of non-Sex Ratio males. Thus in a unit time Sex Ratio males can produce
as many or more sperm than non-Sex Ratio males.
Females mated to Sex Ratio males have more sperm in their storage organs than those
which mated to non-Sex Ratio males. From the sperm transfered females stored selectively
more long than short sperm after mating to a Sex Ratio male than after mating to a non-Sex
Ratio male.
The effect of the high amount of sperm in Sex Ratio males is twofold: the loss of Y
sperm is compensated, and probably in nature a second copulation is prevented by the complete
filling of the female storage organs.
From these results we conlude that the arising and persistence of the Sex Ratio trait in
natural populations depends on the presence of sperm heteromorphism.
M. Chapuisat (Lausanne): Number of matings, genetic relatedness and popu-
lation viscosity in polygynous Formica ants.
Estimating genetic relatedness among nestmates is important to assess the role of kin
selection in the evolution of social behaviour. In social insects, intranest genetic relatedness is
shaped by the reproductive structure of the colony, particularly the number of reproducing
individuals, and by the local population structure.
I studied number of matings, intranest genetic relatedness and local population genetic
structure in Formica lugubris. This red wood ant has hundreds of queens per nest, and therefore
genetic relatedness is expected to decrease to virtually 0. I used microsatellites, which are
highly polymorphic DNA markers well suited for this kind of detailed studies.
Mother-offsping analyses and direct typing of the sperm stored in the spermatheca of
the queens revealed that about one fifth of the queens did mate multiply. This low level of
multiple mating, corresponding to an effective mate number of 1.15, will have little effect on
the kin structure of the colony.
Genetic relatedness among nesmate workers, measured over a sample of 20 nests along
a 1.2 km transect, was 0.17 + 0.04. This value is surprisingly high given the high number of
queens per nest, but a positive inbreeding coefficient (Fis = 0.07 + 0.03) indicates that local
population structure and/or inbreeding boost this relatedness estimate.
The microgeographic population structure is viscous, with a positive correlation
between geographical distance and genetical differentiation. This local population structure is
probably due to dispersal by nest budding and restricted gene flow between existing nests.
O. Daniel (ETH Ziirich): Earth worms and soil fertility.
P. Flook (Zoologisches Institut der Universitat Basel): Sequence Variation in
Orthopteran Mitochondrial Ribosomal RNA Genes.
Sections of the large and small subunit mitochondrial ribosomal genes have been
sequenced in over eighty species of Orthoptera. The taxonomic sample contains representatives
from nearly all of the caeliferan families and two thirds of the subfamilies of the large family
Acrididae, as well as representatives of Ensifera, Phasmida and Grylloblattodea. The wide
range of divergence dates between the different taxa presents a good opportunity to examine
patterns of sequence variation in the mitochondrial rRNA genes and the findings of a compa-
rative analysis of primary and secondary structures are presented. The results indicate that these
sequences are useful for reconstructing deep levels of phylogeny, though filtering of the
sequences is necessary in order to remove sites where multiple hits have occurred. The
application of this information to phylogeny reconstruction is also demonstrated.
ZOOLOGIA 95 857
A. Freitag (Lausanne): Ants availability and the decline of the Wryneck in
Switzerland.
Beginning of the fifties, populations of the wryneck (Jynx torquilla L., Picidae) showed
a steady decrease in Europe and particularly in Switzerland (Glutz von Blotzheim & Bauer,
1980). This bird, who is the only migrant species of the Picidae family in Europe, is mostly
dependent on ants (mainly larvae and pupae) of the genera Lasius, Tetramorium, Myrmica and
Formica for rearing his progeny (Hölzinger, 1992; Bitz & Rohe, 1993). Two hypotheses are
proposed to explain this decline: the first one postulates that due to agricultural practices within
the foraging territory of the bird (who is living in agricultural areas like orchards, meadows
with scattered trees) ant populations also declined gradually. The second one assumes that ant
populations are not declining but food accessibility is reduced because epigaeic nests are rare.
Before testing these hypotheses, we have studied the diet of the wryneck in two areas.
Preliminary results are presented and discussed.
References:
Bitz, A. & ROHE, W., 1993. Nahrungsökologische Untersuchungen am Wendehals (Jynx tor-
quilla) in Rheinland-Pfalz. Beth. Veröff. Naturschutz Landschaftspflege Bad.-Württ. 67,
83-100.
GLUTZ VON BLOTZHEIM, U.N. & BAUER, K.M., 1980. Handbuch der Vögel Mitteleuropas. Bd 9:
Columbiformes - Piciformes. - 1148 p., Wiesbaden.
HÖLZINGER, J., 1992. Zur Nestlingsnahrung des Wendehalses (Jynx torquilla). Beih. Veröff.
Naturschutz Landschaftspflege Bad.-Württ. 66, 47-50.
B. Hägele & M. Rowell-Rahier (Université de Neuchätel, Institut de Zoo-
logie, rue Emile Argand 11, 2007 Neuchätel): The performance of three generalist
herbivores in a single/mixed diet treatment.
The generalist herbivores Miramella alpina (Caelifera, Acrididae), Callimorpha domi-
nula (Lepidoptera, Arctiidae) and Cylindrotoma distinctissima (Diptera, Tipulidae) all differ in
mobility and crypsis. For their last instar five groups of twelve individuals were reared on five
different single food plants and one group of twelve individuals was reared on a mixture of
these food plants. For each individual growth rate, duration of last instar, consumption rate,
digestibility of food and the efficiency of conversion of digested food was calculated and
compared between experimental groups. All three herbivores performed best when they could
feed on a mixture of plants. For certain performance parameters however some single plant
groups performed nearly as good as the mixed diet group. Evidence for detoxification costs
could only be found for three plant species in C. dominula. Neither the nutrient complemen-
tation hypothesis nor the noninteractive toxin hypothesis alone could explain the outcome of
the experiment. A combination of both hypotheses could explain the results for M. alpina, but
still failed to explain all results for C. dominula and C. distinctissima.
L. Heer (Zool. Inst. Univ. Bern, Ornitho-Okologie, Prof. U. Glutz von Blotz-
heim; Zell- und Entwicklungsbiologie, Prof. D. Schiimperli): Cooperative Breeding
by Alpine Accentors (Prunella collaris): Polygynandry, Territoriality and Parentage.
The composition of breeding groups, territoriality, parentage and parental care were
examined in a population of Alpine Accentors in the Swiss Alps. Five (1993) to six (1994)
breeding groups were located on a southward hillside between 2500 and 2850 m above sea
level. 3-5 males had almost fully overlapping territories, wherein 2-3 females hold their
exclusive territory (separate nesting, polygynandry). Males competed in long and intense
chases for the access to fertile females within their group territory. The females copulated
promiscuously with all or several males of the same breeding group. The a- and B-males had
858 ZOOLOGIA 95
more access than other subordinate males. They combined frequent copulations and mate
guarding as types of paternity guards. The female performed most of the parental care. 1-4
males helped feeding the young. Results of DNA-profiling showed that members of one group
were seldom closely related. In three out of five sampled broods, multiple paternity occurred
with up to three males siring offspring within one clutch.
B. Imhoof & P. Schmid-Hempel (Ziirich): Within-Host Dynamics of a
Single-Celled Parasite (Trypanosomatidae) in the Intestine of the Bumblebee Bombus
terrestris.
One of the predictions from theory of virulence is that a fast growing parasite should be
more virulent than a slowly growing one. Also, parasites occurring in mixed infections should
display different dynamics than parasites occurring singly inside a host.
To test these hypotheses bumblebees (Bombus terrestris) of four different colonies
were infected with the Tripanosome Crithidia bombi: one group with a fast growing strain, one
group with a slowly growing strain, a further group with both strains and one group was left
uninfected as a control. The bumblebees were kept individually for 22 days, faeces were
sampled every second day and the number of Crithidia cells in these samples were counted.
An infection developed to a peak level between 4-6 days after infection and decreased
to zero around day 16-18. However, the course of the infection was not significantly different
among the different treatments but differed among colonies. Neither "treatment" nor "colonies"
had a significant effect on the lifespan of the bumblebees.
These findings suggest that Crithidia bombi generally is not a virulent parasite.
Probably because it has to assure the production and mating of young infected queens, as well
as their survival of hibernation and egg-laying, to start a new colony next spring. The virulence
of Crithidia bombi does not seem to depend on its growth rate. Furthermore, it seems that its
within-host dynamics could depend on the genetic background of the host.
P. Jeanneret (Station fédérale de Recherches agronomiques de Changins,
1260 Nyon): Landscape influence on insects flying at the interface between two
ecosystems.
Landscape quality and structure should be an important factor to explain insects dis-
tribution and activity, at a regional (e.g. 20km2) or local sécale (e.g. 1km2). The measurement of
faunal exchanges occuring between an agroecosystem and its surrounding habitats is relevant to
test landscape influence on it. In addition, management practices of the cultivated landscapes may
be an important factor influencing the exchanges between ecosystems. The activity of tortricids
(Lepidoptera), flying between apple orchards and their surrounding habitats were used to test this
hypothesis in the region of Nyon - La Còte. A Malaise trap was placed on each of the North, East,
South and West margins of six apple orchards of two types (three were managed with an
integrated pest control program and three were abandonned for many years). Two collectors were
placed on top of the traps to separate emigrant from immigrant individuals. From the beginning
of Jjune to October 1992 and 1993, 48 samples were collected on 14 successive weeks. There
were a high similarity (Mantel test) between emigrant and immigrant tortricid captures, if they
were caught by the same Malaise trap (Jeanneret and Charmillot, 1995). We conclude that there
were neither emigration nor immigration but an activity of tortricids at the interface of habitats.
The hypothesis of the landscape influence on tortricids activity at the interface was
tested at two scales. The first concerned the proximate surroundings of the orchard and the
second the distant surroundings (300m). Proximate and distant surroundings were assigned to 6
categories including woodland, vineyard, cereal crop, orchard, garden and meadow. At both
scales, multivariate analysis (Correspondence analysis, Canonical correspondence analysis) and
associated statistics (multiple regression and canonical coefficients, Monte Carlo permutations
test), showed that landscape explained a significant part of the variation of tortricids caught at
the interface between both types of apple orchards.
ZOOLOGIA 95 859
M. Katoh & G. Ribi (Ziirich): Genetic variation of sympatric and allopatric
populations of hybridizing freshwater snail species.
We studied genetic structure of sympatric and allopatric populations of hybridizing
freshwater snail species Viviparus ater and V. contectus in central Europe. Six allozyme loci
which were informative in Lake Garda, Italy in the previous study (bGAL, GPI, MDH-I, PGM,
and PNP were diagnostic between the two species and PGDH was highly polymorphic in V.
contectus) were analyzed from nine sympatric populations and four allopatric populations each
for the two species. A total of 900 newly collected snails from Italy, Switzerland, and Germany
was examined electrophoretically. V. ater had low genetic variation (the jacknifed mean of
Wright's FsT + S.E. over four loci was 0.041 + 0.004), and contectus-like alleles were found at
low frequencies in both allopatric and sympatric populations. On the other hand, V. contectus
showed high genetic differentiation especially at four loci (GP/, PGM, PGDH, and PNP; the
jacknifed mean of FgT + S.E. over the six loci was 0.546 + 0.166), and no clear differences
between allopatric and sympatric populations were observed. These results indicate that high
allelic substitution rates in V. contectus seem to be responsible for the high genetic dif-
ferentiation. The high substitution rates can be obtained by high mutation rates, introgression,
or small population sizes.
S. Knoll & M. Rowell-Rahier (Neuchatel): Population structure of Oreina
cacaliae (Coleoptera, Chrysomelidae).
The population structure of O. cacaliae was investigated by means of allozyme-
electrophoresis. The sampling was done in a hierarchical design, patches within localities (dis-
tances < 5km), localities within regions (distances 20 - < 400 km) and regions within the
western species range (from the Pyrenees to Czechia). Analyses of 23 presumptive loci en-
coded by 17 enzyme system revealed that most differentiation already occurs between patches
(hierarchical F-statistics, Fsr =0.121 between patches compared to an Fsr of 0.042 between
localities and of 0.036 between regions). Correlation of genetic distances (Nei's genetic
distance) with geographical distances was weak, but significant. Nm (as an indirect measure of
gene flow) calculated from Fsy and from private alleles was smaller than 1, suggesting
restricted gene flow. In summary, O. cacaliae shows considerable genetic differentiation at a
very small scale with evidence for restricted gene flow and isolation by distance.
B. Koch (Zool. Museum Universitat Ziirich-Irchel, 8057 Zürich): Immigration
and Emigration in a Population of Dippers Cinclus c. aquaticus
From November 91 to October 93 the population dynamics of the Dipper Cinclus c.
aquaticus was investigated in the Sihltal near Ziirich, Switzerland. The total number of
individuals changed seasonally. The density varied from 2.7 ind./km at breedingtime to 5.0
ind./km in winter. The sex ratio was 1.63 males to 1 female. 56% of the immigrating and
emigrating Dippers were juveniles. Females started migrating earlier and moved further than
males.
C. Konig and P. Schmid-Hempel (ETH Ziirich): Immunocompetence and
foraging activity in bumble bees, Bombus terrestris L. (Hymenoptera Apidae)
A general assumption of host-parasite interactions is that resistance is costly.
We tested in our experiments whether foraging activity by workers of the bumble bee,
Bombus terrestris L., is costly in terms of resistance against a parasite. Specifically, we
investigated the cellular immune response of the bumble bees against an artificial parasite
inside the hemocoel. We prevented some workers from foraging by clipping their wings. These
workers had a higher immune response then the unmanipulated workers.
860 ZOOLOGIA 95
A. Kopf and N. Rank (Experimental Ecology, ETH Ziirich, ETH Zentrum -
NW, 8092 Ziirich): Species boundaries and geographic variation in host use.
Allopatric populations of the two leaf beetle species Phratora tibialis and Ph. polaris
(Col.: Chrysomelidae) are specialized on two different willow hosts. Ph. tibialis is described to
feed exclusively on Salix purpurea throughout Central Europe, a willow species that does not
occur in Scandinavia. Ph. polaris is usually found on S. phyllicifolia in Scandinavia.
However males and females from different populations from both species mate readily
when given the change and in 3 way choice tests the variability in host preference is large in all
populations.
Morphological comparisons and sequence data from a mitochondrial gene are currently
being gathered to clarify the host use patterns and the status of both species.
H. Krattli & M. Haffner (Zoologisches Museum der Universitat Ziirich,
Winterthurerstrasse 190, CH-8057 Ziirich, Schweiz): Comparative morphological and
microanatomical investigations on the integument of the feet of the edible dormouse
Glis glis Linné and the laboratory rat Rattus norvegicus Berkenhout.
Because the integument of the feet has direct contact to the ground, it is expected to be
adapted to an animal’s locomotion. We have chosen two representatives of Rodentia habitating
completely different living spaces, and used gross morphology and microanatomy on their feet
to find functional adaptations. While the laboratory rat as a typical terrestrial animal is moving
on a mainly horizontal ground, the climbing edible dormouse is a typical inhabitant of trees.
Our investigations were restored by observations and simple experiments in enclosures in the
edible dormouse.
Locomotion on inclined branches requires a bigger frictional force F than on horizontal
ground to avoid slipping. Frictional force F can be increased by increasing the coefficient of
static friction u or by increasing the normal force N, because F = u*N (Amotons’ first law). In
the edible dormouse we found several adaptations which increase these parameters. In com-
parison with the laboratory rat the footpads of the edible dormouse are large and densely
arranged and dimensionally unstable with numerous, specially arranged epidermal ridges on
their surfaces and abundant eccrine glands, whereby the coefficient of static friction p is in-
creased. Frictional force can also be increased by the force of adduction, if the diameters of the
climbed branches allow to be gripped with the extremities. To avoid problems on balancing on
thin branches the first and the fifth digit of the hind foot can be splayed and a giant double-pad
on the forefoot and an enlarged first tarsal pad project out of the remaining sole of the feet and
increase therefore the area of contact against the ground. In comparison with the rat the small,
pointed claws of the edible dormouse are used for climbing on plain vertical surfaces, fast
locomotion and jumping to eliminate the increased force of repulsion in these situations. The
revolved points of the second and third claw of the hind foot are used as climbing hooks on
round branches.
In both species the forefeet are adapted to manipulate food for example. Because the
pads opposite to the digits contain muscles and cartilage, they are an abutment against the force
of the gripping digits.
R. Maurer (Laboratoire d'éthologie, Genève): Connectionist modelling of
behavioural data: the example of path integration.
The modelling of behaviour has traditionally been grounded on mathematical primi-
tives. In spite of the success of some such models, they have one inherent limitation: they are
more a description of the process to be modelled, than an explanation for it. The trouble lies in
the discrepancy between the building blocks of the models (functions) and those of behaviour
(which may more adequately be described by neural primitives).
ZOOLOGIA 95 861
The relatively recent theoretical framework of neural networks provides a new set of
primitives for modelling. Coming in different flavours, some of them including genetic
(mutation/selection) algorithms, this framework may give, in a number of cases, new answers
to (or at least insights into) why behaviour displays particular properties. The target domains
for connectionist modelling include topics as widely different as signal processing in the retina,
place navigation in rats, or female preference for symmetrical males.
A simple example is discussed here, which pertains to path integration (the process by
which an animal uses self-generated signals to keep track of its position). We created limited-
power neural networks that had to compute path integration for a simulated animal. The way
the networks configured themselves in order to realize the computation reflects theoretical
findings about path integration; and the computation, when carried out by the simulated ani-
mals, resulted in a pattern of errors reminiscent of those we had observed in real hamsters.
M. Moeckli & M. Haffner (Zoologisches Museum Universitat Ziirich-Irchel,
Winterthurerstr. 190, CH-8057 Ziirich): Grobmorphologische und histologische Un-
tersuchungen der Mundschleimhaut im Vertibulum oris einheimischer Fledermaus-
arten (Mammalia, Chiroptera).
Für einzelne Individuen aus verschiedenen Fledermausarten wurde eine bilateralsym-
metrische, polsterartige Struktur im Vestibulum oris beschrieben, die je nach Autor und der von
ihm untersuchten Fledermausart verschieden benannt wurde. In der vorliegenden Arbeit wurde
mit grobmorphologischen und histologischen Methoden eine vergleichende Untersuchung tiber
das Vorkommen, die Grobmorphologie und die Mikroanatomie dieses “Buccal pad” bei
männlichen Myotis myotis, Pipistrellus pipistrellus, Nyctalus noctula, Vespertilio murinus und
Plecotus auritus durchgefiihrt. Es zeigte sich, dass alle untersuchten Vertreter der Gattungen
Pipistrellus, Nyctalus und Vespertilio ein “Buccal pad” besitzen, während es den bisher
untersuchten Vertretern der Gattungen Myotis, Eptesicus und Plecotus fehlt. Der Aufbau des
“Buccal pad” ist bei allen Arten im wesentlichen identisch. Es handelt sich um einen Bereich der
Mundschleimhaut, der durch Vergrösserung des Stratum germinativum und durch Einlagerung
von Lipiden in die Zellen des Stratum superficiale eine spezielle sekretorische Funktion ausübt.
Von rostral nach caudal erstreckt sich das “Buccal pad” von der Höhe des letzten Prämolaren
bzw. ersten Molaren bis auf die Höhe bzw. einige u hinter den letzten Molaren. Caudal stülpt es
sich von der Mundhöhle weg in das umgebende Gewebe ein. Aufgrund verschiedener Hinweise
wird eine olfaktorische Funktion des “Buccal pad” in Betracht gezogen und ein Zusammenhang
mit dem Balzverhalten und der Partnerfindung und -wahl, im Sinne einer individuellen
Erkennung, vermutet.
C. Mühlhäuser, W. Blanckenhorn & P.I. Ward (Zürich): The genetic
component of copula duration in the yellow dung fly.
We assessed the heritability of copula duration (h2=0.39) plus its genetic covariances
and the heritability of body size (h2=0.76), development time (h2=0.23) and fluctuating
asymmetry (h?=0.00; n.s.) in the yellow dung fly Scathophaga stercoraria (L.) using full
sibling analysis. The relative order of magnitude of the values agrees with that of other data:
morphological>behavioural>life history characters. No genetic variation of fluctuating asym-
metry, which is often interpreted as the degree of developmental instability, suggests strong
canalisation. Triply repeated measures of copula duration for each male revealed a general
increase with date or age. The individual pattern of this change ("flexibility") in copula du-
ration, an expression of phenotypic plasticity, was itself heritable (h2=0.19). Significant posi-
tive phenotypic and genetic covariance was only evident between flexibility of copula duration
and body size, and between flexibility of copula duration and copula duration itself. All other
correlations were nil, including that between development time and body size and that between
copula duration and body size. We suggest that in S. stercoraria populations there is some
862 ZOOLOGIA 95
averaging stabilizing selection towards a mean optimal copula duration which can serve as a
behavioural "rule of thumb". At the same time, there is selection for phenotypic plasticity that
allows the individuals to adapt their behaviour to their immediate environment. Both effects
together are consistent both with the assumptions of optimality models and the classic results
on copula duration in yellow dung flies.
J. Pawlowski*, I. Bolivar*, J. Fahrni*, L. Zaninetti* & J.P. Debenay**:
(*Département de Zoologie et Biologie Animale, Université de Genève; 154, route de
Malagnou, CH-1224 Chéne-Bougeries; ** Laboratoire de Géologie, Université d'An-
gers, 49045 Angers Cedex, France): Taxonomy of the Foraminifera Ammonia: a
molecular approach.
DNA was extracted from living foraminifera belonging to the genus Ammonia and
identified as A. parkinsoniana, A. tepida, A. beccarii, and A. batava. The cells were collected at
Camargue and Vendée (France), North Harbour (Long Island, USA), Hamana Lake (Japan),
and Plimoth Sound (England).
A 5'-terminal fragment of the large subunit ribosomal DNA (LSU rDNA) was
amplified by PCR using specific Ammonia primers. The region of about 400 nucleotides,
corresponding to the LSU rDNA divergent domain D1 was sequenced and analysed. According
to the ribosomal sequences the specimens identified as A. beccarii and A. tepida form a group
composed of several genetically different "species". Taxonomic identification of these "spe-
cies", based on morphological characters and sequence data, is proposed. The phylogenetic
relationships of these "species" and other Ammonia species are examined.
J. Pawlowski*, I. Bolivar*, J. Fahrni*, L. Zaninetti*, H. Kitazato** & M.
Tsuchyia** (*Département de Zoologie et Biologie Animale, Université de Genève;
154, route de Malagnou, CH-1224 Chéne-Bougeries; **Institute of Geosciences,
Shizuoka University, Oya, 836, Shizuoka 422, Japan): Taxonomc revision of some
Glabratella (Forminifera) based on the sequence of LSU rDNA divergent domain D1.
DNA was extracted from 14 different morphotypes of Glabratella spp., including G.
erecta, G. elegantissima, G. patelliformis, G. nakamurai, and G. opercularis. The cells were
collected on the French littoral zone of the Mediterranean Sea and on the coast of the Japanese
Islands. A 5'-terminal fragment of the large subunit ribosomal DNA (LSU rDNA) was PCR-
amplified using specific foraminiferal primers, then sequenced.
In examined species, the analysis of D1, the divergent domain of the LSU rDNA shows
a genetic similarity between G. elegantissima and G. patelliformis. It allows also to identify the
morphotypes representing the sexual and asexual life cycle generations; of each species and to
determine the seasonal morphological variations. This study of each species and to determine
the seasonal morphological variations. This study demonstrates that the ribosomal DNA
sequences can be used for identification of foraminifera species, providing a new taxonomic
criterion which is independent from the morphological characters of the tests.
M. Pfenninger & B. Streit (Zoologisches Institut, Johann Wolgang Goethe
Universitat, Siesmayerstr. 70, D-60054 Frankfurt am Main): The impact of habitat-
size, life-history and mobility on the genetic structure of populations of Trochoidea
geyeri (Helicidae, Gastropoda).
T. geyeri (Soös, 1926) is a small, endangered terrestrial gastropod of the subfamily
Helicellinae. We have investigated the genetic structure of two different sized populations by
the use of RAPD-fingerprinting. The genetic structure seems to be determined by ecological
factors, especially life-cycle, mobility and habitat-structure. The habitat-size showed no effects
on the genetic variability of the two populations investigated.
ZOOLOGIA 95 863
G. Ribi (Zoologisches Museum, Winterthurerstr. 190, 8057 Ziirich, Switzer-
land): Barriers to gene flow between the hybridizing freshwater snails Viviparus ater
and V. contectus.
Allozyme patterns of Viviparus ater and V. contectus in Lake Garda, Italy, are consistent
with the hypothesis of introgression in both directions. Here I present data on several mechanisms
influencing the amount of introgression going on, such as hybrid fecundity, species assortative
mating or differential fertilization of eggs. In captivity, V. ater and V. contectus produce hybrids
with a male biased sex ratio (84% males) and a high survival rate (99% in the first year). Hybrids
are fertile inter se and in backcrosses with both parental species. Most backcrosses have a high
fecundity (50-100% of intraspecific control crosses), and backcross offspring have a 1:1 sex ratio
and a survival rate of >80% in the first year. Heterospecific matings are frequent in nature.
Preliminary results of an interspecific sperm competition experiment indicate that V. contectus
sperm have only a slight disadvantage in fertilizing V. ater eggs as compared to V. ater sperm.
All these results taken together would suggest that interspecific hybrids should be common at
places where the two species coexist. However, hybrids are rare in nature (<1%). The possible
role of segregation in space or time as isolating mechanisms is discussed.
C.H.F. Rowell & P. Flook (Basel): The phylogeny and classification of the
Caelifera (Orthoptera): first results of a molecular systematic investigation.
We have used the 12S and 16S mtrRNA genes of some 80 taxa from 37 different
subfamilies to construct a preliminary phylogeny of the caeliferan Orthoptera (short horn
grasshoppers and their allies) at the subfamily level. The results support at the superfamily level
one of the standard morphological classifications, viz. Tridactyloidea (Tetrigoidea (Eumasta-
coidea (Pneumoroidea (Pamphagoidea (Acridoidea))))). The traditional grouping of the sampled
genera into subfamilies or families is also largely confirmed. The Eumastacoidea is probably
paraphyletic; its different lines are genetically distant from each other, suggesting old, well
differentiated families. The Pamphagoidea also appears to be a paraphyletic grouping; the
Pyrgomorphidae is the oldest branch, the Pamphagidae more recent, while the Lentulidae may be
polyphyletic. The Acridoidea sensu strictu is monophyletic, but the divergence of the different
taxa within it is mostly poorly resolved. This may indicate either that the mtrRNA genes are
unsuitable for this purpose, or that the Acridoidea underwent a very rapid early radiation.
L. Rüber! & E. Verheyen? (!Zoological Museum of the University of Zürich;
?Royal Belgian Institute of Natural Sciences, Brussels): Convergent evolution of tro-
phic specializations within the endemic Tanganyikan cichlid tribe Eretmodini, inferred
from mtDNA sequences (Pisces: Cichlidae).
The Tanganyikan cichlid species flocks contain more than 170 species assigned to 49
genera and represent one of the most spectacular examples to study the evolutionary mechanisms
involved in adaptive radiation and ‘explosive’ speciation among living vertebrates. Based upon
morphological features, these species assemblages have recently been classified into 12 tribes. In
an attempt to unravel the phylogenetic interrelationships among and within these tribes a number
of recent studies implement molecular techniques, mainly mtDNA sequencing, to study the rates
and modes of speciation within these speciesassemblages.
Two mitochondrial gene segments from 51 Eretmodini specimens from 32 localities were
amplified and direct sequenced. The obtained mtDNA phylogeny is in partial conflict with the
current generic classification of the Eretmodini that is mainly based on dental morphology. In
contrast to previous studies the present data suggest that the species originally assigned to the
three genera Eretmodus, Spathodus and Tanganicodus represent more than three genetically and
morphologically distinct lineages that show a high degree of intralacustrine endemism. Our data
Suggest that the taxonomy of the Eretmodini should be reinvestigated and that in this tribe,
trophic specializations may be particularly prone to convergent evolution.
864 ZOOLOGIA 95
F. Saucy & J. Studer (Fribourg): Preference for acyanogenic white clover in
the vole, Arvicola terrestris.
A strong emphasis has been recently put on the potential role of plant secondary
metabolites as a force driving the population cycles exhibited by small rodents. In this study we
focus on the reciprocal interactions between the vole, Arvicola terrestris, and white clover,
Trifolium repens (one of the few preferred food items of the vole). The production of cyanogenic
glycosides (liberating HCN when the plant is attacked) as a defence against various herbivores
(insects, slugs, snails and mammals) is controlled by a single genetic system involving two genes.
The maintenance of a polymorphism for cyanogenesis in natural populations of white clover is
poorly understood and could be maintained by a selective pressure of herbivores on different
cyanotypes. In this study, we report a marked preference of the voles for an acyanogenic white
clover cultivar which was preferentially eaten during tests conducted in cages and in outdoor
enclosures. Preliminary field data gathered under low and high vole population densities are also
presented.
V. Séguinot (Laboratoire d'éthologie, Genève): Path integration across spe-
cies: mechanism and ecological implications.
Path integration (or dead reckoning) is an innate navigational strategy by which an
animal that has left some known place (generally its nest or a food source) can return to the
starting-point of its trip. Being entirely based on signals self-generated during locomotion, this
process occurs whether or not visual landmarks or a directional reference from the environment
can be used.
Although path integration has been evidenced both in invertebrates and vertebrates,
little is known concerning the mechanism that lies at the core of this short-distance navigation
system in animals and humans. Since biological systems do not work with infinite precision but
involve errors, one possible way to investigate the rules of the mechanism of path integration is
to analyse the nature and the shape of its errors. Biases observed after an asymmetrical (e.g. L-
shaped) journey are discussed.
Moreover, random and systematic errors alike provide us with some hints about the
functional aspects of the mechanism underlying path integration: ecological implications are
proposed. Indeed, the fact that errors are a widespread phenomenon among species and that
there was no selective pressure to correct them leads us to consider such errors as being the
result of adaptative selection.
R. Spaar (Schweizerische Vogelwarte, Sempach): Flight behaviour of Steppe
Buzzards (Buteo buteo vulpinus) during spring migration in Southern Israel.
Raptor migration was studied in Israel by tracking radar in the Arava Valley near
Hazeva (150 m below sea level) and in the Negev Highlands near Sede Boger (470 m above sea
level). Entire gliding and soaring phases were recorded. The flight altitude of the Steppe
Buzzards Buteo buteo vulpinus depended on the time of day. It increased from morning (first
birds were tracked at about 8 a.m.) until mid afternoon when maximum altitudes, about 2000 m
above ground level in the Arava and 1000 m in the Negev, were reached. Flight altitudes
decreased only during the last two hours before sunset. Migration was lifted slightly higher
above ground and lasted longer towards sunset in the warmer Arava Valley than in the Negev
Highlands. The relatively low mean climbing rate while soaring in thermals of 2 m/s can be
explained by the fact that the whole diurnal circle is included in this average and that the
observation sites were not situated at the edge of large rocky slopes, where strong thermal
updrafts occur, but rather on flat surfaces. Climbing rates ranged from 0.5 up to 4.5 m/s. Strong
thermals were used longer for soaring than weak ones. The climbing rate in thermals was a
decisive trait when explaining the flight behaviour of migrating Steppe Buzzards: the flight
altitude, the airspeed while gliding, and consequently the cross-country speed increased with
ZOOLOGIA 95 865
increasing climbing rate. Thus, the birds adjusted the airspeed while gliding to the actual
climbing rate in order to maximise the cross-country speed. The average cross-country speed
was 9.8 m/s and was considerably enhanced by tail wind.
H.-P. Stauffer, P. Schmid-Hempel (ETH Ziirich): Parasitic larvae of conopid
flies alter flower choice of Bumblebees.
Bumblebees (Apidae, Hymenoptera) are among the most important pollinators of crop
plants. They are also host of a wide range of parasites. In the study presented here, we show
that the flower choice of bumblebees is affected by the larvae of the parasitoid conopid fly
(Conopidae, Diptera), which develops in the abdomen of bees. We offered unparasitised and
parasitised bumblebees different plant species in cages, and noted the behav iour and flower
choice of the bees. Parasitised bees visited the most common plant species T. pratense
(Fabaceae) less often than unparasitised ones. it might be that the parasite impairs the sensory
capabilities of the bees or that parasitised bees avoid 7. pratense, because of its complex
morphology.
L. Sundstrôm (University of Bern, Ethologische Station Hasli, Wohlenstrasse
50a, 3032 Hinterkappelen): Sex allocation in ants - the impact of kinship and
environment.
Hamilton’s rule and the concept of inclusive fitness provide a theoretical basis for
predicting reproductive characteristics of social insect colonies. Conversely sex allocation studies
in social insects can be used to test inclusive fitness theory itself, the most powerful test being the
analysis of individual colonies where the predicted sex allocation varies due to differences in
worker fitness functions. The optimal sex allocation ratios may differ for queens and workers,
causing a worker-queen conflict over sex allocation. The degree on conflict is highest in colonies
headed by one singly mated queen and decreases with the presence of multiple queens as well as
multiple mating of queens. However, also other factors, such as differential dispersal of the sexes
and variations in resource availability, can affect sex allocation. This study analyses simul-
taneously the impact of kin structure, colony productivity and mating pattern on sex allocation
under high vs low levels of worker-queen conflict. The results show that the kin structure is the
main (but not the only) determinant of sex allocation under high levels of worker-queen conflict,
whereas ecological factors best explain the allocation pattern under low levels of conflict.
C. Wedekind’, M. Chapuisat°°, E. Macas* & T. Rülicke** (°Abt. Ver-
haltensökologie, Zoologisches Institut, Universität Bern, 3032 Hinterkappelen;
°°Musée Zoologique and Institut de Zoologie et d'Ecologie Animale, Université de
Lausanne; *Departement fiir Frauenheilkunde, Universitätsspital Zürich; **Biolo-
gisches Zentrallabor, Universitätsspital Zürich): Condition-dependent choice for
MHC combinations during fertilization in mice.
One evolutionary explanation for the success of sexual reproduction assumes that sex is
an advantage in the coevolutionary arms race between pathogens and hosts. Accordingly, an
important criteria in mate choice and maternal selection thereafter is expected to be the allelic
specificity on polymorphic loci involved in parasite-host interactions, e.g. the MHC (major
histocompatibility complex). The MHC has been found to influence mate choice and selective
abortions in mice. However, it could also influence the fertilization process itself, i.e. 1) the
oocyte's choice for the fertilizing sperm and 2) the outcome of the second meiotic division after
the sperm has entered the egg. We tested both hypotheses in an in vitro breeding experiment
with two inbred mouse strains congenic for their MHC. The genotypes of the resulting blasto-
cysts were determined by polymerase chain reaction. We found non-random MHC combi-
866 ZOOLOGIA 95
nations resulting from both possible choice mechanism. The outcome changed over time,
indicating that choice for certain MHC-combinations is influenced by one or several condi-
tional factors, too.
C. Wedekind°, T. Seebeck*, F. Bettens# & A. J. Paepke° (°Abt. Ver-
haltensökologie, Zool. Institut, Universität Bern, 3032 Hinterkappelen; *Institut für
Allg. Mikrobiologie, Universität Bern; # Institut für Immunologie und Allergologie,
Inselspital Bern): MHC-dependent mate preferences in humans.
One substantial benefit of sexual reproduction could be that it allows animals (or
humans) to rapidly react to a continuously changing environmental selection pressure such as
coevolving parasites. This counteraction would be most efficient if the females were able to
provide their progeny with certain allele combinations on loci which may be crucial in the
parasite-host arms race, e.g. the MHC (major histocompatibility complex). Here, we show that
the MHC influences both body odours and body odour preferences in humans, and that the
women's preferences depend on their hormonal status. Female and male students were typed for
their HLA-A, -B and -DR. The men weared a T-shirt during two nights. On the following day,
the women were asked to judge the odours of six T-shirts each. They scored male body odours
as more pleasant when they differed from the men in their MHC than when they were more
similar. This difference in odour assessment was reversed when the judging women were
taking oral contraceptives. Furthermore, the odours of MHC-dissimilar men remind the test
women more often of their own actual or former mates than do the odours of MHC-similar
men. This suggests that the MHC influence human mate choice today. (in press: Proc. R. Soc.
Lond. B, 1995)
E. Wullschleger, N. Zambelli & P.I. Ward (Ziirich): Differences in habitat
choice and life history traits in two shell forms of the freshwater snail Lymnaea
peregra.
The freshwater pulmonate snail Lymnaea peregra (Müller) shows two shell forms
which are favoured in different habitats. Forma peregra with its long and narrow shell can
relatively easily retreat into mud when a water body dries. Forma ovata on the other hand has a
wider and more compact shell which allows it to withstand strong water movements, an
important environmental factor on shores where waves and currents are not reduced by
vegetation.
We compared growth and reproduction of ovata and peregra at constant and at
declining water depth. Reproduction was reduced in both forms by declining water level, while
growth of the form peregra was less affected by this treatment than growth of ovata. Peregra
grew faster also by contstant water level.
To measure their habitat choice, snails were kept in containers with two substrate types
at a number of water depths. Additionally, abundance data of both forms were determined
along the shore gradient of water depth in a sedge reed at Seealpsee (Appenzell, Switzerland)
where they occur sympatrically. Ovata avoided shallow waters compared to peregra in the field
as in the laboratory. Peregra preferred muddy substrate whith vegetation to stones, while ovata
from the same origin habitat showed no substrate preference. Allozyme differences between the
forms suggest that shell form and thus possibly also life history traits or habitat choice are at
least partially determined genetically.
A. Wyttenbach & L. Favre (Lausanne): Genetic polymorphism as revealed
by microsatellite DNA in Sorex araneus (Insectivora, Mammalia) and its use for
population genetic studies
ZOOLOGIA 95 867
Studying the interactions between genetics and ecology one frequently needs to
measure genetic variation in a population. Often low variation is found due to the population
history (bottlenecks, founder effects), due to the reproduction modus of the species (selfing
species) or due to the genetic marker applied in the study. Low levels of genetic variation can
also result from low density populations or from small effective population sizes.
The common shrew (S.araneus) is a case where classical genetic markers such as
allozymes revealed very low genetic polymorphism on a intra/interpopulation level. Because of
its spectacular chromosomal evolution which lead to the existence of more than 30 chromo-
somal races S. araneus became a model species to study the link between genetic and karyo-
typic evolution, the formation of new races and their underlying ecological mechanisms. The
lack of polymorphic markers on a intra/interpopulation level however made it impossible to
study the population structure and its implication in the chromosomal evolution of this species
(e.g. fixation of chromosomal mutants). Similarly, studies in hybrid zones between different
chromosomal races investigating the effects of chromosomes to diminish gene flow and thus
playing a role in speciation processes were hindered. For these reasons microsatellite markers
were developed.
Species specific primer sequences were designed for 10 microsatellite loci. In a first
step 5 microsatellites were tested for their polymorphism on a intra/interpopulation level.
Depending on the locus the number of alleles per locus ranged from 2- 20 and the observed
heterozygosity varied between 0.50- 0.90. Individuals from each side of 2 hybrid zones
between different chromosomal races were also tested in order to look at the level of dif-
ferentiation of microsatellite DNA on a racial scale. Preliminary results suggest that micro-
satellite loci can also be applied in population genetic studies on a racial scale.
Microsatellites are the first highly polymorphic, nuclear markers on a population level in
S. araneus. Therefore microsatellite loci will be an important tool to get insight about the role of
the population genetic structure on the chromosomal evolution in S. araneus. A more detailed
testing of microsatellite polymorphism between different chromosomal races will show to what
extend this kind of genetic marker can be used for studying the fine structure of hybrid zones.
S. Zschokke (Basel): Early stages of orb web construction by Araneus dia-
dematus.
The first stage of orb web construction is the least studied and the most poorly under-
stood one, partly because the behaviour of the spider at that stage lacks the repetitiveness of later
stages, partly because the timing is unpredictable and partly because the moves of the spider
cannot be deduced from the finished web.
In our study, we recorded all moves of the spider during web construction using com-
puterised image analysis. From these recordings and from direct observations we could then
reconstruct the exact and complete web construction for several webs.
We analysed these constructions quantitatively (how long did the spider take and how
long did the spider walk to build a web) and qualitatively (how did the spider build the web).
The first stage (the so called exploration stage) proved to be the more variable in all
respects than the later stages.
We could not find a clear strategy the spider uses to explore a new site, but we found
several behavioural patterns we could observe repeatedly. This lead us to the hypotheses that the
exploring phase of a spider consists of a series of fixed behavioural patterns in random order.
These patterns are ‘designed’ in a way to make it very likely that a proto-hub will emerge. The
proto-hub then forms the basic framework of the web. As soon as a proto-hub had been
established, the behaviour of the spider became more stereo-typed and predictable.
REVUE SUISSE DE ZOOLOGIE, 102 (4) : 869-882; décembre 1995
Allozymes in evolutionary genetics: beyond the twin pitfalls of
“Neutralism” and “Selectionism”’*
Ward B. WATT
Dept. of Biological Sciences, Stanford University,
Stanford, CA 94305-5020 USA,
and
Rocky Mountain Biological Laboratory, P.O. Box 519,
Crested Butte, CO 81224-0519 USA.
Allozymes in evolutionary genetics: beyond the twin pitfalls of “Neu-
tralism” and “Selectionism”. - Population genetics arose as the algebraic
study of genetic transmission in Mendelian populations, showing the
genetic feasibility of Darwinian evolution by natural selection. At the same
time, it established a tradition of bypassing study of the biological mecha-
nisms by which evolution proceeds. This view of evolutionary dynamics as
a primarily algebraic, statistical matter led directly to the “neutralist-
selectionist” controversy over the meaning of extensive allozyme variation
in the wild. Much argument failed to resolve this controversy, demons-
trating that population genetics by itself is necessary but insufficient for
full understanding of evolution, due to inherent limitations in the scope of
its analysis. A broader concept of evolutionary genetics is proposed, using
allozymes and/or other natural genetic variants as biologically specific
probes of putatively adaptive, constrained, or neutral phenotypic states, and
analyzing the translation of resulting genotypic differences in organismal
performance into fitness consequences. This approach includes informed
use of population-genetic concepts, but does not ask them to do more than
they can do. Diverse examples are cited in support of the feasibility and
productivity of this approach. Evolutionary biologists are challenged to
bring mechanistic biology into Darwinian context, to move beyond the
remains of the “neutralist-selectionist” controversy and of purely formal
approaches to evolution, to seek biologically based generalities about the
evolutionary process.
Key-words: Allozymes - Biological specificity - Evolutionary genetics -
Evolutionary mechanisms - Fitness differences.
* Main lecture presented at Zoologia 95.
870 WARD B. WATT
ALLOZYMES AND POPULATION GENETIC HISTORY
POPULATION GENETICS: AN ABSTRACT ELEMENT IN EVOLUTIONARY STUDY
Population genetics, as is well known (PROVINE 1971), developed in the first
part of this century through the joint efforts of FISHER (1930), HALDANE (1932), and
WRIGHT (1931). Its first great success was showing that even small selection pressures
could be effective in evolution, given sufficient effective population size, N., and
time in which to work (FISHER 1930). Another success was explaining the population-
level interaction of causal and non-causal elements — natural selection vs. genetic drift
and/or inbreeding — in evolution (WRIGHT 1931), as in the loss of deleterious but
meiotic-drive-maintained f-alleles in mice, a “helpful” effect of genetic drift in small
populations (LEWONTIN & DUNN 1960). There has even been success in explaining
constraints on evolution which arise from the genetic transmission system — e.g., in
the failure of the favorable hemoglobin CC homozygote to predominate in human
populations under malarial selection, because of the disadvantage of C-carrying
heterozygotes, unless genetic drift and/or inbreeding intervene (TEMPLETON 1982).
However, population genetics by itself has little explanatory power other than
via properties of the transmission system. It does not have language or, facts to probe
the phenotypic bases of why genotypes differ in Darwinian fitness, if indeed they do.
The maintenance of human hemoglobin polymorphism was explained by mechanistic
biology outside population genetics (INGRAM 1962) — which may be part of why some
regard it as an atypical “special case”, rather than simply a specially striking one.
Indeed, population genetics has come to support an a-, or even anti-, mechanistic
tradition in evolutionary study, traceable to FISHER. His “Fundamental Theorem of
Natural Selection” — “the rate of increase in fitness is equal to the genetic variance in
fitness at that time” (1930) — is a base of this tradition, implying as it does that the
control of evolution is primarily a matter of statistics. This trend was reinforced by
the terms in which the “Modern Synthesis” of Neo-Darwinian evolutionary concepts
was expressed (JEPSEN et al. 1949). There, genes were treated as abstracted agents
within a formalized phenotypic architecture, despite a discussion of genotype
phenotype transformations by STERN (1949) which seems insightful even now. Later,
we find MAYR (1958) writing, admittedly speculatively, about genes as “jacks-of-all-
trades” or “good mixers”, irrespective of what functions those genes might actually
support.
Evolutionists’ reluctance to consider the specificity of gene action in evolution
has continued. The last two decades of evolutionary study have seen strong emphasis
on classical quantitative genetics (FALCONER 1989), which bypasses study of most
phenotypic mechanisms by assuming that nearly all traits are governed by many
polygenes whose alleles are of very small individual effect. This is an extension of
Fisher’s “micromutationism”, and in some respects a heuristic one (cf. ENDLER 1986),
but in other respects an overemphasis which goes beyond the evidence (ORR &
Coyne 1992). Happily, leading quantitative geneticists are now asserting the need for
more mechanistic research strategies, and for evaluation of alleles with large effects at
ALLOZYMES IN EVOLUTIONARY GENETICS 871
small numbers of loci, as complements to or extensions of the classical quantitative-
genetics program, e.g. LAI et al. (1994), TRAVIS (1994), VIA (1994 and pers. comm.).
THE “NEUTRALIST-SELECTIONIST” DEBATE: HEAT WITHOUT LIGHT
Among the three founders of population genetics, HALDANE may have been the
most consistently mechanistic in his thinking, though highly committed to the use of
mathematical reasoning about evolution (1932). (WRIGHT was not far behind, though,
given his insight into metabolic organization: 1934). This is not surprising, as
HALDANE helped found the theory of enzyme kinetics and thus laid one important
corner of the foundation of all modern molecular biology. PAULING et al. (1949)
identified the first of what came to be called allozymes, hemoglobin S; by 1954,
HALDANE wrote:
‘“... a number of enzymes have been found... to consist of several constituents separable in an
electric field. Such a mixed enzyme would be a definite advantage in an organism either
subjected to a wide range of environments or with a variety of chemically differentiated
tissues.”
At one stroke, HALDANE here anticipated study of tissue-specific isozymes in
development, physiology, and systematics (e.g. MARKERT et al. 1975, WHITT 1987)
and study of allozyme variants as probes of putative adaptive interactions between
phenotypic mechanisms and the environments in which they evolve (e.g. WATT
1985a, b, Powers et al. 1991). Ironically, population genetics, dominated by formal
theory as it became in part through HALDANE’s advocaty, was for a long time quite
uninterested in his empirical foresight. HUBBY & LEWONTIN (1966), finding much
natural genetic variation in enzymes of digestion, intermediary metabolism, etc.,
nonetheless held a view of allozymes as being a “random sample of the genome”,
which from a functional viewpoint was absurd. Often, studies of function and fitness
impacts of allozymes were met by population geneticists with disinterest (“Oh, that’s
just physiological ecology, not evolution!”). If human hemoglobin variation was seen
as a special case, then the mindset of the field was such as to avoid recognizing
evidence which might threaten that view. This empirical indifference led to the
“neutralist-selectionist debate”, which consumed the attention of many evolutionists
for an unproductively long time.
While admitting the clarity of hindsight, one must see this “debate” as
empirically ill founded from the outset. The power of KIMURA’s theory of neutral
allele evolution (1983) cannot be doubted, as a null model in the same sense that the
Hardy-Weinberg distribution or linkage equilibrium are null models at simpler levels
of study. But whether it describes a given class of variants in the wild is an empirical
question, and neither data nor biological knowledge at large fared well at the hands of
extreme neutralist-selectionist debaters. The distributions of fitnesses asserted by both
extreme positions reveal their failings (GILLESPIE 1991, TURNER 1992; Fig. 1). What,
in phenotypic terms, could cause the absence of genotypes with relative fitnesses of
1.0 1/Ne, as demanded by extreme “selectionism” (Fig. la)? Equally, only a naive
optimalist — holding that this is already the best of all evolutionary worlds — could
872 WARD B. WATT
Selectionist assumption
Number of genotypes
0 1
Relative fitness
Neutralist assumption
Number of genotypes
0 1
Relative fitness
Fic. 1
Alternative distributions of genotypic fitnesses according to the extreme views at issue in the
“neutralist-selectionist” debate, as redrawn from GILLESPIE (1991) and TURNER (1992). “Selec-
tionist assumption” shows essentially no genotypes with relative fitness within + 1/Ne of the
value 1.0. “Neutralist assumption” includes the possibility of the “nearly neutral, slightly dele-
terious” variation of the neutral theory. Fitnesses substantially less than 1 of course represent
the “victims” of stabilizing or purifying selection.
ALLOZYMES IN EVOLUTIONARY GENETICS 873
justify the extreme neutralist view that almost no genetic variants have relative fitness
larger than 1.0 + 1/N, (Fig. 1b). Whatever the shape of the actual distribution may
prove to be for any class of genes, there is no biological basis a priori for
hypothesizing either extreme truncation at arbitrary relative fitness, OR extreme
discontinuity to “avoid” neutrality.
It soon became clear that, given feasible sample sizes, even quite different
models of allozyme variation could not be resolved with any statistical power (EWENS
& FELDMAN 1976, WATT 1985a). Indeed, models with opposed assumptions (e.g. the
infinite-alleles neutral model, KIMURA 1983, and the SAS-CFF fluctuating-environ-
ment selection model, GILLESPIE 1978), which share the statistical property of
exchangeability among genotypes studied, generate closely similar genetic frequency
distributions as a result (ROTHMAN & TEMPLETON 1980, GILLESPIE 1978, 1991, 1994).
For example, SKIBINSKI & WARD (1982) claimed support for applying the infinite-
alleles neutral model to an allozyme data set, because substitution rate was positively
correlated with heterozygosity as that model predicts. But, GILLESPIE (1994) showed
that selection-based models also predict a very similar correlation, and that by choice
of coefficients, models of quite contradictory assumptions can coincide in predictions
of frequency distribution! Thus there seems little point in such efforts as, e.g.,
SKIBINSKI et al.’s (1993) further elaboration of the correlation between intraspecific
heterozygosity and interpopulation or interspecies genetic distances, and claim of
“support” from this for the explanation of allozyme variation by the neutral theory.
“Support” here is ambiguous at best, because exchangeable variation will also “sup-
port” diverse selection models. What is needed for any real test is the ability to
REJECT alternatives, but this is just what the common presence of exchangeability
seems to deny to this entire approach.
KUHN’s (1970) view of science, as dominated by scientists’ commitment to
paradigms and hence their resistance to diverging views, describes this disappointing
history very well. Some biologists or philosophers object to Kuhn’s seeming insis-
tence that science must work this way, but this cannot undermine the observation that
it often does work this way, though it need not do so. It is now clear that the a- or
anti-mechanist paradigm for evolutionary study, explaining natural variation and
evolutionary processes in purely statistical-population-genetic terms as in the neu-
tralist-selectionist debate, has only limited power. It cannot answer some clear ques-
tions about clashing views, and cannot even INCLUDE certain kinds of evolutionarily
important, mechanism-oriented questions. Such problems are clear signs of paradigm
crisis (KUHN 1970). A broader, less-self-limiting, more inclusive evolutionary gene-
tics is needed.
WHAT ALTERNATIVES DO WE HAVE?
FEDER & WATT (1992) argued that population genetics is necessary, but
insufficient, for understanding evolution, because it covers only the last of four stages
in the recursive evolutionary process: fitness — genotypes, wherein fitness coeffi-
cients, effective population sizes, and inbreeding coefficients interact to yield the next
874 WARD B. WATT
generation’s starting genotype array. By inquiring what rules might govern the first
three stages — genotypes — phenotypes, phenotypes — performances, perfor-
mances — fitnesses — we may approach a truly evolutionary-genetic theory within
which population genetics is indispensable, but not predominant.
With respect to allozymes, we should expect a wide range of evolutionary
impacts, in terms of kind and strength of effects on phenotype, hence on organismal
performance — from behavior as “major genes”, down through behavior as part of
“oligo-genic” and polygenic systems (which should be highly “exchangeable”, cf.
above), to some truly null effects. These should produce a corresponding range of
fitness component values. Even “neutrality” itself may have different meanings. To
put the matter in terms of absolute fitnesses (= net reproductive rates, Rg), a genotype
with mean Ro = 1.6000 and another with mean Ro = 1.6001 would be neutral with
respect to one another unless N. is > 10,000, yet either would still be favored
compared to a genotype with Ro = 1.0, even if N. were only 50! This is a very
different situation from that envisioned by the classic neutralist view (Fig. 1b).
Thus, to seek biological generality about evolution, we must study the specific
biology of gene action and phenotype-environment interaction, in ecological and
evolutionary context, more carefully. We cannot ask “population genetics” to do more
than it can. It is indispensable in integrating fitness differences with population-
structure and genetic transmission rules to yield new distributions of genotype
frequencies. The origins of the elements to be integrated must be studied with other
tools.
ALLOZYMES AND THE SCALE OF THE GENE ACTION IN THE WILD
ALLOZYMES AS MAJOR LOCUS-SPECIFIC PROBES OF PUTATIVE ADAPTIVE STATES
In Colias butterflies of lowland North America, 10 allozyme genotypes of
phosphoglucose isomerase, PGI, differ markedly in kinetics and thermal stability (e.g.
WATT 1983, 1992). These properties are negatively correlated, i.e. they “trade off”, in
homozygotes. Some but not all heterozygotes are superior in kinetics. Phenotypic
“neutrality” may occur even in the midst of major differences, e.g. genotypes 4/4 and
4/5 are equally kinetically poorand thermally stable, while they differ by up to 4-fold
in these properties from sister genotypes (cf. above). The biochemical differences
predict genotype-specific differences in flight capacity in the wild (WATT et al. 1983),
as this enzyme is central to energy resupply for flight (see WATT 1992 for a review of
this metabolic context). The flight capacity differences in turn lead to genotype-
specific differences, again up to several-fold in size, in survivorship, male mating
success, and female fecundity (WATT 1983; WATT et al. 1985, 1986; WATT 1992, and
Table 1 as an example.) More fitness component studies in extreme environments will
complete our understanding of this selection regime, which appears now to be a mix
of simple heterozygote advantage with patchy-environment selection such as studied
by LEVENE (1953), GILLESPIE (1978), or HALDANE & JAYAKAR (1963).
ALLOZYMES IN EVOLUTIONARY GENETICS 875
TABLE |
Temperature and fecundity of female Colias vs. PGI Genotype
Comparative daily fecundities of PGI genotypes of female Colias philodice eriphyle Edwards
in August 1989 and August 1990 in an outdoor cage assay at Gothic, Colorado, elevation 2900
m, with monitoring of thermal balance in the cages, re-tabled from WATT (1992). Genotypic
differences were significant with P < 0.001 in a General Linear Model analysis which also
included significant effects of available flight time per day, increasing fecundity, and female
age, decreasing it.
% time in Tp range Mean eggs/day + std. error of mean
flight optimum overheat PGI genotypes
possible 2/35 Dla Bist 3/48 B/S NA AT 74/5
29-40°C 35-39°C > 40°C
6.3-89.7 0-60.3 0-1.4 8.07 7-8 OGM A CAEN C6
ea lige) ee Te See
n= 54 16553915 72589, 297212357 28
We have extended this work to an alpine congener, C. meadii Edwards of the
Rocky Mountains (WATT et al. 1995), which is also highly polymorphic for PGI.
Though C. meadii PGI alleles have similar electrophoretic mobilities to those of
lowland Colias on ordinary-resolution gels, they are electrophoretically distinct in
high-resolution gels. Moreover, while there is heterozygote advantage in kinetics and
a tradeoff of kinetics vs. stability in homozygotes, just as in the lowland species,
“similar” genotypes are not closely similar in function between the taxa. Other North
American Colias taxa are also polymorphic at this allozyme gene. Is this divergence
from ancestral polymorphism, or is it of recent and parallel origin? Whichever answer
emerges from future phylogenetic study, this is an anti-adaptationist, constraint
oriented result: it implies that there are severe limits, perhaps at the level of protein
structure, on the ability of alternative alleles to maximize functional properties when
homozygous.
A number of other allozyme case studies, on diverse creatures, have also found
major genotypic effects on organismal performance and thence on fitness components
(BURTON & FELDMAN 1983, POWERS et al. 1991, WATT 1994). One which has been
pursued by several research groups over the years is that of polymorphism in
Drosophila pentose shunt enzymes (e.g. BULSMA & VAN DELDEN 1977, CAVENER &
CLEGG 1981). LABATE & EANES (1992) demonstrated that a 40% difference in kinetics
among G6PD genotypes results in a 32% difference in in vivo pentose shunt flux,
establishing a very specific mechanistic basis for observed functional differences at
higher phenotypic levels. EANES et al. (1993) have now found molecular-level evi-
dence that not only G6PD allozyme polymorphisms, but also interspecifically diffe-
rent amino acid substitutions in G6PD, are due to the action of selection. Allozymes
can find their main effects in diverse phenotypic subsystems: for example, phospho-
876 WARD B. WATT
glucomutase, PGM, genotypes interact strikingly with social structure alternatives in
fire ants (KELLER & Ross 1993). It has been said that allozymes “may affect
metabolism, but not complex traits of interest to evolutionists” — but look here! the
referenced cases include major allozymic effects, via metabolism, on traits as com-
plex as flight capacity, swimming speed, osmoregulation, even reproductive caste
development — all of which, in turn, have effects on fitness components such as
survivorship, mating success or egg ooutput.
ALLOZYMES AND "EXCHANGEABILITY” IN THE MIDDLE RANGE OF EFFECTS
We do not expect all allozymes to have such dramatic effects as those dis-
cussed above. Moderate magnitudes of genotypic functional differences, intermediate
roles in pathways, and relative peripherality to the energy budget (WATT 1985b)
should cause many variants to fall into the oligogene/polygene range of functional
differences and of fitness consequences. One example studied on a single-locus basis
is that of Drosophila a-glycerophosphate dehydrogenase, whose natural polymorphs
have detectable effects, but rather small ones (BARNES et al. 1989). More such cases
are undoubtedly included in findings of multilocus heterozygosity correlations with
organismal performances of various kinds — oxygen consumption, growth rate, etc.
(e.g. KOEHN & SHUMWAY 1982). In the case of the oyster Crassostrea gigas, an
allozyme locus contributing to such a multilocus pattern has later been examined
genotype-by-genotype, and heterozygote advantage in specific activity was found
(POGSON 1991). Sometimes, of course, such multilocus-heterozygosity correlations
with organismal performance have not been found (e.g. SAVOLAINEN & HEDRICK
1995). Which test organisms, habitat structures or stringencies, and allozyme loci do
or do not display such correlations may, if considered thoughtfully, yield new insight
into the whole phenomenon. This can only hapen if interested workers avoid the trap
of arguing for exclusive generality of their own findings, such as plagued the
neutralist-selectionist debate. If biological specificity is as important as this review
contends, then different allozymes in different creatures may show a diversity of
potentially general patterns, but not simple uniformity.
Another way to test the meaning of such “rank-and-file” allozyme variation is
to compare its biogeographic OR organismal-performance patterns against those of
unrelated groups of genes which can be identified at the molecular level. Thus KARL
& AVISE (1992), studying the oyster Crassostrea, found that uniform distributions or
shallow clines of allozyme frequencies along the Florida coast contrasted to both
nuclear and mitochondrial DNA restriction fragment data showing low gene exchange
between Atlantic and Gulf Coast oysters; they inferred that the allozymes must be
subject to significant selection. POGSON & ZOUROS (1994) restudied the correlation of
allozyme heterozygosity with organismal performances as above. When found, this
has been ascribed to direct allozyme effects, or else to associative effects of genes
closely linked to the allozymes. If the latter were so, the performance correlation
should also occur with heterozygosity of other molecular markers, not just allozymes.
ALLOZYMES IN EVOLUTIONARY GENETICS 877
When tested in the scallop Placopecten, it does not; it is uniquely tied to the
allozymes, arguing clearly in this case for the direct-effect view.
Allozymes of small, albeit consistent, individual impact should certainly make
contributions, to phenotypic variances or to resulting fitness components, which
cannot, because of small size, be distinguished from truly “exchangeable” effects in
statistical analysis. If these are indeed the majority of selectively maintained
allozymes, they would show the ambiguities of multilocus frequency distributions on
which the neutralist-selectionist debate foundered. Such data would fit either to
neutral models or to diverse models of molecular evolution under positive selection,
as summarized by GILLESPIE (1991). Only test strategies which use biological speci-
ficity in one way or another, whether with single gene variants’ specific properties or
in multilocus comparison to other classes of variants, will be able to resolve these
possibilities.
ALLOZYMES AND APPROACHES TO NEUTRALITY
We saw that some Colias PGI variants are equivalent in function and fitness
components, hence neutral with respect to one another, even while they differ greatly
from, and are strongly selected relative to, other genotypes in the same polymorphism.
In E. coli, some allozymes of the same gene, PGI, do NOT differ detectably in function
at 37°C, and as one would therefore expect are the same in fitness evaluations at that
temperature, to a precision of +0.2% (DYKHUISEN & HARTL 1983). Two alleles of
phosphogluconate dehydrogenase in E. coli differ in kinetics by about 30%; they
moved from effective neutrality, on one genetic background in which their enzyme was
peripheral to the overall cellular energy budget, to strong selective difference when on a
second genetic background which made their enzyme central to the cellular energy
budget (DYKHUISEN & HARTL 1980, HARTL & DYKHUISEN 1981). This strongly
supported the use of bioenergetics as a context for evaluating evolutionary impact of
genetic variants (WATT 1985b). DEAN (1989) analyzed other bacterial enzymes in
relation to their placement in metabolic organization, showing which may be subject to
strong selection when variable, and which are more likely to show neutrality for the
same magnitude of allelic functional differences. Once again, biological specificity, far
from causing evolutionary study to bog down in details, is the key to sorting out
patterns of what actually causes neutrality OR selectedness. It appears that general-
izations are waiting to be found, but they must be searched for with mechanistic tools.
ALLOZYMES AS TOOLS FOR OTHER KINDS OF EVOLUTIONARY STUDY
ARE ALLOZYMES USEFUL AS “MARKERS”?
Diverse workers have used allozymes as “markers” for studies of paternity,
“gene flow”, and other aspects of population structure in the wild. What are the
implications for this practice of the above results?
878 WARD B. WATT
For proximate measurement of paternal identities in specific offspring cohorts,
and the like, neutrality or selectedness of the “marker” allozymes usually makes very
little difference to the measurement. For inference of population structure, on the
other hand, distinct or even opposed explanations of any given initial finding are often
possible, though others will very likely be ruled out.
For example, in a very careful and thoughtfully executed study, Costa & Ross
(1994) studied population structure in the eastern N. American moth Malacosoma.
There is only minor differentiation of allele frequencies, indicated by Wrightian F-
statistics calculated on 11 varying allozyme loci, at either 100 km deme separation
scale, or 200-1000 km scale. The authors comment that this shows either major
migrational gene exchange on the scale of hundreds of km, or uniformity of selection
regime over that same scale despite climate and biome differences, or some combi-
nation of the two. Further work, as they say, will be needed to resolve these alter-
natives. Other possibilities (local drift under restricted migration, or local selective
habitat specialization), which could have explained local differences if found, are
ruled out for these characters, so much has already been learned. In ecological/evolu-
tionary contrast, RANK (1992), studying a much less vagile leaf beetle, found sub-
stantial local allozyme frequency differentiation, even among adjacent drainages in
the same mountain range. Rank’s results could be due to local drift, local microhabitat
specialization, or a combination, subject to further study — though gene-specific varia-
tion in the extent of frequency differences led him to postulate local selection, at least
on the PGI gene. In each case, use of allozymes to probe population structure has
ruled out some possibilities, while leaving the need for further specific work to dis-
tinguish among other causal scenarios — as the authors have been careful to state.
These authors properly avoided the temptation to make “simplifying assumptions”
about either selectedness OR neutrality of allozyme probes of population structure;
such assumptions could thoroughly obscure the actual explanations of genetic patterns
seen.
HOW RELIABLE ARE ALLOZYMES AS SYSTEMATICS CHARACTERS?
There is a large literature on the use of allozymes, usually with calculation of
genetic distances (e.g. NEI 1987), as input to systematic analysis. For example, BURNS
(1975) found that the hypervariable Esterase-D gene in Colias (some populations
segregate 20 alleles or more) shows patterns of variation consistent with putative
species complexes among North American taxa. Some of this variation among com-
plexes seems to covary with larval food plant use, suggesting a cause of the pattern
which further study can test. GEIGER (1980), in an initial study of Pieridae (Colias’
family) as a whole, obtained genetic-distance data using 20 allozyme loci, which
provide the only independent support thus far for subgeneric distinctions within the
genus Colias, proposed later by BERGER (1986). But what would be the impact on
such work if, as seems likely, ordinary-resolution electrophoresis gels OVERestimate
the extent of allelic identity among relatives (COYNE 1976, WATT et al. 1995)? Or
what if, contrary to the assumptions of most genetic distance models (e.g. NEI 1987),
ALLOZYMES IN EVOLUTIONARY GENETICS 879
many allozymes are subject to at least moderate natural selection? The effects of such
deviations from “simple” assumptions are not clear. Caution is indicated, and other
types of molecular characters may often be more prudent choices for systematics
work.
THE FUTURE OF EVOLUTIONARY GENETICS
The task before us can be stated simply, though much effort will be needed to
accomplish it. It is to bring mechanistic biology fully into the Darwinian rubric, and
to put it to work asking and answering those questions about evolutionary genetics
and the course of evolutionary change or stasis which only mechanistic study can
approach: the three initial stages of Feder and Watt’s recursive evolution concept,
which complement the fourth stage where “population genetics” holds sway. Allo-
zyme variants can be major tools for this effort — major probes of organismal energy
budgets, putatively adaptive features of metabolic organization, etc. Some fear that
this strategy requires evolutionary geneticists to stretch their knowledge and efforts
too far. The success of diverse research groups, recorded here, at implementing
mechanistic strategies for studying evolution suggests these fears are needless. While
it is true that some further effort may be needed to master more diverse scientific con-
cepts and techniques, this will yield great returns. Individually and collaboratively,
evolutionary geneticists now have the opportunity to move beyond the remains of the
“neutralist-selectionist” controversy in search of new and more general under-
standing.
ACKNOWLEDGEMENTS
I thank Paul Ward and the Swiss Zoological Society for their kind invitation to
present these remarks, numerous members of the Society for stimulating discussions,
and Michael Dohm and Michael Kohane for helpful comments on the draft manus-
cript. I thank the U.S. Dept. of Energy (grant DE FG 03-93-ER-61667) and the U.S.
National Science Foundation (grants DEB 91-19411 and IBN 92-20583) for research
support.
REFERENCES
BARNES, P.T., B. HOLLAND, E V. COURREGES. 1989. Genotype-by-environment and epistatic
interactions in Drosophila melanogaster: the effects of Gpdh allozymes, genetic back-
ground, and rearing temperature on larval development time and stability. Genetics 122,
859-868.
BERGER, L. 1986. Systématique du genre Colias F. (Lepidoptera — Pieridae). Lambillionea 86
suppl., 1-68.
BuLsMma, R. & W. van DELDEN. 1977. Polymorphism at the G6PD and 6PGD loci in
Drosophila melanogaster. I. Evidence for selection in experimental populations. Genet.
Res. 30, 221-236.
880 WARD B. WATT
Burns, J.M. 1975. Isozymes in evolutionary systematics. In MARKERT, C.L., ed., /sozymes IV:
Genetics and Evolution, Academic Press, NY.
BURTON, R.S. & M.W. FELDMAN. 1983. Physiological effects of an allozyme polymorphism:
glumate pyruvate transaminase and response to hyperosmotic stress in the copepod
Tigriopus californicus. Biochem. Genet. 21, 239-251.
CAVENER, D.R. & M.T. CLEGG. 1981. Evidence for biochemical and physiological differences
between enzyme genotypes in Drosophila melanogaster. Proc. Natl. Acad. Sci. USA
78, 4444-4447.
Costa, J.T., & K.G. Ross. 1994. Hierarchical genetic structure and gene flow in macrogeo-
graphic populations of the eastern tent caterpillar (Malacosoma americanum). Evo-
lution 48: 1158-1167.
Coyne, J.A. 1976. Lack of genic similarity between two sibling species of Drosophila as
revealed by varied techniques. Genetics 84, 593-607.
DEAN, A.M. 1989. Selection and neutrality in lactose operons of Escherichia coli. Genetics
123, 441-454.
DYKHUISEN, D.E. & D.L. HARTL. 1980. Selective neutrality of 6PGD allozymes in E. coli and
the effects of genetic background. Genetics 96, 801-817.
DYKHUISEN, D.E., & D.L. HARTL. 1983. Functional effects of PGI allozymes in E. coli. Gene-
tics 105, 1-18.
EANES, W.F., M. KIRCHNER, & J. YOON. 1993. Evidence for adaptive evolution of the G6pd
gene in the Drosophila melanogaster and Drosophila simulans lineages. Proc. Natl.
Acad. Sci. USA 90, 7475-7479.
ENDLER, J.A. 1986. Natural selection in the wild. Princeton Univ. Press, Princeton, NJ.
EWENS, W.J., & M. W. FELDMAN. 1976. The theoretical assessment of selective neutrality. In
KARLIN, S., & E. NEvo, eds., Population genetics and ecology, Academic Press, NY, pp.
303-337.
FALCONER, D.S. 1989. Introduction to quantitative genetics, 3rd. ed. Longmans, London, UK.
FEDER, M.E., & W.B. WATT. 1992. Functional biology of adaptation. In BERRY, R.J., R.J.
CRAWFORD & G.M. HEWITT, eds., Genes in ecology, Blackwell’ s, London, UK, pp. 365-
392!
FISHER, R.A. 1930. The genetical theory of natural selection. Clarendon Press, Oxford, UK.
GEIGER, H.J. 1980. Enzyme electrophoretic studies on the genetic relationships of pierid
butterflies (Lepidoptera, Pieridae). I. European taxa. J. Res. Lepid. 19, 181-195.
GILLESPIE, J. H. 1991. The causes of molecular evolution. Oxford Univ. Press., Oxford, UK.
GILLESPIE, J. H. 1994. Substitution processes in molecular evolution, II. Exchangeable models
from population genetics. Evolution 48, 1101-1113.
HALDANE, J.B.S. 1932. The causes of evolution. Longmans, Green, London, UK.
HALDANE, J.B.S. 1954. The biochemistry of genetics. Allen & Unwin, London, UK.
HALDANE, J.B. S., & S.D. JAYAKAR. 1963. Polymorphism due to selection of varying direction.
J. Genet., 58, 237-242.
HARTL, D.L., & D.E. DYKHUISEN. 1981. Potential for selection among nearly neutral allozymes
of 6-phosphogluconate dehydrogenase in Escherichia coli. Proc. Nat I. Acad. Sci. USA
78, 6344-6348.
Hussy, J.L., & R.C. LEWONTIN. 1966. A molecular approach to the study of genic heterozygo-
sity in natural populations. I. The number of alleles at different loci in Drosophila
pseudoobscura. Genetics 54, 577-594.
INGRAM, W.I. 1962. The hemoglobins in genetics and evolution. Columbia Univ. Press, NY.
JEPSEN, G.L., G.G. SIMPSON, & E. MAYR, eds. 1949. Genetics, palaeontology, and evolution.
Princeton Univ. Press, Princeton, NJ.
ALLOZYMES IN EVOLUTIONARY GENETICS 881
KARL, S., & J.C. AVISE. 1992. Balancing selection at allozyme loci in oysters: implications
from nuclear RFLPs. Science 256, 100-102.
KELLER, L., & K.G. Ross. 1993. Phenotypic basis of reproductive success in a social insect:
genetic and social determinants. Science 260, 1107-1110.
Kimura, M. 1983. The neutral theory of molecular evolution. Cambridge Univ. Press,
Cambridge, UK.
KoEHN, R.K, & S. R. SHumway. 1982. A genetic/physiological explanation for differential
growth rate among individuals of the American oyster Crassostrea virginica (Gmelin).
Mar. Biol. Letters 3, 35-42.
KUHN, T.S. 1970. The structure of scientific revolutions, 2nd ed. Univ. of Chicago Press,
Chicago IL.
LABATE, J., & W.F. EANES. 1992. Direct measurement of in vivo flux differences between
electrophoretic variants of G6PD from Drosophila melanogaster. Genetics 132, 783-
MEN:
Lar, C., R.F. LYMAN, A.D. LONG, C.H. LANGLEY, & T.F.C. MACKAY. 1994. Naturally occurring
variation in bristle number and DNA polymorphisms at the scabrous locus of
Drosophila melanogaster. Science 266, 1697-1702.
LEVENE, H. 1953. Genetic equilibrium when more than one niche is available. American
Naturalist 87, 331-333.
LEWONTIN, R.C., & L.C. DUNN. 1960. The evolutionary dynamics of a polymorphism in the
house mouse. Genetics, 45, 705-722.
MARKERT, C.L., J.B. SHAKLEE, & G.S. WHITT. 1975. Evolution of a gene. Science 189, 102-
114.
Mayr, E. 1958. Change of genetic environment and evolution. In HUXLEY, J., A.C. HARDY, &
E.B. FORD, eds., Evolution as a process, 2nd. ed., Allen & Unwin, London, pp. 188-213.
NEI, M. 1987. Molecular evolutionary genetics. Columbia University Press, NY.
Orr, H.A., & J.A. COYNE. 1992. The genetics of adaptation: a reassessment. American Natu-
ralist 140, 725-742.
PAULING, L., H.A. ITANO, S.J. SINGER, & I.C. WELLS. 1949. Sickle-cell anemia, a molecular
disease. Science 110, 543.
Pocson, G.H. 1991. Expression of overdominance for specific activity at the phospho-
glucomutase-2 locus in the Pacific oyster, Crassostrea gigas. Genetics 128, 133-141.
PoGson, G.H., & E. Zouros. 1994. Allozyme and RFLP heterozygosities as correlates of
growth rate in the scallop Placopecten magellanicus: a test of the associative
overdominance hypothesis. Genetics 137, 221-231.
Powers, D.A., T. LAUERMAN, D. CRAWFORD, & L. DIMICHELE. 1991. Genetic mechanisms for
adapting to a changing environment. Ann. Rev. Genet. 25: 629-659.
PROVINE, W.B. 1971. The origins of theoretical population genetics. University of Chicago
Press, Chicago IL.
RANK, N. 1992. A hierarchical analysis of genetic differentiation in a montane leaf beetle
Chrysomela aeneicollis (Coleoptera: Chrysomelidae). Evolution 46, 1097-1111.
REAL, L. A., ed. 1994. Ecological genetics. Princeton Univ. Press, Princeton, NJ.
ROTHMAN, E.D., & A.R. TEMPLETON. 1980. A class of models of selectively neutral alleles.
Theor. Pop. Biol. 18, 135-150.
SKIBINSKI, D.O.F., & R.D. WARD. 1982. Correlations between heterozygosity and evolutionary
rate of proteins. Nature 298, 490-492.
SKIBINSKI, D.O.F., M. WOODWARD, & R.D. WARD. 1993. A quantitative test of the neutral
theory using pooled allozyme data. Genetics 135, 233-248.
STERN, C. 1949. Gene and character. In JEPSEN et al., eds., op. cit., pp. 13-22.
882 WARD B. WATT
SAVOLAINEN, O., & P. HEDRICK. 1995. Heterozygosity and fitness: no association in Scots pine.
Genetics 140, 755-766.
TEMPLETON, A.R. 1982. Adaptation and the integration of Soßen forces. In R. MILKMAN
(ed.), Perspectives on evolution, Sinauer, Sunderland, MA, pp. 15-31.
TRAVIS, J. 1994. Ecological genetics of life-history traits: variation and its evolutionary
significance. /n REAL, L.A., ed., op. cit., pp. 171-204.
TURNER, J.R.G. 1992. Stochastic processes in populations: the horse behind the cart? In BERRY,
R.J., T.J. CRAWFORD & G.M. HEWITT, eds., Genes in ecology, Blackwell’s, London,
UK, pp. 29-53.
VIA, S. 1994. Population structure and local adaptation in a clonal herbivore, pp. 58-85 in
REAL, L.A., ed., 1994, op. cit.
WATT, W.B. 1983. Adaptation at specific loci II. Demographic and biochemical elements in the
maintenance of the Colias PGI polymorphism. Genetics 103, 691-724.
WATT, W.B., R. C. CASSIN & M. S. Swan. 1983. Adaptation at specific loci III. Field behavior
and survivorship differences among Colias PGI genotypes are predictable from in vitro
biochemistry. Genetics 103, 725-739.
WATT, W.B. 1985a. Allelic isozymes and the mechanistic study of evolution. /sozymes:
Current Topics in Biological and Medical Research 12, 89-132.
WATT, W.B. 1985b. Bioenergetics and evolutionary genetics — opportunities for new synthesis.
American Naturalist 125, 118-143.
WATT, W.B. 1992. Eggs, enzymes, and evolution — natural genetic variants change insect
fecundity. Proc. Nat'l. Acad. Sci. USA 89, 10608-10612.
WATT, W.B. 1994. Allozymes in evolutionary genetics: self-imposed burden or extraordinary
tool? Genetics 136, 11-16.
WATT, W.B., K. DONOHUE, & P.A. CARTER. 1995. Adaptation at specific loci. VI. Divergence
vs. parallelism of polymorphic allozymes in molecular function and fitness-component
effects among Colias species (Lepidoptera, Pieridae). MS in review.
WHITT, G.S. 1987. Species differences in isozyme tissue patterns: their utility for systematic
and evolutionary analyses. /sozymes: Current Topics in Biological and Medical
Research 15, 1-26.
WRIGHT, S. 1931. Evolution in Mendelian populations. Genetics 16, 97-159.
WRIGHT, S. 1934. Physiological and evolutionary theories of dominance. American Naturalist
34, 24-53.
REVUE SUISSE DE ZOOLOGIE, 102 (4) : 883-894: décembre 1995
What causes diversity in male sexual characters?”
Andrew POMIANKOWSKI** & Yoh IWASA***
** Department of Genetics and Biometry, University College London,
4 Stephenson Way, London NWI 2HE, UK.
*** Department of Biology, Faculty of Science, Kyushu University,
Fukuoka 812, Japan.
What causes diversity in male sexual characters? - Closely related
species often differ markedly in sexual morphology and courtship display.
Here we show that Fisher’s runaway process of sexual selection is a good
explanation for this diversity. The runaway process is intrinsically unstable
if selection on the male trait is weak around the natural selection optimism
but increases rapidly beyond a certain limit. Runaway can often result in
continuous change in sexual characters.
Key-words: Sex - Speciation - Courtship - Runaway - Mate choice - Sexual
selection.
INTRODUCTION
Secondary sexual characters are often highly variable between closely related
species (DARWIN 1871; ANDERSSON 1994). This diversity contrasts with morpho-
logical traits that show far less variability over the same taxonomic range. There are
few studies of the comparative rates of change of sexual and non-sexual traits but
there are some notorious cases: the species flocks of the Great Lakes in Africa (FRYER
& ILES 1980), the Hawaiian Drosophila (CARSON 1986), male insect genitalia
(EBERHARD 1985) and the birds of paradise (COATES 1990). These examples suggest
that sexual traits are highly labile and can change very guickly in evolutionary time.
Variation is even seen between sub-populations. A good detailed example is an
isolated, sub-population of sage grouse, from Gunnison, Colarado (YOUNG et al.
1994). The Gunnison birds are smaller than those in the main range from the west
coast to central USA. This has not lead to any apparent ecological divergence (Young
pers. comm.). However, male birds differ markedly in their secondary sexual charac-
ters. Certain elements of the male display are identical - for example strut and song
components. Others have quantitatively diverged - Gunnison males strut at a lower
rate, their tail feather plumes are whiter and their rate of air sac inflation is higher.
Two novel elements are also present - Gunnison males wag their tails from side to
* Main lecture presented at Zoologia 95.
884 ANDREW POMIANKOWSKI & YOH IWASA
side and, more dramatically, toss their neck plumes over their heads. Neither of these
behaviours is seen in sage grouse from the main range (YOUNG er al. 1994). Preli-
minary call playback experiments show concomitant divergence in female preference;
Gunnison females prefer calls of Gunnison males than those made by males from the
main range (Young, pers. comm.).
How has sexual selection lead to the evolution of diverse sexual characters?
Why do some sexual characters remain constant, others show minor quantitative
change, whilst other sexual traits appear to be novel elements? A traditional answer is
that diversity arises to maintain the identity of newly diverged populations or species
(PATERSON, 1985). There is scant theoretical or empirical support that this occurs
through reinforvement to stop hybridization (BUTLIN 1987), though a recent re-
analysis suggests a greater role for reinforcement (HOWARD 1993). Species recogni-
tion cannot be a general explanation as it does not explain divergence in allopatric
populations (eg, sage grouse). High divergence of sexual traits also occurs in island
species where encounters between populations or species are minimal (EBERHARD
1985). But present distributions may hide past associations. We do not rule out a role
for species identity but will not pursue this matter further here.
Another general approach has stressed the importance of random genetic drift
in triggering sexual selection in newly founded populations (TEMPLETON 1980). In the
1980s this view was reinforced by models of Fisher’s runaway process of sexual
selection. These showed that runaway led to an infinite set of outcomes, the so-called
‘line of equilibria’ (LANDE 1981). Divergence from one equilibrium causes runaway
evolution to a novel endpoint. This suggests that allopatric populations will quickly
diverge through the exaggeration of random differences. The problem with this
interpretation is that several studies have shown that ‘lines of equilibria’ collapse to
single points when more realistic model assumptions are made (POMIANKOWSKI 1987;
BULMER 1989; POMIANKOWSKI ef al. 1991). Under these conditions drift no longer
plays a significant role in initiating divergence (see below). As it stands, current
models of sexual selection are inadequate to explain diversity.
In this paper we re-examine the basic assumptions behind models of Fisher’s
runaway process. We show that Fisher’s runaway process is intrinsically unstable and
naturally leads to continual change in sexual traits (IWASA & POMIANKOWSKI 1995).
Female preferences for exaggerated male traits are only semi-stable. Strong preference
slowly decays until runaway is triggered again in a different direction. The only change
we make to previous models is to assume male survival decreases dramatically once
male sexual traits become extreme. This is a general and plausible assumption. It is akin
to Fisher’s (1930) original description of the costs of male sexual traits.
DRIFT AND ENVIRONMENTAL DIFFERENCES
First we investigate the effect of drift and environmental differences in allo-
patric populations, between which there is no gene flow. Using a simple sexual
selection model of Fisher’s runaway we show to what degree divergence is expected.
DIVERSITY IN MALE SEXUAL CHARACTERS 885
Let t be a male trait used by females in mate choice and p be the strength of
female preference. We assume that both ¢ and p are sex-limited and have a polygenic
basis. The standard model of runaway follows the change per generation of the mean
values.
AF\ 1 (GB B, ish N
ee BEIGE IR, 0)’
where G, and G, are additive genetic variances of ¢ and p respectively, B is the addi-
tive genetic covariance between these two traits and u measures mutation bias acting
on the male trait (POMIANKOWSKI et al. 1991; IwAsa er al. 1991). The additive genetic
covariance is responsible for the coevolution of the two traits. The covariance arises
through non-random mating; females with stronger preference (larger p) tend to mate
with males with larger sexual traits (larger r). Given a positive covariance, selection
for larger male traits causes a correlated increase in female preference.
The effect of selection is given by the selection gradients ß, and B,, for the
male trait and female preference respectively. Male fitness is the product of mating
success and survivorship. Mating success is determined by female mate preference. If
the mean female preference is positive p>0, female preference favours males with
larger than average trait values (POMIANKOWSKI et al. 1991). The reverse is true if
p<0, which favours males with smaller than average trait values. When p=0 females
exert no selective pressure on males, as they mate at random. The second component
of male fitness is survivorship. The optimal expression of the male trait under natural
selection is set at ft = 0, with survival chances declining for larger or smaller values.
The coefficients a and c calibrate these two selective effects. Female fitness is deter-
mined by the strength of preference. We assume that female costs are minimised
when there is no discrimination (p = 0) and increase symmetrically with the strength
of preference at a rate b.
The following equations describe the standard mathematical interpretation of
these assumptions (LANDE 1981; POMIANKOWSKI et al. 1991), selection differentials
being evaluated at the population means,
B, = La = 2 (ap(1-1) = ct”) = OD DAG,
ot ot
à 3) 5 (2)
B, = —InW,; = —(-bp?) = -2bp,
op op
where W,, and Wy are the lifetime reproductive success of males and females. The
equations can be substituted into Eg(1) and solved for equilibrium (At = 0, Ap = 0).
We simplify the results by noting that the additive genetic covariance between the
male trait and female preference converges quickly to B = aG, Gp / 2 (POMIANKOWSKI
& IwasA 1993; for a full discussion of the model assumptions see Iwasa et al. 1991).
A single equilibrium exists,
886 ANDREW POMIANKOWSKI & YOH IWASA
a
PE 2b° (3a)
= An:
32,08 (3b)
This is a simple result. Equilibrium is given by the benefits and costs of mate choice.
Female benefits are given by the mutation parameter u which determines the advan-
tage of mating with males bearing a larger trait, multiplied by a the effectiveness of
preference in generating non-random mating. Female choice costs are given by b. The
male trait equilibrium follows the strength of female preference factored by the
benefit to cost ratio of the male trait, a/c.
We can now see that drift has no effect on equilibrium. If this equilibrium is
stable it is globally stable (POMIANKOWSKI, ef al. 1991). All deviations caused by drift
always lead back to the same equilibrium point. (If the equilibrium is unstable then
perpetual runaway occurs, an unrealistic model outcome). So we do not expect drift to
be a major cause of divergence in secondary sexual traits. Drift is only important if b,
u=0 and pp can trake any value along a line of equilibrium given by Eq(3b), as was
originally pointed out by LANDE (1981). Such a requirement is highly restrictive and
not likely to be met in nature.
We can also see the effect of environmental change. This will affect the three
parameters a, b and c. Let us suppose that the environments of two allopatric
populations differ in predation pressure and this alters the cost of the male trait. This
will increase the value of c and reduce the equilibrium exaggeration of the male trait.
If predation also affects the cost of female choice b, then both preference and the male
trait will be reduced. The parameter a is harder to interpret as it is a function of both
male and female behaviour and the environment. From the female’s view point a
reflects the efficiency of preference in identifying males with larger ornaments. From
the male’s view point a reflects the effectiveness of male signalling in causing
increased mating success. a also varies with life history and ecology. For instance,
lower values of a will be common in monogarnous species because there are fewer
mating opportunities compared to polygamous species. Another cause of reduced
values of a could be a densely wooded environment which restricts the efficacy of
visual or auditory signals compared to more open habitats. If two allopatric popu-
lations differ in any of these properties this will cause differences in secondary sexual
characters.
Environmental differences predict divergence in sexual traits. If changes in the
environment are small, divergence in sexual traits will not be dramatic or that much
greater than in non-sexually selected traits, though larger divergence is possible be-
cause sexual selection has an amplifying effect (LANDE 1982). Environmental change
can quantitatively shift the equilibrium values. But it is not obvious how this relates to
the kind of divergence seen in sage grouse populations discussed above. The magni-
tude of these effects requires a different explanation.
DIVERSITY IN MALE SEXUAL CHARACTERS 887
A NEW FISHERIAN MODEL
An assumption common to previous models is that male survival chances
decrease with the quadratic power (-cr2 in Eq(2a)). This follows LANDE’s (1981) original
analysis which assumed normality of the fitness functions. However this need not be the
case. We now examine a more extreme fitness function that fits better with FISHER’s
(1930) original assumption. Fisher supposed that stabilizing selection on the male trait
was weak around the optimum, thereby permitting runaway to get started. But once the
male trait exceeded a certain limit the cost of further exaggeration increased very
quickly. This behaviour can be captured by making fitness decline with the fourth power
rather than the quadratic (IWASA & POMIANKOWSKI 1995). Rewriting Eq(2a).
B = Z(aple-7) - cr) = ap - 4er? (4)
ot
Evolutionary change can be followed by substituting this equation and Eq(2b)
for female fitness into Eq(1). The result is cyclic evolution (Fig. 1). A population
starting near the origin evolves away in a runaway process. The runaway is brought to
a halt once selection on the male trait becomes extreme. The system converges to a
curve, on which evolutionary change slows down. Here the male trait is exaggerated
and females show corresponding strong preference. Female preference then slowly
declines. This decline continues until runaway is triggered again but this time in a
negative direction. The negative runaway stops when selection on the male trait
becomes extreme. At this point both traits are negatively exaggerated. Then a very
slow movement in a positive direction occurs towards p=0 due to the cost of choice.
Eventually runaway starts again toward positive values and the whole process repeats
itself.
The underlying reasons for this evolutionary behaviour are easy to understand.
We assume that the costs to female preference b and mutation bias u are very much
smaller than the other parameters (IWASA et al. 1991). This seems a reasonable assump-
tion as it has proved difficult to measure b and u compared to other parameters (ie, b, u
<< d, C, G;, Gp). So to follow the gross behaviour of the model we can initially neglect
P, and u. This leaves selection on the male triat B, which causes runaway. During
runaway the male trait and female preference evolve together as Ap /Af=B/G,=aGy
/ 2. Runaway ceases when Ap = At = 0 which yields a curve of the equilibria which is
flat around the origin but steeply increases once the male trait becomes exaggerated
(Fig. 2). Part of the curve is stable (solid) and part unstable (dashed). Stability depends
on the slope of the evolutionary trajectory aG, / 2 being less than the slope of the curve
of equilibria.
Once runaway ceases near the curve of equilibria, further evolution is deter-
mined by the costs to female preference b and mutation bias u. First we discuss the case
in which mutation bias is negligibly small (4 = 0), which is likely to be the case for
many sexual traits. After runaway, female preference simply decays towards p = 0
because of the cost of choice. The population never reaches the origin because a
888
ANDREW POMIANKOWSKI & YOH IWASA
a
b
D Aube ;
t 3 i
O2 " ci . ry
2000 * 4000 = 6000 8000
0.2 : ; .
Generation
-0.4 s =
u
Fic. 1
Cyclic evolution caused by Fisher’s runaway process is plotted as a) the joint evolution of the
mean female preference (p) and mean male trait (7) and b) as change in the mean male trait (?)
per generation. The population is plotted every 20 generations. Parameter values are a=0.4,
b=0,001, c=0.05, G,=0.5 G,=0.5, u=0.
DIVERSITY IN MALE SEXUAL CHARACTERS 889
FIG. 2
The curve of equilibria, showing stable (solid) and unstable (broken) regions.
second runaway is initiated once the population reaches the unstable part of the curve
(Fig. 1). The speed of movement around the cycle is determined by the time the
population remains on the semistable arc of the curve of equilibria (runaway in
comparison is very rapid) which is mainly determined by b. Thus cycle time increases
if the cost of choice b is small.
If mutation bias is negligibly small, there is an unstable equilibrium at the
origin. If there is significant mutation bias u>0, the equilibrium lies at,
ua
a (5a)
ST
a 1/3
ie=(£ pr) | (5b)
C
If pg lies in the unstable region of the curve, the population evolves in a cycle as with
u=0. But if mutation bias is strong enough pp lies in the stable region of the curve and
890 ANDREW POMIANKOWSKI & YOH IWASA
the population smoothly converges on the equilibrium which is globally stable (Fig. 3).
Stability is more likely if mutation bias (u) is large and the cost of choice (b) is small.
This model helps to understand the evolution of diversity in sexually selected
characters. It suggests that perpetual change in sexual characters is a simple outcome
of Fisher’ s runaway process. Runaway exaggeration of a particular male trait is only
semi-stable. In our model this leads to cycles, but this result reflects the model’s
constraint to a single dimension. If we allow multiple preferences for different male
traits (eg, song, colour, tail length), the population evolves in a more complex manner
and only cycles over a very long period (IWASA & POMIANKOWSKI, unpub). This
means that slight ecological differences between allopatric populations quickly cause
them to fall out of phase and evolve different sexual phenotypes.
The model also helps us to understand why some sexual traits remain
unchanged. Traits which are subject to large mutation bias (u) will tend to be stable as
will those which are cheap for females to choose (b). This result is of interest to the
debate over the importance of Fisher’s runaway and the handicap process. Mutation
bias is a simple measure of how mutation or environmental change cause a
deterioration in mean trait values. We have argued that Fisher traits (sexual traits that
do not act as indicators of good genes) can be subject to mutation bias when they are
exaggerated or elaborate because most mutations will tend to reduce trait values
(POMIANKOWSKI et al. 1991). But this effect is likely to be small. In contrast sexual
traits that are handicaps are predicted to show large mutation bias because most mu-
tations are known to reduce viability (MUKAI & COCKERHAM 1977; CHARLESWORTH
1987). This leads to the prediction that sexual traits common to members of a clade
are more likely to be handicaps that indicate good genes. Whereas sexual traits which
vary greatly within a clade are more likely to be Fisher traits.
DISCUSSION
Fisher’s runaway model of sexual selection has had a chequered history. In the
reemergence of interest in handicap models there has been a tendency to overlook its
importance (eg, GRAFEN 1990). Others have suggested that runaway is unlikely
because additive genetic variance/covariance in preference and sexual traits are non-
existent or not maintained in the face of strong selection (TAYLOR & WILLIAMS 1982,
KIRKPATRICK & RYAN 1991). But empirical surveys suggest this pessimism is un-
founded. Both traits have been shown to be genetically variable and genetic cova-
riances have now been demonstrated in a number of species (BAKKER &
POMIANKOWSKI 1995; POMIANKOWSKI & MOLLER 1995).
In this paper we have considered how Fisher’s runaway may contribute to the
diversity of sexual traits. Our model shows that runaway is intrinsically unstable and
naturally leads to continual change in sexual traits (IWASA & POMIANKOWSKI 1995).
This result emerges from a simple and non-restrictive change to the standard model
that makes it more consistent with FISHER’s (1930) view. It is assumed that stabilising
selection on the male trait is weak around the natural selection optimum but beyond a
DIVERSITY IN MALE SEXUAL CHARACTERSYAENAS 89]
2000 4000 6000 8000
Generation
Fic. 3
Stable evolution caused by Fisher’s runaway process. In this example mutation bias on the male
trait u is large enough and the cost of choice b small enough for the equilibrium point to lie on
the stable arc of the curve of equilibria. Parameter values are a=0.4, b=0.001, c=0.05, G,=0.5,
Gp=0.5, u=0.001.
892 ANDREW POMIANKOWSKI & YOH IWASA
certain value it increases rapidly. The implication to draw from this result is that traits
exaggerated by runaway will change rapidly in time and quickly diverge in allopatric
populations.
It is possible for runaway to cause the stable exaggeration of sexual characters.
This occurs if there is strong mutation bias and weak choice costs. These conditions
are restrictive. Mutation bias on Fisher traits, which only signal attraction to females,
is unlikely to be very strong - though this has yet to measured. It is more likely that
these conditions are fulfilled by sexual traits acting as indicators of good genes.
Other explanations of the diversity of sexually selected traits have stressed the
importance of random genetic drift about a line of equilibrium (LANDE 1981). We can
rule out the importance of drift in this context because its effect will be dominated by
other weak forces with a directional component, especially choice costs. These reduce
lines of equilibria to single endpoints that are globally stable. At a more general level
we can not rule a role for drift as our model is deterministic. But drift’s general
importance is unproven.
Another explanation has focused on distinct selective pressures in geogra-
phically separated populations as a cause of diversity (LANDE 1982). This is entirely
likely as the exaggeration of sexual traits is a balance between the strength of female
preference and natural selection on the male trait. ENDLER (1980) has shown that this
balance typifies Trinidadian guppy coloration, different populations’ colour patterns
varying with natural selection caused by predation pressure. Recently it has been
established that female preference in the guppy is also constrained by predation pres-
sure (GODIN & BRIGGS 1995). This may explain quantitative differences in the
expression of sexual traits but is not sufficient to explain the extensive diversity of
sexual traits seen in closely related species.
ACKNOWLEDGEMENTS
This research was supported by the Royal Society (AP) the Japanese Ministry
of Education, Science and Culture (YI) and the NERC Centre for Population Biology,
Imperial College at Silwood Park (YI). We are grateful for this support and also thank
Hiroyuki Matsuda, Mark Pagel and Akira Sasaki for their helpful comments. Thanks
to Jessica Young for providing unpublished information about sage grouse. This
paper was originally given as a talk at the Swiss Zoological Society’s meeting in
1995, and I extend my warm thanks to Paul Ward for his encouragement.
REFERENCES
ANDERSSON, M. 1994. Sexual Selection. Princeton University Press, Princeton, New Jersey.
BAKKER, T.C.M. & A. POMIANKOWSKI, 1995. The genetics of female mate preferences. J. Evol.
Biol. 8: 129-171.
BULMER, M. 1989. The structural instability of sexual selection models. Theoret. Pop. Biol. 36:
195-206.
DIVERSITY IN MALE SEXUAL CHARACTERS 893
BUTLIN, R.K. 1987. Speciation by reinforcement. Trends Ecol. Evol. 2: 8-13.
CARSON, H.L. 1986. Sexual selection and speciation. In S. KARLIN & E NEVO (eds). Evolu-
tionary Processes and Theory, pp. 391-409. Academic Press, Orlando.
CHARLESWORTH, B. 1987. The heritability of fitness. In J. W. BRADBURY & M.B. ANDERSSON
(eds) Sexual Selection: Testing the Alternatives, pp. 21-40. Wiley, Chichester.
Coates, B. 1990. The Birds of Papua New Guinea. Volume II Passerines. Dove, Alderley,
Queensland.
Darwin, C.R. 1871. The Descent of Man and Selection in Relation to Sex. John Murray,
London.
EBERHARD, W.G. 1985. Sexual Selection and Animal Genitalia. Havard University Press,
Cambridge, Mass.
ENDLER, J.A. 1980. Natural selection on color patterns in Poecilia reticulata. Evolution 34: 76-
IE
Gopin, J.-G. J. & S.E. BRIGGs. Female mate choice under predation risk in the guppy. Anim.
Behav. In press.
GRAFEN, A. 1990. Biological signals as handicaps. J. Theoret. Biol. 144: 517-546.
FISHER, R.A. 1930. The Genetical Theory of Natural Selection. Clarendon Press, Oxford.
FRYER, G. & T.D. ILES. 1980. The Clichlid Fishes of the Great Lakes of Africa. Oliver & Boyd,
Edinburgh.
HOWARD, D.J. 1993. Reinforcement: the origin, dynamics and fate of an evolutionary hypo-
thesis. In HARRISON, R.G. (ed.) Hybrid Zones and the Evolutionary Process, pp. 118-
142. Oxford University Press.
Iwasa, Y. & A. POMIANKOWSKI.1995. Continual change in mate preferences. Nature 377: 420-
422.
Iwasa, Y., A. POMIANKOWSKI & S. NEE. 1991. The evolution of costly mate preference II. The
‘handicap’ principle. Evolution 45: 1431-1442.
KIRKPATRICK, M. & M.J. RYAN 1991. The evolution of mating preferences and the paradox of
the lek. Nature 350: 33-38.
LANDE, R. 1981. Models of speciation by sexual selection on polygenic traits. Proc. Natl. Acad.
Sci. USA 78: 3721-3725.
LANDE, R. 1982. Rapid origin of sexual isolation and character divergence in a cline. Evolution
36: 213-223.
MUKAI, T. & C.C. COCKERHAM 1977. Spontaneous mutation rates at enzyme loci in Drosophila
melanogaster. Proc. Natl. Acad. Sci. USA 74: 2514-2517.
PATERSON, H.E.H. 1985. The recognition concept of species. In E.S. VRBA (ed.) Species and
Speciation pp. 21-29. Transvaal Museum, Pretoria, South-Africa.
POMIANKOWSKI, A. 1987. The costs of choice in sexual selection. J. theoret. Biol. 128: 195-218.
POMIANKOWSKI, A. 1988. The evolution of female mate preferences for male genetic quality.
Oxford Surv. Evol. Biol. 5: 136-184.
POMIANKOWSKI, A. & Y. IWASA. 1993. Evolution of multiple sexual ornaments by Fisher’s
process of sexual selection. Proc. Roy. Soc. Lond. B 253: 173-181.
POMIANKOWSKI, A. Y. IWASA & S. NEE. 1991. The evolution of costly mate preference I. Fisher
and the biased mutation. Evolution 45: 1422-1430.
POMIANKOWSKI, A. & A.P. MOLLER. 1995. A resolution of the lek paradox. Proc. Roy. Soc.
Lond. B 260: 21-29.
TAYLOR, P.D. & G.C. WILLIAMS. 1982. The lek paradox is not resolved. Theor. Pop. Biol. 22:
392-409.
TEMPLETON, A.R. 1980. The theory of speciation via the founder principle. Genetics 94: 1011-
1038.
Young, J.R., J.W. Hupp, J.W. BRADBURY & C.E. BRAUN. 1994. Phenotypic divergence of
secondary sexual traits among sage grouse, Centrocercus urophasianus, populations.
Anim. Behav. 47: 1353-1362.
REVUE SUISSE DE ZOOLOGIE, 102 (4) : 895-906; décembre 1995
Virilized sexual genitalia as adaptations of female spotted hyaenas*
Heribert HOFER & Marion L. EAST
Max-Planck-Institut für Verhaltensphysiologie,
D-82319 Seewiesen Post Starnberg, Germany.
Virilized sexual genitalia as adaptations of female spotted hyaenas. -
The study of traits as adaptations is reviewed using virilized female sexual
genitalia of spotted hyaenas as an example. Female spotted hyaenas lack a
vagina, have completely fused outer labiae and a clitoris (‘pseudo-penis’)
that closely mirrors the male penis in size and erectability. The evolu-
tionary origin and adaptive value of these structures have eluded scientists
for centuries. The example of virilized genitalia of female spotted hyaenas
demonstrates that questions about the evolutionary origin of a trait benefit
from adopting non-adaptive hypotheses: the virilized genitalia are most
likely a non-adaptive by-product of selection pressures related to the evo-
lution of siblicide. Field studies suggest that virilized genitalia have fitness
costs in the context of parturition and convey benefits in communication
(greeting ceremonies) and mating. The virilized genitalia of spotted
hyaenas highlight the need to pay attention to three aspects in the study of
adaptive processes. First, it is important to study current use and fitness
consequences in the field. Second, an important empirical question is
whether an observed trait is evolutionarily simple or complex in the sense
that the trait may be composed of a number of sub-traits subjected to
independent selection events. In this context comparative studies may be
useful. Third, the evolutionary history of a trait may be strongly influenced
by interactions with other traits.
INTRODUCTION
The study of adaptation is one of the most fundamental aspects of the study of
evolution (WILLIAMS 1966). Adaptation is the process by which the adaptedness of an
organism (its ability to live and reproduce in a given set of environments) improves
over evolutionary time by natural selection (DOBZHANSKY 1968, ENDLER 1986). The
term adaptation has also been used to describe the end-product of this process.
Studying the process of adaptation in the most comprehensive sense requires the
* Main lecture presented at Zoologia 95.
896 HERIBERT HOFER & MARION L. EAST
investigation of the recursive interaction of allele frequencies, the morphological or
physiological design of a trait, its performance, the fitness consequences of per-
formance, and the genetic response to selection in the next generation (FEDER &
WATT 1992). Studies with more modest aims focus on the adaptive value of a trait or
its evolutionary origins. Whatever the purpose, the study of adaptation faces a number
of difficult issues. The purpose of this contribution is to highlight some of these
difficulties and discuss them using the virilized genitalia of female spotted hyaenas as
an example.
Female spotted hyaenas have a pseudo-‘penis’ formed by the clitoris which
closely resembles the male penis in shape and size (FRANK et al. 1990) and can be
erected (MATTHEWS 1939, NEAVES et al. 1980, FRANK ef al. 1990). There is no vulva,
the labiae are fused and the urogenital tract passes through the clitoris. In other words,
female spotted hyaenas urinate, copulate and give birth through their pseudo-‘penis’.
The penis and pseudo-‘penis’ play a special role during greeting ceremonies for both
sexes. Greetings typically involve two hyaenas standing parallel, facing in opposite
directions. The hind leg nearest the animal being greeted is usually raised while each
animal sniffs or licks the erect ‘penis’ (WICKLER 1964, KRUUK 1972, MILLS 1990).
Early ideas were unanimous in that spotted hyaenas were considered to be
hermaphrodites (Table 1). More recent debates about the evolution, ontogenetic
development and function of the pseudo-‘penis’ of spotted hyaenas, on both proximate
and ultimate levels, has centered around the role of the ‘penis’ in greeting ceremonies
(WICKLER 1964, KRUUK 1972, RACEY & SKINNER 1979, GOULD 1981, GOULD & VRBA
1982, LINDEQUE & SKINNER 1982, HAMILTON et al. 1986, LINDEQUE et al. 1986, VAN
JAARSVELD & SKINNER 1987, SHERMAN 1988, FRANK et al. 1991, EAST et al. 1993).
TABLE |
Early ideas about the pseudo-penis of spotted hyaenas.
Idea Source
Spotted hyaenas are simultaneous hermaphrodites Aristotle, Ovid
Spotted hyaenas are sequential hermaphrodites and change sex
every year Plinius
‘The laugh of the hyaena represents mischievous delight over its WILSON 1975, citing
ability to change sex’ classic sources
TESTING HYPOTHESES ABOUT ADAPTATIONS
When we ask whether a trait is an adaptation, we ask whether it evolved
because ancestrally there was selection for this trait and the trait conferred a fitness
advantage because it performed a specific task (SOBER 1993). Hence we need to show
VIRILIZED GENITALIA OF FEMALE SPOTTED HYAENAS 897
that the trait conveyed a fitness advantage and that this fitness advantage arose from
design rather than chance (WILLIAMS 1966). The study of adaptation is then partly a
historical reconstruction. Hence, there can be several difficulties in testing the
hypothesis that a trait is the result of the process of adaptation (the Alternative
Hypothesis) against the null hypothesis that a trait is not the result of the process of
adaptation. Difficulties may arise in several ways:
je The ‘false negatives’: a trait may have historically served as an adaptation but
lacks current utility (i.e. it has no current adaptive value). Lack of current
adaptive value is consistent with the null hypothesis. However, the Alternative
Hypothesis is not falsified by demonstrating a lack of current adaptive value
(SOBER 1993).
DÒ The ‘false positives’: the trait originated as a by-product of selection for some-
thing else but possesses current utility (e.g. the ‘exaptation’ of GOULD & VRBA
1982). While current adaptive value is consistent with the Alternative Hypo-
thesis it is insufficient to reject the null hypothesis (GOULD & LEWONTIN 1979).
3 If the trait has gone to fixation, 1.e. there is no genotypic or phenotypic
variation amongst the members of a population or a taxon, the adaptedness of
such a trait may be difficult to determine because there is no agreed fitness
measure (FEDER & WARD 1992). It may also be difficult to decide which task
was associated with selection on the trait.
4. A trait may be complex in that it fulfills several tasks and may consist of sub-
components subject to independent selection pressures. This may be a common
empirical problem because a trait is by definition identified as an adaptation by
reference to a task while the study of a trait begins with a structure or
behaviour whose task(s) are initially not known.
Sì Selection on the trait may depend on interactions with other traits over evolu-
tionary time.
Faced with these difficulties, a research program could proceed along the fol-
lowing lines. First, study current use and assess the fitness consequences of the trait in
the field. If it is difficult to quantify fitness benefits directly, investigate whether
individuals incur a cost due to its presence. The plausibility argument would then be
that there must be fitness benefits compensating for these costs, otherwise the trait
would have been selected against. Second, construct a non-adaptive scenario that
explains the evolution of the trait without invoking specific tasks that the trait may
have been selected for. The non-adaptive argument is important; it states that we do
not require an adaptive explanation if we can identify a mechanism that creates the
trait as an incidental by-product of selection operating on a different trait. Third,
derive predictions from such a non-adaptive scenario and test them using field and
laboratory studies of single species, or by using the comparative method (HARVEY &
PAGEL 1991) on data from several species. Finally, compare the results with tests of
predictions derived from adaptive hypotheses.
898 HERIBERT HOFER & MARION L. EAST
CURRENT USE
An extensive field study of greeting ceremonies in Serengeti spotted hyaenas
demonstrated the prominent role of the erect ‘penis’ in animals of both sexes (EAST et
al. 1993). Spotted hyaena greetings are a ritualized, active form of submission that
confirm asymmetries in status between greeting partners (EAST et al. 1993). In
mammals, submissive greeting gestures are often derived from behavioral traits
typical of subordinate members of social groups, such as juveniles or females in male-
dominated societies (wolf, Canis lupus: MECH 1970; elephant, Loxodonta africana:
DOUGLAS-HAMILTON 1972; brown hyaena, Parahyaena brunnea: MILLS 1990; pri-
mates: WICKLER 1966, 1967, SUGIYAMA 1971, JoLLY 1972, STAMMBACH 1978,
COLMENARES 1990). The result that in the female-dominated society of spotted
hyaenas a typically male trait, the erect ‘penis’, is used as a signal of submission
during greetings is compatible with this line of thought. In all other mammalian
species that use penile displays during interactions, males are dominant over females,
and the erect ‘penis’ is used as a display of dominance (PLOOG & MACLEAN 1963,
WICKLER 1967, ANTHONEY 1968, JOLLY 1972, WEIGEL 1979). Even among platyrrhine
species (Ateles sp., Alouatta sp.) in which females have an enlarged clitoris, the
enlarged clitoris is used by females to display dominance (PLOOG 1967).
FITNESS CONSEQUENCES
Here we ask first whether anatomical components of the pseudo-‘penis’ or
behavioural features of its use are inexplicable unless we postulate selection pressures
that operate directly on the pseudo-‘penis’. We then review evidence on fitness
consequences of the pseudo-‘penis’ in three contexts.
One plausibility argument considers that large functionless organs are usually
selected against. As the pseudo-‘penis’ comprises circa 1% of adult body mass
(NEAVES ef al. 1980) it is a large organ and unlikely to persist if it conveyed no fitness
benefits (HAMILTON ef al. 1986). HAMILTON ef al. (1986) also pointed out that
experimental induction of virilization of female genitalia in a variety of species did
not produce a penile structure equivalent to that of the male.
Parturition. When female spotted hyaenas give birth, the pseudo-‘penis’ ruptures on the
caudal side, causing a large wound (several centimeters) that may be a source of
infection. Unusually many primiparous females produce stillbirths while subsequent
deliveries are easier (GLICKMAN er al. 1992a, unpublished data). Primiparous females in
the Serengeti also have a significantly smaller litter size than multiparous females
(unpublished data). Losing the whole first litter at parturition imposes a loss in terms of
lifetime reproductive success of at least 5% (unpublished data). These points suggest
that maintenance of the pseudo-‘penis’ in spotted hyaenas entails a cost and that direct
selection pressures operate to maintain the pseudo-‘penis’ in its current form.
Communication. No direct assessment of the fitness consequences of invoking the
pseudo-‘penis’ during greeting ceremonies is available. However, a plausibility argu-
VIRILIZED GENITALIA OF FEMALE SPOTTED HYAENAS 899
ment for fitness benefits may be constructed as follows. The erect ‘penis’ is the only
gesture that reliably signals rank as well as rank difference in greetings between adult
females and cubs (EAST et al. 1993). Thus, cubs can assess the social status of their
mother in relation to that of adult female greeting partners. The erection of the ‘penis’
therefore provides a mechanism which permits the enforcement of ‘rank inheritance’
(HoLEKAMP & SMALE 1991), a notable benefit at least to offspring of higher ranking
mothers.
Mating system. EAST et al. (1993) proposed a selective advantage for females that
have the opening to their reproductive tract (the urogenital meatus) displaced to a
position forward of their hind legs at the tip of their hypertrophied clitoris. We discuss
this in more detail below.
VIRILIZED FEMALE GENITALIA: NON-ADAPTIVE HYPOTHESES
The evolutionary origins of the pseudo-‘penis’ in spotted hyaenas have been
accredited to both non-adaptive processes and adaptive selection pressures. We begin
with null models.
The first non-adaptive hypothesis states that female virilization was a con-
sequence of a punctuated genetic translocation (VAN JAARSVELD & SKINNER 1987).
This scenario does not explain how the trait would have gone to fixation and makes
no predictions about trait design.
A second scenario suggests that female virilization was an incidental by-
product of selection for elevated prenatal androgen levels leading to large size and
social dominance of females over males (GouLp 1981, GouLp & VRBA 1982).
HAMILTON et al. (1986) thought that the initial virilization of female genitalia evolved
as a by-product of selection for larger body size and aggressiveness to obtain a
competitive feeding advantage. The second scenario predicts
je species with virilized females are more likely to have female-dominated
societies than those without;
2, females are larger than males in female-dominated societies;
3 large, aggressive individuals become dominant;
4. virilized females have higher concentrations of androgens than non-virilized
females.
Virilization of female external genitalia is rare in mammalian species and
where it occurs it is not associated with female dominance in any species other than
the spotted hyaena (e.g. European moles Talpa europaea: MATTHEWS 1935, platyr-
rhine monkeys: PLOOG 1967, OPPENHEIMER 1977). In mammalian species, female
dominance is also uncommon, but does occur in a wide variety of species (RALLS
1976, HRDY 1981) and is typical for Lemuriformes (JOLLY 1984, RICHARD 1987,
KAPPELER 1990, YOUNG et al. 1990). Masculinization of female external genitalia has
not been recorded in any female-dominated species except for the spotted hyaena.
Female dominance in mammals is not dependent on females being larger than males,
900 HERIBERT HOFER & MARION L. EAST
and dominance is not a function of aggressiveness (RowELL 1974). Even in spotted
hyaenas, females are not necessarily larger than males (HAMILTON et al. 1986,
unpublished data) and body size in males and females is not related to social rank in
our Serengeti population nor in captive animals (FRANK et al. 1989). In several
primate species (Saimiri, Cebus, Cercopithecus talapoin, Macaca mulatta, Erythro-
cebus patas) coalitions of females can successfully challenge dominance of larger
males (HALL 1967, KAPLAN & ZUCKER 1980, HARVEY et al. 1987, ROBINSON et al.
1987, SMUTS 1987), while in spotted hyaenas coalitions are important in the
maintenance of female rank (see above), and may have been important in the
development of female dominance (FRANK et al. 1989). Adult female spotted hyaenas
do not have significantly higher levels of androgens than adult males (RACEY &
SKINNER 1979, FRANK ef al. 1985, LINDEQUE et al. 1986, VAN JAARSVELD & SKINNER
1991; GLICKMAN et al. 1992b). Androgen levels in female spotted hyaenas are not
elevated in relation to androgen levels in female brown or striped hyaenas (VAN
JAARSVELD & SKINNER 1987). The striking aspect of the hormonal profile of adult
spotted hyaenas are the low androgen levels of adult males compared with males of
other hyaena species (VAN JAARSVELD & SKINNER 1987, 1991). In summary, neither
comparative evidence nor current knowledge of spotted hyaena behavior and endocri-
nology appear to support the non-adaptive scenario that links selection for social
dominance with elevated pre-natal androgen levels via body size or aggressiveness.
A third scenario links pre-natal androgen levels to siblicide (HOFER & EAST
1992, East et al. 1993) as there is frequent fatal neonatal aggression between
members of a twin litter (FRANK et al. 1991). The third scenario predicts that
Ne spotted hyaenas have higher androgen concentrations during the period of
siblicide (several weeks beginning shortly after birth) than species without
siblicide;
Dee patterns of maternal investment favour strong neonatal aggression leading to
siblicide;
3% species with neonatal siblicide have traits that undergo accelerated foetal
development compared to species without;
4. species with neonatal siblicide are more likely to be virilized than those
without.
Siblicide is associated with extremely high neonatal aggression and high
androstenedione levels at birth and during ontogeny (FRANK et al. 1991, LICHT et al.
1992). FRANK et al. (1991) proposed that siblicide is "another phenomenon that may
have originated as a by-product of selection for female androgenization". In contrast,
we suggested that the exceptionally high (GITTLEMAN & OFTEDAL 1987) investment in
lactation and long nursing period in spotted hyaenas (EAST et al. 1989, HOFER & EAST
1993) favours strong selection for high neonatal aggression leading to siblicide
(HOFER & EAST 1992). Following this line of argument, the elevated levels of andro-
gens (androstenedione in females, testosterone in males) recorded during foetal deve-
lopment, at birth, and during the initial months of life (GLICKMAN et al. 1987, FRANK
VIRILIZED GENITALIA OF FEMALE SPOTTED HYAENAS 901
et al. 1991, LIcHT et al. 1992) have been selected for to produce neonates primed for
severe agonistic encounters with fully erupted teeth and open eyes. Spotted hyaenas
are the only species of carnivores currently known to be born with fully erupted teeth
and open eyes. Developmental acceleration of siblicide-relevant anatomical structures
and behaviours appear to place spotted hyaenas on the borderline between altricial
and precocial species (MARTIN & MACLARNON 1985, van JAARSVELD et al. 1988).
Spotted hyaenas are currently the only mammalian species known with frequent
neonatal siblicide (FRANK et al. 1991), and the only species with such elaborately
virilized genitalia. We conclude that current evidence is consistent with the Hofer &
East hypothesis.
VIRILIZED FEMALE GENITALIA: ADAPTIVE HYPOTHESES
The first adaptive hypothesis attributed the evolution of the pseudo-‘penis’
directly to selective advantages of a structure that becomes conspicuous through erec-
tion, focussing the attention of greeting partners and keeping them together over a
longer period, thereby facilitating the re-establishment of social bonds (WICKLER
1965, KRUUK 1972). The second adaptive hypothesis attributed the evolution of the
pseudo-‘penis’ indirectly to a two-stage process where after the initial evolution of
virilization (see above) a penile signal was incorporated into greetings, favouring
selection for the enlargement and erectile nature of the pseudo-‘penis’ (HAMILTON et
al. 1986). These two adaptive hypotheses predict:
Il, Adults that meet infrequently should display an erect penis more often than
adults that meet frequently. This means that males should erect their penises in
all-male greetings more often than females in all-female greetings (HOFER &
EAST 1993).
2 Greetings should last longer and be more likely to be asymmetric with respect
to ‘penis’ erection in cases where the benefits of greetings are likely to be
larger. This means that greetings ought to last longer and be more likely to be
asymmetric in greetings between adult females than other age/sex categories
(EAST et al. 1993).
3. The erection of the penis provides information not available through other
gestures during the greeting ceremony.
All three predictions were confirmed by EAST er al.'s (1993) field study.
Greetings between immigrant males comprised a substantially higher proportion of
cases with both individuals erecting their ‘penis’ than greetings between adult females.
Greetings between adult females lasted significantly longer and comprised a higher
proportion of asymmetric greetings with respect to ‘penis’ erection than greetings
between males. There are gestures besides the erect ‘penis’ that are available to signal
submission. However, the erect ‘penis’ is the only gesture that reliably signals absolute
rank as well as rank difference in greetings between adult females and cubs. This
emphasizes the influence of the rank of a cub's mother on the socialization of the cub
902 HERIBERT HOFER & MARION L. EAST
because it permits cubs to assess the social status they obtain from their mother in
relation to that of an adult female greeting partner. Thus, it provides a mechanism that
permits the enforcement of ‘rank inheritance’ (HOLEKAMP & SMALE 1991).
The third adaptive hypothesis proposed that after the initial virilization a
selective advantage accrued to those females that had the opening to their repro-
ductive tract displaced to a position forward of their hind legs because they gained
control over the mechanics of copulation and male mating success now depended on
the goodwill of the females (EAST ef al. 1993). The third hypothesis predicts that
je males expend extensive effort on courting females;
DA aggression amongst contemporary males is low;
3 male spotted hyaenas have unusually low androgen levels;
4. asymmetries in gestures in greetings between males are less common than in
greetings between females;
I. number of forced greetings between immigrant males is low.
The structure and position of the pseudo-‘penis’ dictates that males have consi-
derable difficulty locating and penetrating the clitoris opening (KRUUK 1972, own
observations). Although the pseudo-‘penis’ is not erect during mating, its presence
and position prevent males from achieving intromission without the complete co-
operation of the female, i.e. forced copulations are impossible. Females thus gain
control over the mechanics of copulation and male mating success becomes
dependent on the relationships they develop with females. Then, the importance of
sexual selection for male fighting prowess would be minimal because an amicable
relationship with a female is a non-transferrable resource. As males achieve little by
fighting with each other game theory predicts that fights should be rare and/or
fighting intensity low (MAYNARD SMITH 1982, HARPER 1991). Androgen levels of
males are unusually low and aggression amongst contemporary males is low (FRANK
1986, EAST & HOFER 1991). Males do expend extensive effort on courting females
(EAST & HOFER 1991). Asymmetries in gestures in greetings between males are less
common than in greetings between females and the number of forced greetings
between immigrant males was low (EAST et al. 1993). In short, maintenance of the
pseudo-‘penis’ may have facilitated the evolution of male submission by providing
females with control over mating in a species where males do not contribute to
parental care.
A SIMPLE OR A COMPLEX TRAIT?
Several lines of evidence suggest that the pseudo-‘penis’ may represent a
complex trait comprising different components subjected to independent selection
events. For instance, while parturition in primiparous spotted hyaenas is associated
with unusually high litter losses, there are no reports that parturition is impeded in
primiparous primates with virilized genitalia. Female European moles have a separate
vaginal opening at the base of their pseudo-‘penis’ (MATTHEWS 1935, NIETHAMMER
VIRILIZED GENITALIA OF FEMALE SPOTTED HYAENAS 903
1990) which facilitates both mating and parturition. There are no reports that females
of species with virilized genitalia can fully erect the enlarged clitoris other than
spotted hyaenas. This is probably a consequence of the sexual dimorphism found in
Onufs nucleus, the area in the sacral spinal chord responsible for erecting the ‘penis’
which is well-developed in males but not in females in all species studied so far
(BREEDLOVE & ARNOLD 1980, FORGER & BREEDLOVE 1986). We predict that in
spotted hyaenas, Onuf's nucleus is monomorphic in structure. If this is the case, then
the ability to erect the pseudo-‘penis’ might be another trait unexplained by the cur-
rent non-adaptive scenarios.
CONCLUSIONS
The example of virilized genitalia of female spotted hyaenas shows that the
study of adaptations benefits from carefully considering non-adaptive scenarios on the
evolutionary origins of a trait. What was initially considered one trait (the virilized
genitalia) may well turn out to be a complex of several sub-traits subjected to dif-
ferent selection pressures. Here the analysis of comparative data plays a crucial role in
deciding whether the trait is simple or complex in this sense. This difficulty is likely
to arise frequently because a trait is identified as an adaptation by reference to a task
while the study of a trait begins with a structure or behaviour whose task(s) are ini-
tially not known. We have argued that the pseudo-‘penis’ has negative fitness conse-
quences in the context of parturition but the argument is less tight than it could be if
hyaena populations consisted of females with and without pseudo-‘penises’. In other
words, lack of variation in a trait because it has gone to fixation makes it difficult to
assess the adaptedness of a trait. Several arguments suggest that the virilized genitalia
possess current utility as well as specificity in the sense that this trait evolved (after its
initial origination) to fulfill two specific tasks in communication and mating.
The discussion of one trait has led us to use arguments from many disciplines
dedicated to questions about both proximate and ultimate causes. It also became
important to consider the possible links and interactions to other anatomical, beha-
vioural and life history traits. While the reconstruction of a historical process will
remain incomplete without a complete fossil record, careful formulation of hypo-
theses and predictions can take us some way to establishing traits as adaptations.
REFERENCES
ANTHONEY, T.R. 1968. The ontogeny of greeting, grooming, and sexual motor patterns in
captive baboons (superspecies Papio cynocephalus). Behaviour 31: 358-372.
BREEDLOVE, S.M., A.P. ARNOLD. 1980. Hormone accumulation in a sexually dimorphic motor
nucleus of the rat spinal cord. Science 210: 564-566.
COLMENARES, F. 1990. Greeting behaviour in male baboons, I: Communication, reciprocity and
symmetry. Behaviour 113: 81-114.
DOBZHANSKY, T. 1968. On some fundamental concepts of Darwinian biology. Evol. Biol. 2:
1-34.
904 HERIBERT HOFER & MARION L. EAST
DOUGLAS-HAMILTON, I. 1972. On the ecology and behaviour of the African elephant: the
elephants of Lake Manyara. DPhil. thesis, Oxford University.
East, M.L., H. HOFER, A. TURK. 1989. Functions of birth dens in spotted hyaenas (Crocuta
crocuta). J. Zool. Lond. 219: 690-697.
East, M.L., H. HOFER. 1991. Loud calling in a female-dominated mammalian society: II.
Behavioural contexts and functions of whooping of spotted hyaenas, (Crocuta crocuta).
Anim. Behav. 42: 651-669.
EAST, M.L., H. HOFER, W. WICKLER. 1993. The erect ‘penis’ is a flag of submission in a
female-dominated society: greetings in Serengeti spotted hyenas. Behav. Ecol. Socio-
biol. 33: 355-370.
ENDLER, J.A. 1986. Natural selection in the wild. Princeton University Press, Princeton.
FEDER, M.E., W.B. WATT. 1992. Functional biology of adaptation. /n: BERRY R.J., CRAWFORD
T.J., HEWITT G.M. (eds), Genes in ecology. Blackwell Scientific, Oxford, pp. 365-392.
FORGER, N.G., S.M. BREEDLOVE. 1986. Sexual dimorphism in human and canine spinal cord:
role of early androgen. Proc. Nat. Acad. Sci. USA 83: 7527-7531.
FRANK, L.G. 1986. Social organisation of the spotted hyaena (Crocuta crocuta). II. Dominance
and reproduction. Anim. Behav. 34: 1510-1527.
FRANK, L.G., J.M. DAvIDSON, E.R. SMITH. 1985. Androgen levels in the spotted hyaena
Crocuta crocuta: the influence of social factors. J. Zool. Lond. A206: 525-531.
FRANK, L.G., S.E. GLICKMAN, P. LICHT. 1991. Fatal sibling aggression, precocial development
and androgens in neonatal spotted hyaenas. Science 252: 702-704.
FRANK, L.G., S.E. GLICKMAN, I. PowcH. 1990. Sexual dimorphism in the spotted hyaena
(Crocuta crocuta). J. Zool. Lond. 221: 308-313.
FRANK, L.G., S.E. GLICKMAN, C.J. ZABEL. 1989. Ontogeny of female dominance in the spotted
hyaena: perspectives from nature and captivity. Symp. zool. Soc. Lond. 61: 127-146.
GITTLEMAN, J.L., O.T. OFTEDAL. 1987. Comparative growth and lactation energetics in carni-
vores. Symp. zool. Soc. Lond. 57: 41-77.
GLICKMAN, S.E., L.G. FRANK, J.M. DAVIDSON, E.R. SMITH, P.K. SITERI. 1987. Androstenedione
may organize or activate sex-reversed traits in female spotted hyaena. Proc. Nat. Acad.
Sci. USA 84: 3444-3447.
GLICKMAN, S.E., L.G. FRANK, P. LICHT, T. YALCINKAYA, P.K. SIITERI, J. DAVIDSON. 1992a.
Sexual differentiation of the female spotted hyena. Ann. New York Acad. Sci. 662:
135-159.
GLICKMAN, S.E., L.G. FRANK, S. PAVGI, P. LICHT. 1992b. Hormonal correlates of ‘masculini-
zation’ in female spotted hyaenas (Crocuta crocuta). I. Infancy to sexual maturity. J.
Reprod. Fert. 95: 451-462.
GOULD, S.J. 1981. Hyena myths and realities. Nat. Hist. 90: 16-24.
GOULD, S.J., R.C. LEWONTIN. 1979. The spandrels of San Marco and the Panglossian paradigm:
a critique of the adaptionist program. Proc. Roy. Soc. Lond. B205: 581-598.
GOULD, S.J., E.S. VRBA. 1982. Exaptation — a missing term in the science of form. Paleobiol.
8:4-15.
HALL, K.R.L. 1967. Social interactions of the adult male and adult females of a patas monkey
group. Jn: ALTMANN S.A. (ed.), Social communication among primates. University of
Chicago Press, Chicago, pp. 261—280.
HAMILTON, W.J., R.L. TILSON, L.G. FRANK. 1986. Sexual monomorphism in spotted hyaenas,
Crocuta crocuta. Ethology 71: 63-73.
HARPER, D.G.C. 1991. Communication. /n: KREBS J.R., DAVIES N.B. (eds), Behavioural eco-
logy: an evolutionary approach. Blackwell Scientific, Oxford, pp. 374-397.
HARVEY, P.H., R.D. MARTIN, T.H. CLUTTON-BROCK. 1987. Life histories in comparative perspec-
tives. Jn: SMUTS B.B., CHENEY D.L., SEYFARTH R.M., WRANGHAM R.W., STRUHSAKER
T.T. (eds), Primate societies. University of Chicago Press, Chicago, pp. 181-196.
VIRILIZED GENITALIA OF FEMALE SPOTTED HYAENAS 905
Harvey, P.H., M.D. PAGEL. 1991. The comparative method in evolutionary biology. Oxford
University Press, Oxford.
HOFER, H., M.L. EAST. 1992. Siblicide of spotted hyaenas is a consequence of allocation of
maternal effort and a cause of skewed litter sex ratios. Abstracts Fourth Int. Behav.
Ecol. Congr. Princeton, New Jersey.
HOFER, H., M.L. EAST. 1993. The commuting system of Serengeti spotted hyaenas: how a
predator copes with migratory prey. III. Attendance and maternal care. Anim. Behav.
46: 575-589.
HOLEKAMP, K.E., L. SMALE. 1991. Dominance acquisition during mammalian social develop-
ment: The "inheritance" of maternal rank. Am. Zool. 31: 306-317.
HRDY, S.B. 1981. The woman that never evolved. Harvard University Press, Cambridge.
JAARSVELD, A.S. VAN, J.D. SKINNER. 1987. Spotted hyaena monomorphism: an adaptive
‘phallusy’? S. Afr. J. Sci. 83: 612-615.
JAARSVELD, A.S. VAN, J.D. SKINNER, M. LINDEQUE. 1988. Growth, development and parental
investment in the spotted hyaena Crocuta crocuta. J. Zool. Lond. 216: 45-53.
JAARSVELD, A.S. VAN, J.D. SKINNER. 1991. Plasma androgens in spotted hyaenas (Crocuta cro-
cuta): influence of social and reproductive development. J. Reprod. Fert. 93: 195-201.
JoLLY, A. 1972. The evolution of primate behaviour. Macmillan, New York.
JoLLY, A. 1984. The puzzle of female feeding priority. In: SMALL M. (ed.), Female Primates:
studies by women primatologists. Alan R. Liss, New York.
KAPLAN, J.R., E. ZUCKER. 1980. Social organization of a group of free-ranging Patas monkeys.
Folia Primatol. 34: 196-213.
KAPPELER, P.M. 1990. Female dominance in Lemur catta: more than just female feeding
priority? Folia Primatol. 55: 92-103.
KRUUK, H. 1972. The spotted hyena. University of Chicago Press, Chicago.
LicuT, P., L.G. FRANK, S. PAVGI, T.M. YALCINKAYA, P.K. SUTERI, S.E. GLICKMAN. 1992.
Hormonal correlates of ‘masculinization’ in female spotted hyaenas (Crocuta crocuta)
2. Maternal and fetal steroids. J. Reprod. Fert. 95: 463-474.
LINDEQUE, M., J.D. SKINNER. 1982. Fetal androgens and sexual mimicry in spotted hyaena
(Crocuta crocuta). J. Reprod. Fert. 65: 405-410.
LINDEQUE, M., J.D. SKINNER, R.P. MILLAR. 1986. Adrenal and gonadal contribution to circu-
lating androgens in spotted hyaenas (Crocuta crocuta) as revealed by LHRH, hCG and
ACTH stimulation. J. Reprod. Fert. 78: 211-217.
MARTIN, R.D., A.M. MACLARNON. 1985. Gestation period, neonatal size, and maternal
investment in placental mammals. Nature 313: 220-223.
MATTHEWS, L.H. 1935. The oestrus cycle and intersexuality in the female mole Talpa
europaea. Linn. Proc. zool. Soc. Lond. 106: 347-383.
MATTHEWS, L.H. 1939. Reproduction in the spotted hyaena (Crocuta crocuta Erxleben). Phil.
Trans. Roy. Soc. Lond. B230: 1-78.
MAYNARD SMITH, J. 1982. Evolution and the theory of games. Cambridge University Press,
Cambridge.
MECH, L.D. 1970. The wolf. University of Minnesota Press, Minneapolis.
Mitts, M.G.L. 1990. Kalahari hyaenas. The comparative behavioural ecology of two species.
Unwin Hyman, London.
NEAVES, W.B., J.E. GRIFFIN, J.D. WILSON. 1980. Sexual dimorphism of the phallus in spotted
hyaena (Crocuta crocuta). J. Reprod. Fert. 59: 509-513.
NIETHAMMER, J. 1990. Talpa europaea Linnaeus, 1758 — Maulwurf. In: NIETHAMMER J., KRAPP
F. (eds), Handbuch der Säugetiere Europas 3/I: Insektenfresser, Primaten. Aula Verlag,
Wiesbaden, pp. 99-133.
906 HERIBERT HOFER & MARION L. EAST
OPPENHEIMER, J.R. 1977. Communication in New World monkeys. /n: SEBEOK T.A. (ed.), How
animals communicate. Indiana University Press, Bloomington, pp. 851-889.
PLooG, D.W. 1967. The behaviour of squirrel monkeys (Saimiri sciureus) as revealed by
sociometry, bioacoustics, and brain stimulation. /n: Altmann S.A. (ed.), Social commu-
nication among primates. University of Chicago Press, Chicago, pp. 149-184.
PLO0G, D.W., P.D. MACLEAN. 1963. Display of penile erection in squirrel monkeys (Saimiri
sciureus). Anim. Behav. 11: 32-39.
Racey, P.A., J.D. SKINNER. 1979. Endocrine aspects of sexual mimicry in spotted hyaenas
(Crocuta crocuta). J. Zool. Lond. 187: 315-326.
RALLS, K. 1976. Mammals in which females are larger than males. Quart. Rev. Biol. 51:
245-276.
RICHARD, A.F. 1987. Malagasy prosimians: female dominance. /n: SMuTS B.B., CHENEY D.L.,
SEYFARTH R.M., WRANGHAM R.W., STRUHSAKER T.T. (eds), Primate societies.
University of Chicago Press, Chicago, pp. 25-33.
ROBINSON, J.G., P.C. WRIGHT, W.G. KINZEY. 1987. Monogamous cebids and their relatives:
intergroup calls and spacing. Jn SMUTS B.B., CHENEY D.L., SEYFARTH R.M., WRANG-
HAM R.W., STRUHSAKER T.T. (eds), Primate societies. University of Chicago Press,
Chicago, pp. 44-53.
ROWELL, T.H. 1974. The concept of social dominance. Behav. Biol. 11: 131-154.
SHERMAN, P.W. 1988. The levels of analysis. Anim. Behav. 36: 616-618.
Smuts, B.B. 1987. Gender, aggression, and influence. /n: SMUTS B.B., CHENEY D.L., SEYFARTH
R.M., WRANGHAM R.W., STRUHSAKER T.T. (eds), Primate societies. University of
Chicago Press, Chicago, pp. 400-412.
SOBER, E. 1993. Philosophy of biology. Oxford University Press, Oxford.
STAMMBACH, E. 1978. On social differentiation in groups of captive female hamadryas
baboons. Behaviour 67: 322-338.
SUGIYAMA, K. 1971. Characteristics of the social life of bonnet macaques (Macaca radiata).
Primates 12: 247-266.
WEIGEL, R.M. 1979. The facial expression of the brown capuchin monkey (Cebus apella).
Behaviour 68: 250-276.
WICKLER, W. 1964. Vom Gruppenleben einiger Säugetiere Afrikas. Mitt. Max-Planck-Ges. 5-6:
296-309.
WICKLER, W. 1965. Die Evolution von Mustern der Zeichnung und des Verhaltens. Naturwiss.
52: 335-341.
WICKLER, W. 1966. Ursprung und biologische Deutung des Genitalpräsentierens männlicher
Primaten. Z. Tierpsychol. 23: 422-437.
WICKLER, W. 1967. Socio-sexual signals and their intra-specific imitation among primates. In:
MORRIS D. (ed.), Primate ethology. Weidenfeld & Nicolson. London, pp. 69-147.
WILLIAMS, G.C. 1966. Adaptation and natural selection. Princeton University Press, Princeton.
WILSON, E.O. 1975. Sociobiology. Harvard University Press, Cambridge Mass.
Young, A.L., A.F. RICHARD, L.C. ATELLO. 1990. Female dominance and maternal investment in
strepsirrhine primates. Am. Nat. 135: 473-488.
REVUE SUISSE DE ZOOLOGIE, 102 (4) : 907-912; décembre 1995
Recovery from eutrophication delayed in the profundal
of Lake Neuchatel: evidence from the oligochaete communities
Claude LANG & Olivier REYMOND
Conservation de la faune, Marquisat 1,
CH-1025 St-Sulpice, Switzerland.
Recovery from eutrophication delayed in the profundal of Lake Neu-
chatel: evidence from the oligochaete communities.- In Lake Neuchatel
(Switzerland), total phosphorus (TP) concentrations decreased from 42 mg
m-3 in 1982 to 18 mg m-3 in 1994. In 1994, responses of oligochaete
communities to this decrease were analysed in the deepest area (74 m—139
m) of the northern basin. Mean relative abundance of oligotrophic species
(mostly Spirosperma velutinus and Stylodrilus heringianus) was around
14%, as in a meso-eutrophic lake, and well below 39%, the value predicted
from TP. In contrast, at a depth of 40 m, mean relative abundance of oligo-
trophic species increased from 16% in 1984 to 33% in 1992, as predicted
from TP. In Lake Neuchatel as in Lake Geneva, responses of oligochaete
communities to the abatement of eutrophication were slower and less clear
in the deepest areas than in the sublittoral.
Key-words: Biomonitoring - Eutrophication - Indicator species - Lake -
Zoobenthos.
INTRODUCTION
In Lake Neuchatel (Switzerland), mean total phosphorus concentrations de-
creased from 42 mg m- in 1982 to 18 mg m> in 1994 (PoKORNI, 1994). At a depth
of 40 m, oligochaete communities responded clearly to the increase (LANG &
REYMOND, 1993): mean relative abundance of oligotrophic species increased from
16% in 1984 to 33% in 1992. At the same depth, the same trend was observed in Lake
Geneva between 1982 and 1991 (LANG & REYMOND, 1992) but, at a depth of 150 m,
the response of worms was slower and less clear (LANG & REYMOND, 1995).
In the present study, we analyse the response of oligochaete communities to
the abatement of eutrophication in the deepest area (74 m - 139 m) of the northern
basin of Lake Neuchatel. Our goal is to determine if the recovery, observed at a depth
of 40 m, has extended its effects down to the profundal. In this case, the observed
relative abundance of oligotrophic species will be close to the value predicted from
total phosphorus concentrations (LANG, 1990).
Manuscript accepted 03.04.1995.
908 CLAUDE LANG & OLIVIER REYMOND
STATIONS AND METHODS
Sixty-five sampling sites were visited in June 1994 in the northern basin of
Lake Neuchatel (Fig. 1). Sites, located with a GPS (Global Positioning System) re-
ceptor, were regularly spaced (500 m apart) on 5 parallel transects. These five
transects (500 m apart) covered the whole profundal (depth of sites: 74 - 139 m, mean
127 m), forming a regular grid system.
In each site, one 16 cm? sediment core (30 cm long) was taken from the
surface with a gravity corer. In the laboratory, the sediment was sieved (mesh size:
0.2 mm) and the retained material preserved in 5% formalin. The collected macro-
fauna consisted mainly of tubificid and lumbriculid worms which were picked and
counted. Worms whose diameter was greater than 0.29 mm were mounted (REYMOND,
1994) and identified to species. Juvenile worms (diameter less than 0.3 mm) were
excluded to decrease the effects of seasonal variability on species abundance (LANG,
1991). The biological analysis was based on 48 cores in which adult worms were
present. Seventeen cores were excluded because macrofauna was absent (7 cores) or
only a few juvenile worms were present.
Species whose numerical dominance in tubificid and lumbriculid communities
indicates, respectively, oligotrophic, mesotrophic or eutrophic conditions (LANG,
1991) were designated oligotrophic, mesotrophic and eutrophic species in Table 1.
Relative abundance (%) of oligotrophic species (OS), calculated as a percentage of
the total number of tubificid and lumbriculid worms present in each core, was
predicted from total phophorus (TP) concentrations (mg m73) recorded in the water
during the five years preceding the sampling of worms (LANG, 1990):
Equation | OS = 80.29 — 8.35 TP05
where TP = 51.5 mg m-3 (mean for 1979 - 1983), TP = 29.4 mg m-3 (mean for
1987-1991), TP = 24.6 mg m-3 (mean for 1989 - 1993), (POKORNI, 1994).
The area studied in 1994 has not been visited previously, except in part by MONARD
(1919) in 1918. Therefore, we used the area visited in 1984 and 1992, at a depth of 40
m, as a reference (Fig. 1).
RESULTS
Sampling sites visited in 1994 were located deeper (127 m) than those studied
in 1984 and 1992 (40 m). Therefore some of the differences recorded in 1994 can be
attributed to depth rather than to eutrophication. For instance, abundance of species 5
to 8 of table 1 tends to decrease with depth, even in oligotrophic lakes (LANG, 1990).
Potamothrix vejdovskyi, absent from Lake Neuchatel in 1984, was relatively frequent
in 1992 (LANG & REYMOND, 1993). In Lake Geneva, the abundance of this species
decreases below 100 m, probably because its reproduction is limited by low tempe-
rature (LANG & REYMOND, 1995). Therefore its scarcity in 1994 can also be ascribed
to depth, as indicated by its distribution (Fig. 1).
OLIGOCHAETES IN LAKE NEUCHATEL 909
Fic. 1
A. Location of sampling areas (with sampling years indicated) in Lake Neuchatel. Area: 215 km2,
mean depth: 64 m, max. depth: 153 m, volume: 13.8 km3 (SOLLBERGER, 1974). La Motte is an
underwater hill whose top is 10 m under the surface. B. Location of the sampling sites visited in
1994. Presence of oligotrophic worm species (large circle), of mesotrophic species (intermediate
circle), and of eutrophic species only (small circle), site without macrofauna (dash).
910 CLAUDE LANG & OLIVIER REYMOND
In 1994, oligotrophic species, which are not limited by depth in oligotrophic
lakes (LANG, 1990), were mainly present on the edge of the deepest area (Fig. 1). In
contrast, sites in which only eutrophic species were present or sites without
macrofauna were mostly located in the deepest area. These different distributions
could be explained by a deterioration of oxygen conditions with depth.
Occurence of oligotrophic species indicated that the 1994 situation, expressed
in terms of trophic state, was closer to that of 1984 than to that of 1992 (Tab. 1).
Indeed, mean relative abundance of oligotrophic species was the same in 1984 and in
1994, but it was higher in 1992 (Tab. 2).
TABLE |
Changes recorded between 1984 and 1994 in lumbriculid and tubificid communities of Lake
Neuchatel. For species 1 and 2, results pooled together. Samples collected at a mean depth of
40 m in 1984 and 1992, of 127 m in 1994. Values for species 6 to 10 based only on mature
individuals.
Occurrence (%)
1984 1992 1994
Code Species (n=64) (n=171) (n= 48)
1 Bichaeta sanguinea Bretscher
2 Stylodrilus heringianus Claparede 37765 46.8 6.2
3} Spirosperma velutinus (Grube) 12 195 25.0
4 Potamothrix vejdovskyi (Hrabe) 0 23.4 4.2
5 Spirosperma ferox (Eisen) 1.6 13.5 0
6 Potamothrix moldaviensis (Vejdovsky, Mrazek) 0 93 0
7 Limnodrilus hoffmeisteri (Claparède) 6.2 4.1 0
8 Limnodrilus profundicola (Verrill) 7.8 0 0
9 Potamothrix hammoniensis (Michaelsen) 10.9 152 221
10 Tubifex tubifex (Müller) 25.0 7.6 8.3
Species 7-8 (immatures included) 40.6 28.1 ZA
Species 9-10 (immatures included) 100.0 67.3 DIL
Oligotrophic species 1-3 43.8 SOA LS
Mesotrophic species 4-6 1.6 3291 4.2
97,
Eutrophic species 7-10 100.0 VW
TABLE 2
Changes recorded in oligochaete communities of Lake Neuchatel between 1984 and 1994.
Significantly (P = 0.05) different means underlined. Right of the mean: standard error.
Species h
Mean abundance Year Oligotrophic Mesotrophic Eutrophic
Absolute 1984 1.01 0.19 0.02 0.02 4.97 0.39
(No. per core) 1992 1215 0.10 0.58 0.08 1.89 0.15
1994 0.33 0.07 0.04 0.03 2.50 0.38
Relative 1984 15.8 2.8 0.3 0.3 83.9 2.8
(%) 1992 33.0 DA] 14.6 1.9 52.4 29
1994 14.4 39 Sal 253 82.5 4.3
OLIGOCHAETES IN LAKE NEUCHATEL 911
DISCUSSION
Around 1900, Lake Neuchatel was oligotrophic as most of the large and deep
lakes of Switzerland (FRICKER, 1980). As in these other lakes (PIGUET & BRETSCHER,
1913), oligotrophic species (mostly Spirosperma velutinus and Stylodrilus herin-
gianus) were present in most (90%) samples collected in 1918 in the northern basin of
Lake Neuchatel (MONARD, 1919). In this first study, Spirosperma velutinus demons-
trates already its indicator value: this species was absent from areas receiving heavy
organic inputs (dead leaves) from the Areuse river.
Mean relative abundance of oligotrophic species was not computed for this
first study because individuals were not counted. Instead, the mean abundance of
these species, observed in Lake Superior (North America), was used as a reference for
oligotrophic conditions (LANG, 1990). According to results from several lakes (LANG,
1979, 1990), mean values decrease from 70% in oligotrophic lakes to around 35% in
mesotrophic lakes, to around 17.5% in meso-eutrophic lakes, and to zero in the
profundal of eutrophic lakes. These values were used as references to interpret results
from Lake Neuchatel.
Following the increase of phosphorus concentrations from 12 mg m-3 in 1962
to 67 mg m-3 in 1980 (LANG, 1989), mean relative abundance of oligotrophic species
decreases to 9% in 1980 in the whole Lake Neuchatel, i.e. close to zero, the value
indicative of eutrophic conditions. In the deepest area (150 m), the abundance was as
low as 2.4%, but on the Motte, an underwater hill swept by currents (Fig. 1), mean
values higher than 30% persisted.
After the decrease of phosphorus concentrations, mean abundance of oligo-
trophic species, recorded at a depth of 40 m, indicated that the lake was meso-
eutrophic in 1984, but mesotrophic in 1992 (Tab. 2). In contrast, the lake remained
meso-eutrophic in 1994, at a mean depth of 127 m, according to oligotrophic species.
Mean relative abundance of these species, predicted from total phosphorus concen-
trations (Equation 1, see Stations and methods), was 20.4% for 1984, 35% for 1992,
and 38.9% for 1994 respectively. Predicted and observed values were close in 1984
and 1992, but the difference was large in 1994 (Tab. 2). The low abundance observed
in 1994 indicated that the recovery from eutrophication was delayed in the profundal
of Lake Neuchatel. The same trend was observed in Lake Geneva (LANG &
REYMOND, 1995).
A shortage of oxygen at the interface between sediment and water could
explain this delay. Mortality of whitefish (Coregonus sp.) eggs, a species typical of
oligotrophic lakes, was high in some Swiss lakes (MULLER, 1992) even if, as in Lake
Neuchatel (POKORNI, 1994), oxygen concentrations, measured 1 m above the bottom,
were always higher than 7 mg 1-!. However, oxygen uptake by the sediment was
large enough in these previously productive lakes to create a thin but almost anoxic
layer at the interface. Hence, embryogenesis of fish eggs was inhibited. If such an
anoxic layer exists above the deep sediments of Lake Neuchatel, it could also affect
oligotrophic species, especially juveniles, which respire at the interface level.
912 CLAUDE LANG & OLIVIER REYMOND
In conclusion, the recovery of Lake Neuchatel will be complete only if the
sediments, especially those of the profundal, are recolonized by the oligotrophic
species which prevailed therein, before the onset of eutrophication. This
recolonization will indicate that the sediment has recovered its pristine state, as far as
this goal is realistic.
ACKNOWLEDGEMENTS
Raymond Ducret helped to collect samples.
REFERENCES
FRICKER, HJ. 1980. Eutrophication programme regional project, alpine lakes. Swiss Federal
Board for environmental Protection, Bern.
LANG, C. 1989. Eutrophication of Lake Neuchatel indicated by the oligochaete communities.
Hydrobiologia 174: 57-65.
LANG, C. 1990. Quantitative relationships between oligochaete communities and phosphorus
concentrations in lakes. Freshw. Biol. 24: 327-334.
LANG, C. 1991. Decreasing phosphorus concentrations and unchanged oligochaete communities
in Lake Geneva: how to monitor recovery? Arch. Hydrobiol. 122: 305-312.
LANG, C. & O. REYMOND. 1992. Reversal of eutrophication in Lake Geneva: evidence from the
oligochaete communities. Freshw. Biol. 28: 145-148.
LANG, C. & O. REYMOND. 1993. Recovery. of Lake Neuchatel (Switzerland) from eutro-
phication indicated by the oligochaete communities. Arch. Hydrol. 128: 65-71.
LANG, C. & O. REYMOND. 1995. Contrasting responses of oligochaete communities to the
abatement of eutrophication in Lake Geneva. Hydrobiologia 308: 77-82.
MONARD, A. 1919. La faune profonde du Lac de Neuchâtel. Thèse Univ. Neuchâtel. 176 pp.
MULLER, R. 1992. Trophic state and its implications for natural reproduction of salmonid fish.
Hydrobiologia 243/244: 261-268.
PIGUET, E. & K. BRETSCHER. 1913. Oligochètes. Catalogue des invertébrés de la Suisse. Georg.
& Cie, Genève, 215 pp.
POKORNI, B. 1994. Surveillance des eaux de surface. Rapport annuel du Département de la
gestion du territoire. Service cantonal de la protection de l'environnement, CH-2034
Peseux.
REYMOND, O. 1994. Préparations microscopiques permanentes d'oligochètes: une méthode
simple. Bull. Soc. Vaud. Sc. Nat., 83: 1-3.
SOLLBERGER, H. 1974. Le Lac de Neuchatel (Suisse). Ses eaux, ses sédiments, ses courants
sous-lacustres. Thèse Univ. Neuchâtel. 415 pp.
REVUE SUISSE DE ZOOLOGIE, 102 (4) : 913-942; décembre 1995
Oribatids from Brunei I (Acari: Oribatida).
New and interesting mites from the Geneva Museum LXXV
Sandor MAHUNKA
Zoological Department, Hungarian Natural History Museum, Baross utca 13,
H-1088 Budapest, Hungary.
Oribatids from Brunei I (Acari: Oribatida). New and interesting mites
from the Geneva Museum LXXV. - Eleven species from Brunei are dis-
cussed; nine of them are new to science, one of them representing a new
genus: Teraja gen. n. (Microzetidae). Extended description of Temburongia
Mahunka, 1990 is given. The following new combinations are proposed:
Sellnickochthonius planus (Chinone) comb.n = Brachychthonius planus
Chinone, 1974; Teraja fimbriata (Mahunka) comb.n. = Microzetes fim-
briatus Mahunka, 1989.
Key-words: Acari - Oribatida - Taxonomy - New species, new genera -
Brunei.
INTRODUCTION
Oribatids are a major group of soil microarthropods with a high degree of
diversity. Over 6000 described species in more than 1000 genera have been described
(BALOGH & BALOGH 1992). They occur in many habitats, occupying a variety of
niches, and their widespread distribution affords great scope for biogeographical
studies (e.g. BERNINI, AVANZATI & BERNINI 1988; SUBIAS, ARILLO & GIL-MARTIN
1992; MAHUNKA 1993a).
Oribatida is an ancient group (150 - 200 million years) and the global distri-
bution patterns of the species need to be interpreted within the context of continental
drift. The oribatid fauna of South-East Asia is partly of Gondwanan and partly of
Laurasian origin which makes zoogeographical interpretations somewhat problematic
(HAMMER & WALLWORK 1979). A more thorough knowledge of the fauna of Borneo
is critical for a clearer understanding of these matters.
Few data have been available for the North Bornean territories Sabah and
Brunei (e.g. AOKI 1967; RAMSAY & SHEALS 1969), and none at all for Sarawak, until,
in 1982. Dr. Bernd Hauser, Head of the Arthropod Department of the Museum
d'Histoire naturelle, Geneva, began a systematic exploration of the soil fauna,
Manuscript accepted 30.09.1995.
914 SANDOR MAHUNKA
especially Microarthropoda, of the rain forests of South-East Asia and surrounding
areas. Subsequently, 121 species records for Sabahn oribatids were published from his
material, 94 of which were new to science (MAHUNKA 1987a, b, 1988, 1991, 1994b).
Other published results on oribatids from this research programm have been those
from Singapore (MAHUNKA 1989a), Sumatra (MAHUNKA 1989b) and Madagascar
(MAHUNKA 1993b, 1994a).
In the present paper!, oribatids collected from Brunei in 1988 by Dr. B. Hauser
during his 1988 expedition to Brunei, organized together with Dr. Charles Lienhard,
Research Officer at the same Department, are catalogued. Eleven species are listed,
nine of them new to science.
I would like to thank Dr. M. Luxton for critically reading of the manuscript
and for his valuable recommandations.
LIST OF LOCALITIES
Bru-88/12: Brunei (Brunei-Muara District): près du pont sur le ruisseau “Sungai Lubang
Barus” sur la route venant de Tutong, à 33 km de Bandar Seri Begawan, prélèvement de sol
dans les angles formés sur les contreforts de deux grands arbres proches des habitations, env.
20 m; 16.X1.1988; leg. B. H. (B)2
Bru-88/21: Brunei (Belait District): “Andulau Forest Reserve”, à 3,5 km au sud de Sungai
Liang (= à 39,5 km de Labi), forêt primaire (“Mixed dipterocarp forest”), K-7 (“Kompartment
7°), prélèvement de sol dans les angles formés par les contreforts de grands arbres, 50 m;
19.XI.1988; leg. B.H. (B)?
Bru-88/24: Brunei (Brunei-Muara District): “Berakas Forest Reserve” au nord de Bandar Seri
Begawan sur la route, à 19,5 km de Muara (= à 102,5 km de Kuala Belait), forêt “Kerangas”(=
“Tropical heath forest”), prélèvement de sol au pied de Casuarina nobilis Whitmore (Casuari-
naceae), 30 m; 20.X1.1988; leg. B. H. (B)?
Bru-88/29: Brunei (Belait District): Sungai Liang, “Arboretum Forest Reserve”, forét primaire
(“Mixed dipterocarp forest”), prélèvement de sol dans les angles formés par les contreforts de
deux arbres appelés “Nyatho”, 90 m; 21.X1.1988; leg. B.H. (B)?
Bru-88/32: Brunei (Belait District): “Labi Hills Forest Reserve”, “Teraja”, à 42 km au sud de
Sungai Liang (= 12 km au Sud de Labi), environs de “Rumah Panjang” (= Longhouse du Kam-
pong Teraja), forêt primaire (“Mixed dipterocarp forest”), prélèvement de sol dans les angles
formés par les contreforts d’un très grand arbre, 40 m; 22.X1.1988; leg. B.H. (B)?
Bru-88/34: Brunei (Belait District): “Badas Forest Reserve”, à env. 10 km sur la route secon-
daire qui bifurque, à 32 km de Kuala Belait, vers le sud, forét “Kerangas” (= “Tropical heath
forest”) formée presque exclusivement par Agathis dammara (Lambert) L.G. Rich. (Arau-
cariaceae), sur et sous écorces, 10 m; 23.X1.1988; leg. B. H.
Bru-88/35: Brunei (Belait District): “Badas Forest Reserve”, à env. 10 km sur la route secondaire
qui bifurque, à 32 km de Kuala Belait, vers le sud, forêt “Kerangas” (=“Tropical heath forest”)
formée presque exclusivement par Agathis dammara (Lambert) L.G. Rich. (Araucariaceae),
prélèvement de sol au pied de Agathis dammara, 10 m; 23.X1.1988; leg. B.H. (B}
Bru-88/38: Brunei (Temburong District): “Peradayan Forest Reserve” (="Bukit Patoi”), à 14,5
km de Bangar (= 2,5 km de Labu) forêt primaire (“Mixed dipterocarp forest”), prélèvement de sol
dans les angles formés par les contreforts de grands arbres morts, 80 m; 24.X1.1988; leg. B. H.
(BY
! Partly sponsored by the Hungarian National Scientific Research Fund (OTKA 17629).
2 (B) = extraction par appareil Berlese à Bandar Seri Begawan (Brunei).
3 (B) = extraction par appareil Berlese à Hong Kong.
ORIBATIDS FROM BRUNEI 915
LIST OF IDENTIFIED SPECIES
Brachychthoniidae Thor, 1934
Sellnickochthonius muara sp. n.
Localities: Bru-88/12; Bru-88/24.
Sellnickochthonius planus (Chinone, 1974) comb. n.
Locality: Bru-88/24: 2 specimens.
Phthiracaridae Perty, 1841
Hoplophthiracarus (Plonaphacarus) aculeatus sp. n.
Locality: Bru-88/38.
Notophthiracarus hauseri sp. n.
Locality: Bru-88/21.
Temburongiidae Mahunka, 1990
Temburongia patoi Mahunka, 1990
Localities: Bru-88/21; Bru-88/38.
Sabahtritiidae Mahunka, 1988
Sabahtritia lienhardi sp. n.
Locality: Bru-88/29.
Eremaeozetidae Balogh, 1972
Eremaeozetes maculosus sp. n.
Locality: Bru-88/24.
Microzetidae Grandjean, 1936
Teraja wongi gen. n., sp. n.
Locality: Bru-88/32
Carabodidae C. L. Koch, 1837
Hardybodes flabellatus sp. n.
Locality: Bru-88/21; Bru-88/32; Bru-88/38.
Hardybodes penicillatus sp. n.
Locality: Bru-88/35.
Pasocepheus eremaeozetoides sp. n.
Locality: Bru-88/32
DESCRIPTIONS AND IDENTIFICATIONS
Sellnickochthonius muara sp. n. (Figs 1-4)
Material examined: Holotype: Bru-88/24, 2 paratypes from the same
sample; 3 paratypes: Bru-88/12. Holotype and 3 paratypes: MHNG4, 2 paratypes
(1364-PO-89): HNHMS.
Measurements. - Length: 140-148 um; width: 70-75 um.
Prodorsum: Rostrum conical, rostral teeth not visible in dorsal view.
Prodorsal surface well sclerotized, ornamented by conspicuously outlined areolar
patches, typical for the genus, as shown in Fig. |. Prodorsal setae phylliform, essen-
tially subequal. Interlamellar setae arising far from each other, near to the bothridium.
Head of sensillus (Fig. 2) symmetrical, covered with large spines arranged in longi-
tudinal rows.
4MHNG = deposited in the Muséum d’histoire naturelle, Genève.
SHNHM = deposited in the Hungarian Natural History Museum, Budapest, with identi-
fication number of the specimens in the Collection of Arachnida.
916 SANDOR MAHUNKA
Fics 1-4
Sellnickochthonius muara sp. n. - 1: body in dorsal view, 2: sensillus, 3: body in ventral view,
4: body in lateral view.
Notogaster: Border of notogastral shield Na partly reduced or absent
(Fig. 1). In the midline of shield Na the first, second and fifth pair of patches com-
pletely fused; borders of some fields sinuous. Border of ring-spot strong, also sinuous.
Ventral region: Formand position of ventral setae as shown in Fig. 3.
Remarks: The new species belongs to the “zelawaiensis” - group [con-
taining e. g. also S. zelawaiensis (Sellnick, 1928), S. foliatus (Hammer, 1958), S. elsos-
neadensis (Hammer, 1958), S. hauserorum (Mahunka, 1979), S. elisabethae Mahunka,
1973 and S. griseus (Hammer, 1958). The group is characterized by the foliate
notogastral setae, the symmetrical sensillus and the medially partly fused patches on the
notogastral shield Na.
The species is named after the city Muara.
ORIBATIDS FROM BRUNEI 917
Sellnickochthonius planus (Chinone, 1974) comb.n. (Figs 5-8)
= Brachychthonius planus Chinone, 1974.
Material examined: Bru-88/24: 2 specimens.
The species was described by CHINONE from Japan. The newly collected
specimens entirely correspond to the description and figures given by CHINONE
(1974). This species is particularly characterized by the basally curved prodorsal and
dorsal setae (Figs 5-8).
On the basis of the form of adanal setae this species belongs to the genus
Sellnickochthonius Krivolutzky, 1964.
Measurements. - Length: 155-157 um; width: 85-89 um.
Fics 5-8
Sellnickochthonius planus (Chinone, 1974) - 5: body in dorsal view, 6: sensillus, 7: body in
ventral view, 8: body in lateral view.
918 SANDOR MAHUNKA
Hoplophthiracarus (Plonaphacarus) aculeatus sp. n. (Figs 9-15)
Material examined: Holotype: Bru-88/38; 4 paratypes from the
same sample. Holotype and 2 paratypes: MHNG, 2 paratypes (1365-PO-89): HNHM.
Measurements:- Length of aspis: 290-335 um; length of notogaster:
526-729 um; height of notogaster: 364-567 um.
Aspis: Dorsal outline gradually convex, but rostrum clearly incised in
lateral view. Median crista absent, lateral carina short, not reaching beyond the sinus
line and not directed to the lateral rim (Fig. 12). Dorsal surface ornamented by weak
alveoli, but near to the posterior margin some stronger ones exist (Fig. 11). All
prodorsal setae - excepting seta ex-thick and finely ciliate.
Notogaster (Fig. 9): Surface always covered by cerotegument with
debris. Cuticle ornamented by alveoli. Forty (!) pairs of mostly straight and erect
notogastral setae present. Setae cj slightly longer than the others, but no essential
difference existing between them. The lyriffisures and the alveoli of the vestigial setae
(f) hardly observable, but two lyriffisures (ia, im) and the alveoli of setae fj visible.
Anogenital region: The position of the genital setae (Fig. 10) is
typical for the subgenus Plonaphacarus. All four pairs of setae on the ano-adanal
plates roughened or finely ciliate. Setae ad; somewhat shorter than the anal ones.
Legs: Seta d on femur I hooked (Fig. 13). Seta / missing on tarsus I (Fig.
14). The setal fomulae are:
I: 1-4-2+2-5+1-16+3-1
IV: 2-1-1-2+1-10-1 (Fig. 15).
Remarks: The large number of notogastral setae is unique in this family.
Considering the other features, the new species is correctly placed in the genus Hoplo-
phthiracarus Jacot, 1933, and therein in the subgenus Plonaphacarus Niedbala, 1986.
The species is named after the form of the notogastral setae.
Notophthiracarus hauseri sp. n. (Figs 16-24)
Material examined: Holotype: Bru-88/21: 5 paratypes from the
same sample. Holotype and 3 paratypes: MHNG, 2 paratypes (1366-PO-89): HNHM.
Measurements. - Length of aspis: 162-221 um; length of notogaster:
315-433 um; height of notogaster: 207-261 um.
Aspis: Its outline angulate in lateral view (Fig. 16), median crista strong,
distinctly protruding from the surface; a weaker lateral crista also present. Anterior
surface areolate, aspis basally ornamented by longitudinal rugae amongst which are
oblong spots (Fig. 19). Lateral carina absent, lateral rim very short, sinus-line distinct.
All prodorsal setae (excepting setae ex) phylliform. Sensillus long, thin, approxi-
mately setiform, distinctly spinose; spines arranged in two longitudinal rows.
Notogaster: Anterior part of notogaster very wide, protruding forward, in
dorsal view covering the basal part of aspis (Fig. 18). Surface of notogaster with large
protuberances and three longitudinal ribs gradually disappearing anteriorly (Fig. 18).
Fifteen pairs of phylliform notogastral setae present, their distal end sharply pointed.
ORIBATIDS FROM BRUNEI 919
Fics 9-12
Hoplophthiracarus (Plonaphacarus) aculeatus sp. n. - 9: body in lateral viw, 10: anogenital
region, 11: aspis in dorsal view, 12: aspis in lateral view.
920 SANDOR MAHUNKA
Fics 13-15
Hoplophthiracarus (Plonaphacarus) aculeatus sp. n. - 13: basal articles of leg I, 14: tarsus of
leg I, 15: leg IV.
Gnathosoma: Chelicera and palp as shown in Figs 17, 24. Palp three-
segmented, its setal formula: 2-2-8+1.
Anogenital region (Fig. 22): Genital plate hollowed laterally, its
surface areolate. Posterior five pairs of genital setae much longer than the others, the
anterior 2-3 pairs minute, hardly discernible (Fig. 23). Among the setae of the ano-
adanal plates 4 pairs are reduced so much as to be represented only by their alveoli,
only setae ad3 visible.
Legs: The legs chaetotaxy is of the “complete type” showing the subse-
quent setal formulae:
I: 1-4-2+2-5+1-16+3-1 (Fig. 21)
II: 1-3-2+1-3+1-12+2-1
III: 2-2-1+1-2+1-10-1
IV: 2-1-1-2+1-10-1 (Fig. 20)
Remarks: The new species corresponds well with the genus Notophthira-
carus Ramsay, 1968, and belongs to a species group characterized by the phylliform
notogastral setae [(e.g. N. multituberculatus (Balogh & Mahunka, 1966)].
921
ORIBATIDS FROM BRUNEI
Fics 16-19
Notophthiracarus hauseri sp. n. - 16: body in lateral view, 17: chelicera, 18: notogaster in
dorsal view, 19: aspis in dorsal view.
922 SANDOR MAHUNKA
Fics 20-24
Notophthiracarus hauseri sp. n. - 20: leg IV, 21: leg I, 22: anogenital region, 23: genital plate,
24: palp.
ORIBATIDS FROM BRUNEI 923
I dedicate the new species to my friend Dr. Bernd Hauser, the collector of this
very important and interesting material.
Temburongia Mahunka, 1990
Diagnosis®: Aspis with two pairs of median (!) and one pair of lateral
(divided) cristae. One lateral carina and a sinus-line on both sides long, well de-
veloped. All three pairs of median notogastral setae modified, setae ro originating far
from the rostrum. Notogastral shield much extended ventrally and passing between
the genito-aggenital and ano-adanal plates, dividing them on the surface. The two
ventral lobes of the notogaster independent (not fused), but touching medially.
Fourteen pairs of notogastral setae present, the vestigial setae (f,) are behind setae hy),
neither lyriffisures nor opening of glandules observable. Genital and aggenital plates,
like anal and adanal plates, completely fused with each other. The genito-aggenital
plates also fused basally with each other, only a short division between them ante-
riorly, the narrowed basal part arched and forming a deep sinus with the similar part
of the ano-adanal plates inside the body, covered by the ventral lobes of the noto-
gaster. Seven pairs of genital (aggenital) setae present. Ano-adanal plates forming a
semicircle, six pairs of setae arising on them, all arranged in longitudinal rows along
the inner margin of the plates. Chelicerae well developed, both the fixed and the
movable digits are dentate. Palp four-segmented. Legs: monodactylous. Setae on
femur I display characteristic reduction, otherwise, the legs belong to a characteristic
chaetotaxy group.
Type species: Temburongia patoi Mahunka, 1990.
Remarks: On the basis of the above features the new taxon is so far from
all distinct groups of the superfamily Euphthiracaroidea, that it could not be acco-
modated in any of the heretofore known families; therefore, a new family (Tem-
burongiidae Mahunka, 1990) was established. It represents a so far unknown line of
evolution within the “phtiracaroid” oribatids shedding some light on how the “ventral
plate” of the higher oribatids could have developed.
The genus is named after the Temburong District.
Temburongia patoi Mahunka, 1990 (Figs 25-36)
Material examined: Holotype: Bru-88/21, 19 paratypes from the
same sample; 16 paratypes: Bru-88/38. Holotype and 22 paratypes: MHNG, 13
paratypes (1367-PO-89): HNHM.
Measurements. - Length of aspis: 182-276 um: length of notogaster:
276-448 um; height of notogaster: 197-335 um.
Aspis: Its outline concave medially and strongly convex anteriorly. Whole
surface finely granulate. Two pairs of median cristae long, the third, outer one is
6 This description completes a preliminary one in MAHUNKA (1990).
924 SANDOR MAHUNKA
Fics 25-28
Temburongia patoi Mahunka, 1990 - 25: body in lateral view, 26: aspis in dorsal view, 27:
aspis in lateral view, 28: chelicera.
ORIBATIDS FROM BRUNEI 925
divided into two arched parts (Fig. 26). Lateral carina long, running along lateral
margin anteriorly, instead of the lateral rim. This lateral rim very short, hardly obser-
vable; sinus-line very long and strong. Three pairs of strong, sabre-shaped, distinctly
spiculate and one pair (ex) of thin, simple, prodorsal setae present. Sensillus long,
stick-shaped, only slightly arched (Fig. 27). Bothridial squama large, approximately
guttiform.
Notogaster: Whole surface covered by a cerotegument layer mostly
forming granules imitating alveoli. Collar line not observable. Fourteen pairs of
Fics 29-32
Temburongia patoi Mahunka, 1990 - 29: anogenital region, 30: genital plates, 31: anterior
margin of genital plate, 32: coxisternal region.
926 SANDOR MAHUNKA
Fics 33-36
Temburongia patoi Mahunka, 1990 - 33: tarsus of leg I, 34: basal articles of leg I, 35: palp, 36:
leg III.
ORIBATIDS FROM BRUNEI 927
notogastral setae present, all highly similar, but setae p1, p2, and especially p3, shorter
than the others. All setae phylliform, with median vein, their surface and margin
smooth. Lyrifissures ia and im present, other lyrifissures absent (?). Ventral lobes of
notogaster approximate medially, ending very near to each other (Fig. 29), their
margin slightly emarginate medially.
Gnathosoma: Chelicera as shown in Fig. 28. Palpal setal formula: 0-1-
2-9+1.
Epimeral region (Fig. 32): Epimeral setal formula: 3-0-1-2, but one
pair of setae on epimere 1 (perhaps /a) reduced, represented only by alveoli, and far
removed posteriorly from the other two setae.
Anogenital region: Genital plates fused basally, but anteriorly
separated from each other, in front of a sacculiform structure (Fig. 30). The narrowed
posterior part of genito-aggenital plates arched and delving into deeper layers of the
body and not directly connected with the similarly arched anterior part of ano-adanal
plages (Fig. 29). Seven pairs of genital setae present, one of them smooth and arising
on the anterior border of genital plates (Fig. 31); four pairs of these conspicuously
ciliate and originating in a more or less transverse furrow, two pairs arising near to the
sacculiform structure. Six pairs of identical ano-adanal setae observable, all six
arranged in a longitudinal row; all short, thin and simple.
Legs: Claw of legs thick but simple, without ventral teeth. Setation of legs
characteristic, and different from all related taxa. Setal formulae:
I: 1-2(!)-5+2-5+1-17+3-1
If: 1-3-4+1-5+1-12+2-1
IH: 2-2-3+1-3+1-11-1
IV: 2-2-2-3+1-11-1
On tarsus I a single seta arises near the basis of w (Fig. 33). Setae p’ and p" on tarsi
II (Fig. 36) and IV are of different type.
Remarks: The species is named after the hill of Bukit Patoi.
Sabahtritia lienhardi sp. n. (Figs 37-43)
Material examined: Holotype: Bru-88/29, 23 paratypes from the
same sample. Holotype and 15 paratypes: MHNG, 10 paratypes (1368-PO-89): HNHM.
Measurements. - Length of aspis: 133-169 um; length of notogaster:
194-266 um; height of notogaster: 97-163 um.
A spis: Its outline gradually convex in lateral view (Fig. 37). Surface
ornamented by a conspicuous network dorsally and by irregular wrinkles basally; a
narrow lateral part smooth. Lateral rim inconspicuous, a weak, short lateral carina and a
sinus line on each side observable. Median prodorsal setae slightly dilated, with a thin
velum. Sensillus setiform, long. Setae in the shortest of all prodorsal setae. Bothridium
small, but protruding conspicuously from the outline in dorsal view (Fig. 40).
928 SANDOR MAHUNKA
<
NE yar
Ua
sy. in
|
Fics 37-40
Sabahtritia lienhardi sp. n. - 37: body in lateral view, 38: anogenital region, 39: sculpture of
notogaster, 40: aspis in dorsal view.
Notogaster: Cuticle foveolate, cerotegument layer consisting of small
tubercles arranged so as to form a rugose, almost reticulate, pattern (Fig. 39) around
the foveolae. 14 pairs of phylliform notogastral setae present. Four pairs of lyri-
fissures (ia, im, ih, ip) visible; no vestigial setae observable.
Anogenital region (Fig. 38): Anogenital plates typical for the genus,
ornamented by a polygonal reticulation. Seven pairs of genito-aggenital, 3 pairs of
adanal setae visible.
ORIBATIDS FROM BRUNEI 929
Fics 41-43
Sabahtritia lienhardi sp. n. - 41: leg I, 42: palp, 43: leg IV.
Gnathosoma: Chelicera normal, both digits equally strong. Palp (Fig.
42) three-segmented, setal formula: 2-2-7+1.
Legs: Chaetotaxy of the “complete type”, with following setal formulae:
I: 1-3-5+2-4+1-19+3-1 (Fig. 41)
If: 1-2-2+1-3+1-13+2-1
II: 2-1-0-(3+1)-12-1 (Fig. 43)
Remarks: S. lienhardi is the third species of this genus. It is clearly
distinguished from both known species by the shape of its notogastral setae (also
phylliform but much broader in S. hauseri) or the shape of ano-adanal setae (serrated,
very long in S. mirabilis).
I dedicate the new species to Dr. Charles Lienhard for his continous help in
correcting my manuscripts and for his collaboration with Dr. B. Hauser’s field
activity.
Eremaeozetes maculosus sp. n. (Figs 44-52)
Material examined: Holotype: Bru-88/24, deposited in the MHNG.
Measurements. - Length: 403 um: with: 216 um.
930 SANDOR MAHUNKA
Fics 44-48
Eremaeozetes maculosus sp. n. - 44: body in dorsal view, 45: body in ventral view, 46: genital
plate, 47: anal plate in lateral view, 48: palp.
Prodorsum: Surface (as the whole surface of the body) covered by
reticulate cerotegument, which mostly reflects the sculpture of the cuticular surface.
Lamellae wide, fused basally and covering the whole prodorsum, excepting the basal
region (Fig. 44). Lamellar setae dilated, originating on the bent distal end of lamellae.
Rostral setae (ro) horn-shaped’. Interlamellar and exobothridial setae well developed,
like small spines. Sensillus very large, flabellate, with spiculate surface.
7 Some setae (ro, ge, an and ad) of the body are robust. Il seems to me, that they only
consist of secretions or are covered by a thick secretion layer. When these “setae” were broken
during preparation, no “real” setae could be found inside.
ORIBATIDS FROM BRUNEI 931
Fics 49-52
Eremaeozetes maculosus sp. n. - 49: tarsus of leg II, 50: tibia of leg II, 51: leg. III, 52: leg. I.
Notogaster: Humeral process well developed, bent downwards to the
epimeral surface. Lenticulus round. Some irregular, partly fused smooth areas
medially, over them the cerotegument layer lost (Fig. 44), all other surfaces areolate.
Ten pairs of dilated, round notogastral setae, the lyrifissures im and the glandular
openings visible in dorsal view.
Ventral region: Epimeral setal formula: 3-1-2-2. All setae short,
spiniform, hardly observable because of the thick cerotegument. Genital (Fig. 46) and
anal plates also areolate and covered by cerotegument. Anogenital setal formula: 6-1-
2-3. The posterior pair of genital setae and all anal and adanal setae (ad), robust but
their form varying (Fig. 47).
Gnathosoma: Mentum foveolate, seta h bending backwards, originating
laterally. Palpal eupathidium acm arising on a large apophysis, palpal setal formula:
2-1-3-9+1 (Fig. 48).
Legs: All legs tridactylous, with strong heretodactyly. Tarsus of legs I-IV
(Fig. 49) with a triangular dorsal spur, e.g. leg III (Fig. 51). Solenidium +, originating
on a long apophysis, the latter giving rise to a smaller anteriorly directed apophysis on
932 SANDOR MAHUNKA
which 2 is situated. Seta ff" on tarsus I (Fig. 52) strongly and characteristically
curved, famulus (e) long, arising in front of ao.
Remarks: The new species is clearly characterized by the special
formation of the dorsal cerotegument and the form and position of the genital and anal
setae. On the basis of these characters the new species is quite distinct from all other
members of the genus.
Teraja gen. n.
Diagnosis: Family Microzetidae. Rostrum conical. Lamellae wide with
long, horn-shaped, outer, and, completely reduced, inner cuspis; their median borders
partly touching medially, excavated basally and at their basal ends connected by an
arched transversal band. Lamellar setae spiniform, the phylliform interlamellar seta
arising on the dorsal surface of the lamellae. Tutorium with a velum-like formation
anteriorly. Sensillus setiform, directed backwards. Pteromorphae small, triangular.
Epimeral setal formula 3-1-3-3. A strong x-shaped sejugal formaton present in front
of the genital aperture. Ano-adanal setal formula: 6-1-2-2. Lyrifissures iad very long.
All legs monodactylous.
Type species: Teraja wongi sp. n.
Remarks: The new taxon is related to Microzetes Berlese, 1913 and
Megazetes Balogh, 1959, but both are distinguished from the new genus by the lack
of the characteristic velum-like formation of the tutorium. Only Microzetes fimbriatus
Mahunka, 1989 shows the some velum-Ike formation of the tutorium and conse-
quently has to be transferred to the new genus: Teraja fimbriata (Mahunka, 1989)
comb.n.
The new genus is named after the village of Teraja.
Teraja wongi sp. n. (Figs 53-56)
Material examined: Holotype: Bru-88/32; 6 paratypes from the
sample. Holotype and 4 paratypes: MHNG, 2 paratypes (1370-PO-89): HNHM.
Measurements. - Length: 242-251 um; width: 197-202 um.
Prodorsum: _ Rostrum conical in dorsal, beak-shaped in lateral view.
Rostral setae long, distinctly pilose. Lamellae wide, covering prodorsum in dorsal
view. Outer lamellar apex very long, strongly curved inwards and always reaching
beyond lamellar seta (Fig. 55). Interlamellar setae fine and densely pilose. Sensillus
long, setiform, its outer margin distinctly pilose, excepting basal part. Distal end of
tutorium with some (3-4) digitiform extensions.
Notogaster: Pteromorphae (Fig. 56) small, approximately triangular,
anterior surface wrinkled, basally ornamented by some small tubercles. Posterior
outline incised medially and laterally; these hollows continue in a flat furrow, directed
medially. These parts are densely granulate (Fig. 53). Nine pairs of fine, short
notogastral setae present.
ORIBATIDS FROM BRUNEI 933
Fics 53-54
Teraja wongi gen. n., sp. n. - 53: body in dorsal view, 54: body in ventral view.
934 SANDOR MAHUNKA
Fics 55-56
Teraja wongi gen. n., sp. n. - 55: prodorsum in lateral view, 56: pteromorpha.
Ventral region (Fig. 54): Pedotecta 1, 2-3 and discidium with
transversal rugae, surface of the epimeres and the ventral plate smooth. Epimeral
setae fine, pilose. Epimeral region divided by a strong x-shaped band only. Genital
and aggenital setae pilose. Epimeral region divided by a strong x-spahed band only.
Genital and aggenital sera pilose, anal and adanal setae simple and very short.
Remarks: The new species stands very near to 7. fimbriata (Mahunka,
1989) comb. n.: it is distinguished by the shape of its lamellar cuspis (much shorter in
T. fimbriata) and the number of the digitiform extensions (9-10 in 7. fimbriata).
I dedicate the new species to Dr. K. M. Wong from the Forest Research
Centre, Sungai Liang in recognition of this great assistance to the Geneva Expedition.
Hardybodes flabellatus sp. n. (Figs 57-61)
Material examined: Holotype: Bru-88/38, 3 paratypes from the
same sample; 2 paratypes: Bru-88/32; 4 paratypes: Bru-88/21. Holotype and 6
paratypes: MHNG, 3 paratypes (1371-PO-89): HNHM.
Measurements. - Length: 266-319 um; width: 123-153 pm.
Prodorsum: Whole surface (as whole surface of the body) covered by a
granular layer of secretions. Rostrum very wide, its anterior margin nearly straight or
slightly convex medially. Rostral and lamellar setae phylliform, both pairs smooth,
the latter larger than the former. Rostral setae originating very far from each other,
near to the lamellar setae. Lamellae running parallel, thus prodorsum, approximately
ORIBATIDS FROM BRUNEI 935
Fics 57-61
Hardybodes flabellatus sp. n. - 57: body in dorsal view, 58: leg I, 59: body in ventral view, 60:
prodorsum in lateral view, 61: leg II.
936 SANDOR MAHUNKA
rectangular in dorsal view (Fig. 57). No sharp lamellar cuspides, anterior outer margin
of lamellae with some alveoli and with some weak rugae. Interlamellar region with
some spots. Basal part of prodorsum with a strong transversal rib. Interlamellar setae
lanceolate, originating on the lamellar surface, directed forwards, their surface
spiculate. Sensillus large, spatulate (Fig. 60), directed outwards, its surface distinctly
barbed.
Notogaster: No dorsosejugal suture observable medially. Median part
well framed and protruding from notogastral surface. In this part some pairs of
additional elevations present and on the posterior part some large light spots. Fifteen
pairs of notogastral setae present, all dilated distinctly spiculate, similar in shape to
the intelamellar setae, but all are much smaller than the latter. Two pairs of setae
arising in humeral, 4 pairs in posteromarginal, position (Fig. 57).
Coxisternal region: Epimeral borders and apodemes perceptible,
as are some irregular spots and ribs. All epimeral setae minute, but their alveoli readi-
ly observable. Epimeral setal formula: 3-1-3-3. Setae /a originating characteristically
in front of /b (Fig. 59).
Anogenital region: Shorter in longitudinal direction than the
epimeral region. Surface between the genital and anal aperture, and also lateral parts,
with some wrinkles. Genital and aggenital setae slightly pilose, adanal setae dilated
(similarly to the notogastral setae), anal setae minute. Setae ad3 originating very far
anteriorly from the anal aperture.
Legs: Femur of leg I and II areolate, seta /" phylliform. Seta d” on genu
also wide, dilated, their surface distinctly spiculate. Seta u on all legs modified, short,
no other spiniform seta on tarsi I and II (Figs 58, 61). Setal formulae:
I: 1-4-3+1-4+2-16+2-1
II: 1-4-3+1-2+1(!)-15+2-1
Femur III with thinner, femur IV with broader ventral edge, femur III dilated dorsally.
Legs IH and IV with no special characteristics.
Remarks: See the remarks after the next Hardybodes species.
Hardybodes penicillatus sp. n. (Figs 62-66)
Material examined: Holotype: Bru-88/35, 6 paratypes from the
same sample. Holotype and 3 paratypes: MHNG, 3 paratypes (1371-PO-89): HNHM.
Measurements. - Length: 295-379 um; width: 138-167 um.
Prodorsum: Rostrum wide, anterior margin approximately straight.
Prodorsum oblong in dorsal view, lamellae narrow. Outer margin of lamellae dis-
tinctly rugose. Prodorsal setae similar to the preceding species, but sensillus much
longer and thinner, distal end not spatulate (Fig. 65).
Notogaster: Ornamented by large tubercles, forming a polygonal network
(Fig. 62). Fifteen pairs of notogastral setae present, similar to the preceding species.
ORIBATIDS FROM BRUNEI 937
FIGS 62-66
Hardybodes penicillatus sp. n. - 62: body in dorsal view, 63: leg I, 64: body in ventral view, 65:
prodorsum in lateral view, 66: leg II.
938 SANDOR MAHUNKA
Coxisternal region (Fig. 64): Similar to H. flabellatus.
Anogenital region: Surface between the genital and anal apertures
and lateral parts with some round tubercles and short wrinkles. Genital and aggenital
setae plumose, adanal setae phylliform and spiculate, anal setae minute.
Legs: The form and the chaetotaxy of all legs (leg. I, II: figs 63, 66) are
exactly the same as those of Hardybodes flabellatus.
Remarks: These two new species were easy to place in the genus
Hardybodes Balogh, 1970, altough some corrections and additions to the generic
diagnosis are necessary. Consequently, the peculiar form of the rostrum and the
rostral setae is characteristic only for the type species (H. mirabilis Balogh, 1970).
The notogaster is not always flat, in the new species its median part distinctly
projects.
The three species may be separated by the following key:
I Rostral part of prodorsum elongated, rostral and lamellar setae T-shaped;
interlamellar region with a complex structure. . mirabilis Balogh, 1970
— Rostral part of prodorsum normal, wide, almost straight anteriorly. ...
à
2 Sensillus very wide, spatulate; notogaster without tubercles; genital
seraeswithisome short cilia: 2. PEER eas ee flabellatus sp. n.
_ Sensillus long, narrow, without a spatulate head; notogaster with tubercles;
Pemitallsetac plumMOse RE TE ee penicillatus sp. n.
Pasocepheus eremaeozetoides sp. n. (Figs 67-73)
Material examined: Holotype: Bru-88/32; 3 paratypes from the
same sample. Holotype and | paratype: MHNG, 2 paratypes (1373-PO-89): HNHM.
Measurements. - Length: 379-482 um; width: 261-335 um.
Prodorsum: Lamellae conspicuously protruding above the prodorsal
surface; their basal parts fused into a typical U-shaped formation in dorsal view (Fig.
67). Prodorsal surface coarsely wrinkled anteriorly and (in common with the rest of
the prodorsal surface) finely granulate basally. Lateral part of prodorsum and surface
between lamella and tutorium ornamented by some areolae. Basal part of prodorsum
with two depressions. Sensillus (Fig. 70) long, distinctly ciliate, directed outwards and
slightly backwards.
Notogaster: Dorsosejugal suture clearly visible. Median part of
notogaster distinctly protruding (Fig. 69); this part consists of two elevations, between
them there is a hollow divided by three transversal costulae, and from these elevations
three gradually decreasing cristae are directed forward. Whole surface ornamented,
anteriorly and anterolaterally polygonated, with symmetrically arranged tubercles
around the irregularly wrinkled median elevation. Posterolateral margin granulate.
Fifteen pairs of small, but slightly dilated, phylliform notogastral setae present, two
pairs of them arising from the humeral apophysis, four of them in a posteromarginal
position and the rest arising on the median elevation.
ORIBATIDS FROM BRUNEI 939
Fics 67-70
Pasocepheus eremaeozetoides sp. n. - 67: body in dorsal view, 68: body in ventral view, 69:
body in lateral view, 70: sensillus.
Ventral region: Mentum with a wide transverse crest, its surface
coarsely foveolate. Apodemes and epimeral borders short, with a wide median hollow
between the epimeral surfaces (Fig. 68). Three pairs of large tubercles present on
epimeres | and 4 in the sejugal region, being clearly visible also in lateral view.
Epimeral setal formula: 3-1-3-3, but setae 3a originating on fourth epimeres; all setae
minute. The distance between the median setae (/a-/a, 2a-2a, etc.) is great, due to the
940 SANDOR MAHUNKA
Fics 71-73
Pasocepheus eremaeozetoides sp. n. - 71: leg I, 72: lamella in lateral view, 73: leg II.
presence of the median hollow. Anogenital region framed by two strong, distinct
crests, the inner one running around the genital aperture and continuing to the anal
aperture, the other one running far laterally. Surface of the genital and anal apertures
wrinkled. Anogenital setal formula: 4-1-2-3. Adanal lyrifissures were not observed.
Adanal setae phylliform, all the other setae simple and minute.
Legs: All trochanters and femora pitted with areolae, the other segments
smooth. Chaetotaxy of all legs typical for the family, setae u’ on every tarsus bigger
than u”. The position of the setae of legs I and II is shown in Figs 71 and 73.
Remarks: On the basis of the characteristic structure of the body the new
species is easily classifiable into the until now monotypical genus Pasocepheus Aoki,
1976. According to the original description this genus is characterized by 13 pairs of
notogastral setae, but possibly (type specimens not seen) two pairs of setae in the
anterior part of notogaster were overlooked by AOKI. The new species is readily
ORIBATIDS FROM BRUNEI 94]
distinguished from the type species of Pasocepheus (P. triarcuatus Aoki, 1976) by
the form of its notogastral elevation (three elevations exist in P. triarcuatus) and by
its very large humeral apophysis (absent in P. triarcuatus).
REFERENCES
AOKI, J. 1967. A preliminary revision of the family Otocepheidae (Acari, Cryptostigmata) II
Subfamily Tetracondylinae. Bull. natn. Sci. Mus. Tokyo 10: 297-359.
AOKI, J. 1976. Oribatid mites from the IBP Study Area, Pasoh Forest Reserve, West Malaysia.
Nature Life S.E. Asia 7: 39-59.
BALOGH, J. 1970. New Oribatids (Acari) from New Guinea II. Acta zool. hung. 30: 291-344.
BALOGH, J. & P. BALOGH, 1992. The Oribatid mites genera of the world. Hungarian Natural
History Museum, Budapest, vol.1: 263 pp., vol. 2: 375 pp.
BALOGH, J. & S. MAHUNKA. 1966. New Oribatids (Acari) from South Africa. Acta zool. hung.
12: 1-23.
BERNINI, F., A. M. AVANZATI & S. BERNINI, 1988. Notulae Oribatologicae XXXVII. Gli Acari
Oribatei del Massiccio del Pollino (Italia Meridionale): aspetti faunistici e
biogeografici. Lav. Soc. ital. Biogeogr. N.S. 10 (1987): 379-488.
CHINONE, S. 1974. Further contribution to the knowledge of the family Brachychthoniidae from
Japan. Bull. biogeogr. Soc. Japan 30: 1-28.
GRANDJEAN, F. 1936. Les Microzetidae n. fam. (Oribates) Bull. Soc. zool. Fr. 61: 60-93.
HAMMER, M. 1958. Investigations on the Oribatid Fauna of the Andes Mountains I. The
Argentine and Bolivia. Biol. Skr. 10: 1-129.
HAMMER, M. & J. A. WALLWORK. 1979. A review of the world distribution of Oribatid Mites
(Acari: Cryptostigmata) in relation to Continental Drift. Biol. Skr. 22: 1-31.
MAHUNKA, S. 1973. Neue und interessante Milben aus dem Genfer Museum. XI. Neue und
wenig bekannte Oribatiden aus Rhodesien (Acari). Archs Sci. Genève 26: 205-225.
MAHUNKA, S. 1979. Neue und interessante Milben aus dem Genfer Museum XLI. Vierter
Beitrag zur Kenntnis der Oribatiden-Fauna Griechenlands (Acari: Oribatida). Revue
suisse Zool. 86: 541-571.
MAHUNKA, S. 1987a. Neue und interessante Milben aus dem Genfer Museum LV. Oribatids
from Sabah (East Malaysia) I (Acari: Oribatida). Archs Sci. Genéve 40: 293-305.
MAHUNKA, S. 1987b. Neue und interessante Milben aus dem Genfer Museum LX. Oribatids
from Sabah (East Malaysia) II. (Acari: Oribatida) Revue suisse Zool. 94: 765-817.
MAHUNKA, S. 1988. New and interesting mites from the Geneva Museum LXI. Oribatids from
Sabah (East Malaysia) III (Acari: Oribatida). Revue suisse Zool. 95: 817-888.
MAHUNKA, S. 1989a. New and interesting mites from the Geneva Museum LXIV. Oribatids
from Singapore (Acari: Oribatida). Revue suisse Zool. 96: 381-402.
MAHUNKA, S. 1989b. New and interesting mites from the Geneva Museum LXV. Oribatids of
Sumatra (Indonesia) I (Acari: Oribatida). Revue suisse Zool. 96: 673-696.
MAHUNKA, S. 1990. A survey of the superfamiliy Euphthiracaroidea Jacot, 1930 (Acari: Oriba-
tida). Folia ent. hung. 51: 37-80.
MAHUNKA, S. 1991. New and interesting mites from the Geneva Museum LXVIII. Oribatids
from Sabah (East Malaysia) IV (Acari: Oribatida). Revue suisse Zool. 98: 185-206.
MAHUNKA, S. 1993a. Hungaromotrichus baloghi gen. et sp. n. (Acari: Oribatida), and some
suggestions to the faunagenesis of the Carpathian Basin. Folia ent. hung. 54: 75-83.
MAHUNKA, S. 1993b. Oribatids from Madagascar I. (Acari: Oribatida) - New and interesting
mites from the Geneva Museum LXXVI. Revue suisse Zool. 100: 289-315.
MAHUNKA, S. 1994. Oribatids from Madagascar II. (Acari: Oribatida). (New and interesting
mites from the Geneva Museum LXXIX.) Revue suisse Zool. 101: 47-88.
942 SANDOR MAHUNKA
MAHUNKA, S. 1995. Oribatids from Sabah (East Malaysia) V (Acari: Oribatida: Parakalum-
moidea stat. n. and Galumnoidea). Trop. Zool. 8: 269-308.
NIEDBALA, W. 1986. Catalogue des Phthiracaroidea (Acari), clef pour la détermination des
espèces et descriptions d’espèces nouvelles. Annis zool. Warsz. 40: 309-370.
Ramsay, G. W. & J. G. SHEALS. 1969. Euphthiracaroid mites (Acari, Cryptostigmata) from
North Borneo. Bull. Br. Mus. nat. Hist. Zoology 18: 91-115.
SELLNICK, M. 1928. Formenkreis: Hornmilben, Oribatei. Jn: P. Brohmer, P. Ehrmann & G.
Ulmer (eds.) Tierwelt Mitteleur. 3 (IX): 1-42.
SUBIAS, L. S., A. ARILLO & J. GIL-MARTIN. 1992. Consideraciones biogeograficas sobre los
Oribatidos (Acari, Oribatida) de Marruecos y Sahara Occidental. Jn: A. Alemany (ed.):
Historia Natural ‘91 (Biologia de Organismos y Sistemas) p. 189-202.
REVUE SUISSE DE ZOOLOGIE, 102 (4) : 943-959; décembre 1995
Sur les Euthiini et Cephenniini (Coleoptera, Scydmaenidae)
de l'Extrèéme-Est de la Russie et du Japon
Serguei A. KURBATOV
Severodvinskaya 9-84, Moscou 129224, Russie.
On the Euthiini and Cephenniini (Coleoptera, Scydmaenidae) from
Russian Far East and Japan. - Following new species are described:
Veraphis fatiloquus n. sp. (South Primorje), Cephennodes vafer n. sp.
(South Kuriles), Chelonoidum pullatum n. sp. (South Kuriles, Shikoku),
Chelonoidum moderatum n. sp. (South Primorje), Chelonoidum besucheti
n. sp. (Shikoku), Chelonoidum loebli n. sp. (Shikoku) and Chelonoidum
torosum n. sp. (Honshu). Cephennodes graeseri Reitter is transferred to
Chelonoidum Strand. Keys to the Palaearctic Veraphis and East Palaearctic
Chelonoidum are given.
Key-words: Coleoptera - Scydmaenidae - Taxonomy - Far East of Russia -
Japan.
INTRODUCTION
Les Scydmaenidae sont des prédateurs des acariens répandus dans le monde
entier. Toutefois, ils sont encore peu étudiés, sauf en Europe. Les tribus des Euthiini
et Cephenniini, placées a la base de la famille, sont restées presque inconnues dans
l'est de la région paléarctique. Ainsi, cette étude basée sur les récoltes récentes faites
dans l'Extrême-Est de la Russie et au Japon, porte le nombre des espèces recensées de
3 à 4 pour les Veraphis et de 2 a 9 pour les Cephenniini. Les données disponibles sur
la diversité et la distribution du groupe reflètent actuellement plus l’activité des
récolteurs que la réalité biologique.
L'auteur remercie vivement Monsieur C. Besuchet qui lui a remis le matériel
du Muséum d'histoire naturelle de Genève (MHNG) et qui lui a permis d'utiliser les
données non-publiées relatives à Cephennodes japonicus Sharp. Le matériel typique
est déposé également au Muséum zoologique de l'Université de Moscou (MZUM) et
dans la collection de l'auteur (SK)
Manuscrit accepté le 22.12.1994.
944 SERGUEI A. KURBATOV
Veraphis Casey
Genre holarctique avec 7 espèces aux Etats-Unis (Casey, 1897) et 4 espèces de
la taïga paléarctique (FRANZ, 1971; HISAMATSU, 1985), toutes très rares dans les
collections. Les Veraphis montrent un dimorphisme sexuel secondaire bien accusé
affectant surtout la proportion des articles antennaires.
Veraphis fatiloquus n. sp.
Matériel. Holotype d: Russie: Primorsky Kray, à l'est d'Oussouriisk, environs de
Kamenouchka, Abies pourri, 22.VI.1990, leg. S. Kurbatov (MZUM).
Paratypes: Russie: Primorsky Kray, au sud de Spassk-Dalnij, environs de Merkou-
chevka, Pinus coreanus pourri, 18.VI.1990, 1 4 (SK); pente occidentale du mont Olkhovaya,
conifère pourri, 29.VI.1990, 1 d (SK); même localité, Abies pourri, 30.VI.1990, 1 2 (MZUM);
25 km au nord-ouest de Terney, Abies pourri, 3.VII.1992, 1 9 (SK); tous leg. S. Kurbatov.
Longueur 1,25-1,30 mm. Brun, coloration des élytres parfois un peu plus
claire.
Téte (yeux compris) large de 0,26 mm, nettement déprimée entre les yeux,
dépression prolongée en avant jusqu'au bord antérieur du front, progressivement plus
large et moins marquée; téguments lisses sauf sur la partie antérieure de la dépression,
pourvue d'une ponctuation floue; bord antérieur du front nettement anguleux; yeux
grands, peu saillants; tempes en vue dorsale aussi longues que les 2/3 du diamètre des
yeux; tubercules antennaires petits; antennes sans massue nette, s'élargissant
progressivement vers le sommet.
Pronotum long de 0,30-0,33 mm, large de 0,33-0,35 mm, bord basal légère-
ment et régulièrement convexe, bords latéraux faiblement élargis jusqu'au niveau du
quart antérieur puis arrondis vers le bord antérieur; pas de ponctuation; centre du
disque muni d'un sillon longitudinal très faible.
Elytres sur la suture longs de 0,69-0,74 mm, larges de 0,50-0,53 mm, à
ponctuation effacée, régulièrement arrondis sur les côtés, disque faiblement convexe,
non aplati.
Mâle. Article 1 de l'antenne au moins 2 fois plus long que large; article 2 un
peu plus étroit, allongé; 3 un peu transverse, plus étroit que 2; 4 de la méme largeur,
isodiamétrique; 5 isodiamétrique ou légèrement transverse, nettement plus large que
4; 6 a peine plus large que 5, presque isodiamétrique; 7 plus large que 6, faiblement
transverse; 8 encore plus large, faiblement transverse; 9 plus large que 8, faiblement
transverse; 10 plus large que 9, aussi faiblement transverse; 11 plus long que large, de
la méme largeur que l'article précédent, faiblement concave a l'extrémité de son còté
extérieur (Fig. 1). Métasternum a carène médiane prolongée du bord antérieur au
centre et munie sur toute sa longueur d'une rainure fine mais nette; cette carène est
suivie postérieurement par une large et faible dépression longitudinale. Edéage (Figs
3, 4) long de 0,24-0,25 mm.
Femelle. Articles 1 à 3 de l'antenne de même grandeur que chez le mâle; 3 et 4
égaux; 5 et 6 de la méme longueur, mais un peu plus larges; 7 encore plus large,
fortement transverse; 8 de la méme longueur, a peine plus étroit que 7, fortement
EUTHIINI ET CEPHENNIINI (COLEOPTERA, SCYDMAENIDAE) 945
Fics 1-4
Veraphis fatiloquus n. sp.; 1 : extrémité de l'antenne mâle; 2 : idem, antenne femelle; 3 et 4 :
édéage, face dorsale et latérale.
transverse; 9 bien plus large et long que 8, transverse; 10 transverse, plus volumineux
que 9; 11 aussi long que large, plus large que 10, muni d'une vaste échancrure dans la
moitié distale du côté extérieur (Fig. 2); carène métasternale longitudinale moins
accusée que chez le male, bien marquée seulement jusqu'au niveau du bord postérieur
des hanches 2, s'atténuant peu a peu; dépression longitudinale moins accusée que chez
le male.
Par la forme de l'édéage, V. fatiloquus est proche à V. engelmarki Franz. Il en
diffère par la structure de la partie apicale de l'édéage et par les proportions des
articles antennaires, ainsi que par le disque élytral non aplati. Veraphis ishikawai
Hisamatsu du Japon (Hokkaido) dont une seule femelle est connue, diffère de l'espèce
nouvelle par sa grande taille, par les proportions de l'article antennaire 6 et par le
pronotum aussi long que large.
946 SERGUEI A. KURBATOV
Quatre femelles, trouvées dans la méme région et déterminées comme
Veraphis irkutensis Reitter (Kurbatov, 1990), sont très semblables aux femelles de la
nouvelle espèce, mais plus petites (longues de 1,2 mm au maximum) et présentent la
dépression du vertex lisse sur sa partie antérieure.
CLE DES Veraphis PALEARCTIQUES
1 Males) oe ate a ip oc cc 60 0 000 = 2
— Femellest ho ont à A een SO IR ET, 4
2 Articles antennaires 4 à 7 plus longs que larges. Longueur 1,35-1,40
mmasuedeiboréale N 2 CA RENE ene engelmarki Franz
= Articles antennaires 4 a 7 isodiamétriques ou légèrement transverses....... 5
3) Articles antennaires 8 à 10 bien plus larges que longs. Longueur 1,0-1,1
mm. Massif Sayan oriental, Mongolie boréale ............ irkutensis Reitter
- Articles antennaires 8 à 10 à peine plus larges que longs. Longueur
1625153 Orme Primorskey Kray PE PE PMP ee eee fatiloquus n. sp.
4 Article antennaire 6 plus étroit que 5. Longueur 1,8 mm. Hokkaido
ea celano PAR NC pela ie eae arate gar ishikawai Hisamatsu
- Article antennaire 6 au moins aussi large que 5. Longueur maximale 1,4 mm. . 5
5) Articles antennaires 7 à 10 au moins 3 fois plus larges que longs.
Horo WELT RC PRET CR RP deo irkutensis Reitter
— Articles antennaires 7 à 10 au maximum 2 fois plus larges que longs ....... 6
6 Articles antennaires 3 à 5 isodiamétriques. Longueur 1,35-1,4 mm
et mere ee ee ee coh Ua engelmarki Franz
— Articles antennaires 3 à 5 légèrement transverses. Longueur 1,25-1,3 mm
aa agita agi o SER Tal MO RES ao fatiloquus n. sp.
Cephennodes Reitter
Ce genre est caractérisé par la présence d'une carène parallèle au bord latéral
du pronotum et par la structure de l'édéage plat aux paramères asymétriques. 16
espèces de l'Asie méridionale et orientale, de l'Afrique orientale et des îles Hawaii.
Cephennodes vafer n. sp.
Matériel. Holotype, d: Russie, Kuriles du Sud, île de Kounachir, environs de Tretja-
Kovo, litière près d'un ruisseau, 2-5.VII.1991, leg. S. Kurbatov (MZUM).
Paratypes: Russie: Kuriles du Sud, comme l’holotype, 18 ex. et même données mais du
18-21.VII.1990, 26 ex. (MZUM, MHNG et SK).
Longueur 0,95-1,05 mm. Coloration brun rougeatre, pubescence dense, longue,
oblique, jaunâtre. Front aplati, à ponctuation très fine, reste de la tête non ponctué;
tubercules antennaires nets, séparés du front par les dépressions longitudinales légères;
la partie centrale de la tête convexe longitudinalement; yeux bien accusés, en vue
EUTHIINI ET CEPHENNIINI (COLEOPTERA, SCYDMAENIDAE) 947
dorsale bien plus longs que l'article antennaire 1; ce dernier plus long que large; 2
aussi plus long que large, plus petit que 1; 3 a 7 de méme largeur, un peu plus étroits
que 2; 3 a 4 aussi longs que larges; 5 a 6 a peine plus longs que les précédents; 7
encore plus long; article 8 plus large que 7, à peu près isodiametrique; massue formée
par les articles 9 a 11, comprimée; 9 en vue latérale a peine plus long que large,
nettement plus large que 8; 10 un peu plus large que long, environ 1,5 fois plus large
que 9; 11 environ 1/5 plus large que 10, 1,7 fois plus long que large, a peine plus long
que 9 et 10 réunis; en vue dorsale 9 plus long que large, 10 également plus long que
large, 11 a peine plus large que 10, presque 2 fois plus long que large.
Fics 5-7
Cephennodes vafer n. sp.; 5 et 6 : édéage, face dorsale et latérale. Cephennodes japonicus
(Sharp); 7 : édéage, face dorsale.
948 SERGUEI A. KURBATOV
Pronotum long de 0,35-0,36 mm, large de 0,48-0,51 mm, à points petits, ronds
et denses, à proximité des angles antérieurs plus fortement ponctué; côtés du
pronotum très légèrement élargis des angles basaux jusqu'un peu en avant du milieu,
puis arrondis vers le bord antérieur; carène latérale faiblement visible, étendue
presque de l'angle postérieur au tiers antérieur du pronotum; fossette latérale aussi
éloignée du bord latéral que du bord postérieur.
Elytres longs de 0,59-0,67 mm sur la suture, larges de 0,50-0,54 mm, leur plus
grande largeur située au quart antérieur; disque légèrement déprimé dans sa partie
antérieure; ponctuation nettement plus grosse que celle du pronotum; carène humérale
très courte, plus courte que l'article antennaire 7; carène subhumérale dépassant le
niveau du quart antérieur des élytres et aussi éloignée de la carène humérale que du
centre de la fossette basale.
Brachyptère, aile un peu plus courte et 4 fois plus étroite que l'élytre; méta-
sternum régulièrement convexe, finement ponctué; apophyse métasternale antérieure
prolongée en arrière par deux carènes courtes et divergentes, formant un angle de 60°
environ.
Pas de caractères sexuels secondaires visibles. Edéage (Figs 5, 6) long de 0,20-
0,22 mm.
La seule espèce paléarctique orientale connue de ce genre, C. Japonicus
(Sharp, 1886) de Nagasaki, diffère par les caractères suivants : taille plus grande (lon-
gueur 1,25 mm), ponctuation élytrale nulle, pronotum ponctué dans sa partie centrale
seulement, par des points elliptiques, orientés longitudinalement. Edéage (Fig. 7) long
de 0,17-0,18 mm, plus petit et de forme distincte (Besuchet, com. pers.).
Chelonoidum Strand
Diffère du genre précédent par l'absence de la carène latérale du pronotum et
par l'édéage à capsule basale volumineuse et aux paramères symétriques ou légère-
ment asymétriques. Une douzaine d'espèces holarctiques.
Chelonoidum graeseri (Reitter), comb. nov.
Cephennodes graeseri REITTER, 1887:270.
Cette espèce, d’un brun rougeätre, présente un pronotum dépourvu de carène
latérale et l'édéage typique des vrais Chelonoidum (Figs. 8, 9). Macroptère, aile 2 fois
plus longue et environ 1,5 fois plus large que l'élytre. Répandue en Extréme-Est de la
Russie, du Territoire Autonome Juif a l'extréme sud du Primorsky Kray. Vit dans la
litière forestière et dans le bois pourri, parfois en compagnie de Lasius spp.
Chelonoidum pullatum n. sp.
Matériel. Holotype d : Russie, Kuriles du Sud, île de Kounachir, environ de Tretjakovo,
litière près d'un ruisseau, 2-5.VII.1991, leg. S. Kurbatov (MZUM).
EUTHIINI ET CEPHENNIINI (COLEOPTERA, SCYDMAENIDAE) 949
Fics 8-9
Chelonoidum graeseri (Reitter); édéage, face dorsale et latérale.
Paratypes: Russie, Kuriles du Sud, méme données que l’holotype, 6 ex. (SK); méme
données mais du 19-21.VII.1990, 24 ex.(SK, MHNG, MZUM); environs de Mendeleevo,
10.VII.1985, 1 ex., leg. V. Belov (SK); Japon, Ehime, via Mt. Ishizuchi, 1000 m, 14.VIIL.1980,
1 ex., leg. I. Löbl (MHNG).
Longueur 1,25-1,30 mm. Trapu; coloration brun foncé a brun noiràtre, pubes-
cence dense, courte, couchée, a soies isolées plus longues et plus dressées près des
angles postérieurs du pronotum, sur les côtés et dans la moitié postérieure des élytres.
Front et vertex densément mais très finement ponctués; tubercules antennaires
nets; yeux bien développés, en vue dorsale nettement plus longs que l'article
antennaire 1; ce dernier a peine plus long que large; 2 environ 1,5 fois plus long que
large, plus étroit que l'article 1; 3 plus étroit que 2, plus long que large; articles 4 a 6
de même largeur que 3, mais moins longs, chacun plus long que large; 7 à peine plus
large et nettement plus long que 6; 8 plus large que le précédent, aussi long ou un peu
plus long que large; 9-11 comprimés, formant une massue pas très nette car la taille
950 SERGUEI A. KURBATOV
de 8 est intermédiaire entre celle de 7 et 9; 9 a peine plus long que large, 10 aussi long
que large, un peu plus large que 9; article 11 à peine plus large que 10, presque 2 fois
plus long que large, aussi long que 9 et 10 réunis.
Pronotum long de 0,46-0,49 mm, large de 0,65-0,70 mm, à ponctuation dense
et fine, plus forte sur les côtés; ceux-ci très légèrement élargis des angles basaux
jusqu'au milieu, puis arrondis vers le bord antérieur; fossette latérale aussi éloignée du
bord latéral que du bord postérieur.
Elytres longs de 0,76-0,79 mm sur la suture, larges de 0,72-0,76 mm, très
convexes, à largeur maximale située au niveau du tiers antérieur; ponctuation effacée;
carène humérale très courte, pas plus longue que l'article antennaire 2, carène
subhumérale dépassant le niveau du tiers antérieur des élytres, aussi éloignée de la
carène humérale que du centre de la fossette basale.
Fics 10-11
Chelonoidum pullatum n. sp.; édéage, face dorsale et face latérale.
EUTHIINI ET CEPHENNIINI (COLEOPTERA, SCYDMAENIDAE) 951
Macroptère, aile plus que 2 fois plus longue et 1,5 fois plus large que l'élytre;
métasternum régulièrement convexe, finement ponctué; apophyse métasternale anté-
rieure comme chez Cephennodes vafer.
Pas de caractères sexuels secondaires visibles. Edéage (Figs 10, 11) long de
0,33-0,34 mm.
Chelonoidum moderatum n. sp.
Matériel: holotype d: Russie, Kuriles du Sud, île de Kounachir, environs de Men-
deleevo, 15ème km, litière près d'un ruisseau, 3.VII.1991, leg. S. Kurbatov (MZUM).
Paratypes: Russie, Kuriles du Sud, méme données, 1 9; environ de Tretjakovo, litière
près d'un ruisseau, 20.VII.1990, 1 9, leg. S. Kurbatov (SK); Primorsky Kray, à l'est d'Oussou-
riisk, environs de Kamenouchka, Abies pourri, 2.VI.1990, 1 2, leg. S. Kurbatov (SK); environs
du cap Gamov, litière dans la chénaie, 30.VII.1988, 1 8,1 2, leg. S. Kurbatov (SK); réserve
"Kedrovaya pad", tilleul pourri, 2.VI.1991, 1 9, leg. S. Kurbatov (SK); Japon, Gunma, Onio-
shadashi Rock Garden, 21.VII.1980, 1 d, leg. I. Löbl (MHNG); Ehime, via Mt. Ishizuchi, 1000
m, 14.VIIL.1980, 2 8 et 3 2, leg. I. Löbl (MHNG, SK); Ishizuchi N. Park, Mt. Ishizuchi, 1550
m, 13.VIIL.1980, 1 d et 2 @, leg. I. Löbl (MHNG); Ishizuchi N. Park, Omogo, 900 m,
2.VII.1980, 1 d, leg. I. Löbl (SK); Gifu, 8 km SE Osaka, 750 m, 1.VIIL.1980, 4 à et 22 9,
leg. I. Löbl (MHNG).
Longueur 1,05-1,15 mm. Trapu, coloration brun rougeatre, pubescence courte,
couchée, a soies isolées plus longues et plus dressées dans la moitié basale du bord
latéral du pronotum, sur les côtés et sur la moitié postérieure des élytres.
Front a ponctuation dense et fine, vertex ponctué plus éparsement; tubercules
antennaires nets; article antennaire | un peu plus long que large, nettement moins
long que l'oeil en vue dorsale; 2 plus étroit que le prédécent, 1,5 fois plus long que
large; 3 a 6 de méme largeur, plus étroits que 2, chacun un peu plus long que large; 7
un peu plus large que 6, aussi long ou un peu plus court que 2; 8 un peu plus large que
7, aussi long ou légèrement plus long que large; 9 a 11 légèrement comprimés, en vue
latérale 9 aussi long que large, plus large que 8; 10 aussi long que large, un peu plus
grand que 9, en vue dorsale 9 et 10 à peine plus longs que larges; 11 a peine plus large
que 10, moins de 2 fois plus long que large, aussi long que 9 et 10 réunis.
Pronotum long de 0,38-0,43 mm, large de 0,54-0,58 mm; finement ponctué;
ponctuation sur les angles antérieurs nettement plus dense; côtés du pronotum
légèrement élargis des angles postérieurs jusqu'au milieu puis arrondis vers le bord
antérieur; fossette latérale aussi éloignée du bord latéral que du bord postérieur.
Elytres longs de 0,67-0,70 mm sur la suture, larges de 0,59-0,63 mm; très
convexes, leur largeur maximale située au niveau du tiers antérieur; ponctuation
effacée; carene humérale très courte, pas plus longue que l'article antennaire 2; carène
subhumérale atteignant plus ou moins le niveau du milieu de l'élytre, aussi éloignée
de la carène humérale que du centre de la fossette basale.
Macroptère, aile comme chez l'espèce précédente; métasternum régulièrement
convexe, finement ponctué; apophyse métasternale antérieure comme chez Cephen-
nodes vafer.
Pas de caractères sexuels secondaires visibles. Edéage (Figs 12, 13) long de
0,28-0,29 mm.
952 SERGUEI A. KURBATOV
12 13
Fics 12-13
Chelonoidum moderatum n. sp.; édéage, face dorsale et latérale.
Chelonoidum ussuricum n. sp.
Matériel. Holotype d: Russie, Primorsky Kray, réserve "Kedrovaya pad", bois
vermoulu et litière près d'un vieux tilleul, 2.VI.1991, leg. S. Kurbatov (MZUM).
Paratypes: Russie, Primorsky Kray, mêmes données, 1 © (SK); mêmes données mais
litière dans la forêt de la vallée, 24. VII.1987, 3 ex. (SK, MHNG); mêmes données mais litière,
26.VII.1987, 1 ex. (MHNG); mêmes données mais litière parmi les pierres, 1. VIII.1987, 1 ex.
(MHNG); mêmes données mais 8.IX.1964, 1 ex., leg. Dlusski (MHNG); environs du cap
Gamov, litière près d'un tilleul au bord d'un ruisseau desséché, 30.VII.1988, 1 ex., leg. S.
Kurbatov (SK); a l'ouest de Nezhino, sources de la rivière Malaya Ananjevka (anciennement
Malaya Eldouga), litière dans la forêt, 6.VIII.1988, 2 ex., leg. S. Kurbatov (SK); à l'est
d'Oussouriisk, environs de Kamenouchka, souche pourrie d'un feuillu, 3.VI.1989, 2 ex., leg. S.
Kurbatov (SK); méme localité, Abies pourri, 10.VI.1989, 7 ex., leg. S. Kurbatov (SK, MZUM);
mémes données mais tilleul pourri, 6.VI.1990, 7 ex., leg. S. Kurbatov (SK, MHNG, MZUM);
mémes données, tilleul pourri, 9.VI.1991, 1 ex. leg. S. Kurbatov (SK); mémes données, litière
dans la forêt de la vallée, 21.VI.1992, 2 ex., leg. S. Kurbatov); 60 km à l'ouest de Partisansk,
environs Anissimovka, forét de la vallée, litière près d'un peuplier, 21.VII.1988, 2 ex., leg. S.
Kurbatov (SK); pente occidentale du mont Olkhovaya, bois vermoulu et litière près d'un
peuplier, 29.VI.1990, 1 ex., leg. S. Kurbatov (MHNG).
EUTHIINI ET CEPHENNIINI (COLEOPTERA, SCYDMAENIDAE) 953
Longueur 1,0-1,10 mm. Coloration brun rougeätre clair, pubescence assez
courte, couchée, a soies isolées plus longues et plus dressées dans la moitié basale du
bord latéral du pronotum, sur les côtés et dans la moitié postérieure des élytres.
Front à ponctuation dense et fine, vertex lisse, non ponctué; tubercules
antennaires nets; yeux petits, à peine plus courts que l'article antennaire 1, celui-ci
nettement plus long que large; article 2 allongé, plus court et plus étroit que 1; 3 à 6
de même largeur, nettement plus étroits que 2; 3 un peu plus long que large; 3 à 6
progressivement raccourcis; 6 et 7 isodiamétriques, 7 plus grand que 6, plus long et
un peu plus large que 8 (en vue latérale); 8 plus large que long; 9 à 11 formant une
massue nette, 9 et 10 à peu près de même taille, à peine plus larges que longs, 11 de la
même largeur, un peu plus court que 9 et 10 réunis.
Fics 14-15
Chelonoidum ussuricum n. sp.; édéage, face dorsale et latérale.
954 SERGUEI A. KURBATOV
Pronotum long de 0,36-0,37 mm, large de 0,50-0,51 mm; ponctuation très
légère, presque invisible; côtés du pronotum faiblement élargis des angles postérieurs
jusqu'au milieu puis arrondis vers le bord antérieur; fossette latérale plus près du bord
postérieur que du bord latéral.
Elytres longs de 0,36-0,37 mm sur la suture, larges de 0,50-0,51 mm, assez
convexes; largeur maximale située un peu en arrière du tiers antérieur; ponctuation
effacée; carène humérale très courte, plus courte que l'article antennaire 2; carène
subhumérale atteignant le niveau du tiers antérieur de l'élytre, plus près de la carène
humérale que du centre de la fossette basale.
Brachyptère, aile de la même longueur que l'élytre; métasternum régulièrement
convexe, pratiquement sans aucune ponctuation; apophyse métasternale antérieure
comme chez Cephennodes vafer.
Pas de caractères sexuels secondaires visibles. Edéage (fig. 14, 15) long de
0,44-0,45 mm.
Chelonoidum besucheti n. sp.
Matériel. Holotype d : Japon, Ehime, Ishizuchi N. Park, Mt Ishizuchi, 13-14.VIII.1980,
leg. C. Besuchet (MHNG).
Longueur 1,15 mm. Coloration brun rougeätre, pubescence assez courte,
couchée.
Téte densément chagrinée, clypéus lisse; vertex un peu aplati entre les yeux;
tubercules antennaires nets; grands yeux, en vue dorsale aussi longs que l'article
antennaire 1; celui-ci à peine plus long que large; 2 a peine plus long que large, un
peu moins large que 1; 3 a 6 de méme taille, isodiamétriques, nettement plus étroits
que 2; 7 allongé, plus long et un peu plus large que 6, a peine plus long que 8; celui-ci
carré; 9-11 formant une massue nette, 9 et 10 isodiamétriques, 10 à peine plus grand
que 9; 11 plus large que 10, aussi long que 9 et 10 réunis.
Pronotum long de 0,37 mm, large de 0,56 mm, à ponctuation extrêmement
fine, presque invisible, plus forte près des angles antérieurs; côtés du pronotum
faiblement élargis des angles basaux jusqu'à peu près au milieu, puis arrondis vers le
bord antérieur; fossette latérale également éloignée des bords latéral et basal.
Elytres longs de 0,71 mm sur la suture, larges de 0,60 mm, assez convexes; à
points assez grands mais effacés; largeur maximale située au niveau du tiers antérieur;
carène humérale très courte, de la méme longueur que l'article antennaire 2; carène
subhumérale atteignant le niveau du tiers antérieur de l'élytre, située un peu plus près
de la carène humérale que du centre de la fossette basale.
Métasternum dépourvu de ponctuation, régulièrement convexe; apophyse
métasternale antérieure comme chez Cephennodes vafer. Ailes non examinées.
Pas de caractères sexuels secondaires visibles. Edéage (Figs 16, 17) long de
0,44 mm.
Chelonoidum loebli n. sp.
Matériel. Holotype d : Japon, Ehime, via Mt. Ishizuchi, 1000 m, 14.VIIL.1980, leg. I.
Löbl (MHNG).
Paratype: mémes données que l’holotype, 1 ex. (MHNG).
EUTHIINI ET CEPHENNIINI (COLEOPTERA, SCYDMAENIDAE) 955
Fics 16-17
Chelonoidum besucheti n. sp.; édéage, face dorsale et latérale.
Longueur 1,10 mm. Coloration brun rougeätre, pubescence oblique, plus
longue que chez l'espèce précédente.
Téte très finement ponctuée, clypéus lisse; vertex entre les yeux régulièrement
convexe; tubercules antennaires nets; yeux en vue dorsale aussi longs que la largeur
de l'article antennaire 1; celui-ci a peine plus long que large; article 2 allongé,
nettement moins large que 1; 3 à 6 subégaux, allongés, nettement plus étroits que 2; 7
nettement plus long et aussi large que 6 ou (en vue latérale) un peu plus large; article
8 a peine plus large et moins long que 7, a peine plus long que large; 9-11 formant
une massue nette, 9 et 10 isodiamétriques ou a peine plus longs que larges, 10 plus
grand que 9, 11 plus large que 10, aussi long que 9 et 10 réunis.
956 SERGUEI A. KURBATOV
Pronotum long de 0,39 mm, large de 0,57 mm, à ponctuation formée de points
minuscules, plus forte près des angles antérieurs; côtés du pronotum légèrement
élargis des angles basaux jusqu'au milieu, arrondis après vers le bord antérieur;
fossette latérale aussi éloignée du bord latéral que du bord basal.
Elytres longs de 0,69-0,71 mm sur la suture, larges de 0,61-0,63 mm,
convexes, à ponctuation effacée, largeur maximale située au niveau du tiers antérieur;
carène humérale plus courte que l'article antennaire 2; carène subhumérale atteignant
le niveau du tiers antérieur de l'élytre, également éloignée de la carène humérale et du
centre de la fossette basale.
Métasternum lisse, sans aucune ponctuation; apophyse métasternale antérieure
comme chez Cephennodes vafer. Ailes non examinées.
Pas de caractères sexuels secondaires visibles. Edéage (Figs 18, 19) long de
0,43 mm.
Fics 18-19
Chelonoidum loebli n. sp.; édéage, face dorsale et latérale.
EUTHIINI ET CEPHENNIINI (COLEOPTERA, SCYDMAENIDAE) 957
Chelonoidum torosum n. sp.
Matériel. Holotype d: Japon, Gunma, J.E. Kogen N Park, Shirane, 1750 m,
22.V11.1980, leg. I. Löbl (MHNG).
Longueur 1,3 mm. Coloration brun rougeätre, pubescence de longueur moyenne,
couchée, avec quelques soies isolées plus longues et plus dressées sur la moitie
postérieure des élytres.
Front à ponctuation très fine, effacée; clypéus et vertex lisses; tubercules
antennaires nets; en vue dorsale, oeil plus court que l'article antennaire 2; article 1 a
peine plus long que large; 2 allongé, nettement moins large que 2; 3 à 6 de même
taille, à peine plus longs que larges, un peu moins larges que 2; 7 plus long et aussi
large que le précédent ou, en vue latérale, à peine plus large; 8 nettement plus large
que 7, à peine plus long que large; 9-11 formant une massue pas très nettement
limitée, 9 nettement plus large que 8, isodiamétrique, 10 à peine plus large que long,
plus large que 9, 11 a peine plus large que 10, aussi long que 9 et 10 réunis (l'exem-
plaire unique est difforme: son antenne droite est raccourcie, composée de dix
articles, à cause de 9 et 10 non-séparés; tous les autres articles anormalement raccour-
cis; la description est basée sur l'antenne gauche).
m _
_
20 21
Fics 20-21
Chelonoidum torosum n. sp.; édéage, face dorsale et latérale.
958 SERGUEI A. KURBATOV
Pronotum long de 0,53 mm, large de 0,76 mm, volumineux, pratiquement sans
aucune ponctuation y compris sur les angles antérieurs; côtés du pronotum nettement
élargis des angles basaux jusqu'au milieu, puis arrondis vers le bord antérieur; fossette
latérale aussi éloignée du bord latéral que du bord basal.
Elytres longs de 0,75 mm sur la suture, larges de 0,77 mm (mais semblant
moins larges que le pronotum), convexes, à ponctuation effacée, leur largeur maxi-
male située avant le niveau du tiers antérieur; carène humérale très petite, de la même
longueur que l'article antennaire 2; carène subhumérale n'atteignant pas le niveau du
tiers antérieur de l'élytre, aussi éloignée de la carène humérale que du centre de la
fossette basale.
Métasternum à ponctuation indistincte; apophyse métasternale antérieure
comme chez Cephennodes vafer. Ailes non examinées.
Pas de caractères sexuels secondaires visibles. Edéage (Figs 20, 21) long de
0,36 mm, très semblable à celui de Chelonoidum moderatum, mais inversé.
CLE DES Chelonoidum DE LA REGION PALEARCTIQUE ORIENTALE
1 Pronotumia ponciuationifiime MAIS nette: i... 202 2 2
- Pronotum pratiquement sans aucune ponctuation ...................... 4
2 Articles antennaires 3 à 6 isodiametriques, 7 plus court que 6, 11 un peu
plus court que 9 et 10 réunis. Elytres a pubescence dense, longue et
oblique, composée de soies uniformément orientées. Carène sub-
humérale n'atteignant pas le niveau du tiers antérieur de l'élytre.
Longueur 1,5-1,6 mm. Sud de Khabarovsky Kray, Primorsky Kray
RARES ORC Re QUO SO A E BN graeseri (Reitter)
= Articles antennaires 3 à 6 nettement plus longs que larges, 7 plus long
que 6, 11 aussi long que 9 à 10 réunis. Elytres à pubescence assez
courte, couchée, munis d'une vingtaine de soies plus longues et plus
dressées sur les côtés et sur la moitié postérieure. Carène subhumérale
dépassant le niveau du tiers antérieur de l'élytre. Longueur ne dépassant
pas 1 mm. ide RR RITO, 3
3 Coloration brun noirâtre, longueur 1,25-1,30 mm. Kuriles du Sud,
Smkokuds porca LD eb eae eae EEE pullatum n. sp.
~ Coloration brun rougeatre, longueur 1,05-1,15 mm. Primorsky Kray,
Kumlésidu Sud, Honshu Sio moderatum N. sp.
4 Articles antennaires 4 a 6 et 8 isodiamétriques ou légérement trans-
verses, 7 un peu plus large que 8 (surtout en vue latérale)................ 5
Articles antennaires 4 à 6 et 8 plus longs que larges, 7 moins large que 8... . 6
5) Front a ponctuation dense et fine, vertex lisse, régulièrement convexe;
article antennaire 8 plus large que long, article 11 plus court que 9 et 10
réunis. Longueur 1,0-1,1 mm. Sud de Primorsky Kray ...... ussuricum n. Sp.
— Front et vertex densément chagrinés, vertex entre les yeux un peu
aplati; article antennaire 8 carré, 11 aussi long que 9 et 10 réunis.
ongueur 15 Smm Shikoku 2.2 2 2 eee besucheti n. sp.
EUTHIINI ET CEPHENNIINI (COLEOPTERA, SCYDMAENIDAE) 959
6 Elytres 1,8 fois plus longs et un peu plus larges que le pronotum; carène
subhumérale atteignant le niveau du tiers antérieur de l'élytre. Longueur
PPM TTI TKO KU EEE E RENNEN a loebli n. sp.
- Elytres 1,4 fois plus longs, apparemment plus étroits que le pronotum;
carene subhumérale dépassant a peine le niveau du quart antérieur de
helytres Eoneueur 153 mm Honshucentral 2255544 see torosum n. sp.
BIBLIOGRAPHIE
CASEY, T.L. 1897. Coleopterological Notices. VII. Annals N. Y. Acad. Sci. 9: 285-684.
FRANZ, H. 1971. Untersuchungen über die paläarktischen Arten der Gattungen Euthia Steph.
und Veraphis Casey (Col. Scydmaenidae). EOS 46: 57-84.
HISAMATSU, S. 1985. Notes on some Japanese Coleoptera, I. Trans. Shikoku Ent. Soc. 17: 5-13.
KURBATOV, S.A. 1990. More on the tribe Euthiini (Coleoptera, Scydmaenidae) in the USSR Far
East. Zool. Zhurnal, Moscow. 69: 136-140 (in Russian).
REITTER, E. 1887. Neue Coleopteren aus Europa, den angrenzenden Ländern und Sibirien, mit
Bemerkungen tiber bekannte Arten. III Theil. D. ent. Z. 31: 241-288.
SHARP, D. 1886. The Scydmaenidae of Japan. Ent. Monthly Mag. 23: 46-51.
fan fece mi fit
= | an Qu
REVUE SUISSE DE ZOOLOGIE, 102 (4) : 961-968; décembre 1995
The types of some species of Lathrobium Gravenhorst
and of Xantholinus sejugatus G. Benick
(Coleoptera: Staphylinidae)
Volker ASSING
Gabelsbergerstr. 2, D-30163 Hannover, Germany.
The types of some species of Lathrobium Gravenhorst and of Xantholinus
sejugatus G. Benick (Coleoptera: Staphylinidae). - An examination of the
types of Lathrobium microps L. Benick, L. patris G. Benick, L. dilutum
Erichson, L. longulum Gravenhorst, Xantholinus sejugatus G. Benick and X.
longiventris Heer yielded the following synonymies: Lathrobium dilutum Er.
= L. microps L. Benick syn. n., L. longulum Grav. = L. patris G. Benick syn.
n. and Xantholinus longiventris Heer = X. sejugatus G. Benick syn. n.
Lectotypes are designated for L. dilutum Er., L. longulum Grav. and X. longi-
ventris Heer. The male sexual characters of L. longulum Grav. and the
female terminalia of L. pallidum Nordm. and L. dilutum Er. are figured.
Key-words: Coleoptera - Staphylinidae - Lathrobium - Xantholinus -
Germany - Taxonomy.
INTRODUCTION
Among the numerous species of Central European Staphylinidae there are, not
only in the Aleocharinae, several taxa which never or only once or twice have been
recorded again since their description. The density of coleopterologists and their
collecting activity in this region are among the highest in the world. Such utmost
rarity is, therefore, remarkable and could have the following reasons: a) extinction, b)
strongly restricted areas of distribution (endemics) and/or special bionomics or life-
histories (e.g. subterraneous species or those with very short-lived adults), c)
confusion with similar, more common species (particularly in groups whose species
are difficult to identify) and d) the description is based on artefacts.
In the case of Lathrobium microps L. Benick, L. patris G. Benick and
Xantholinus sejugatus G. Benick no records exist after their description, with one
exception: HABERMANN (1983) reported L. patris from Estland. Even so, they were
considered and treated in major taxonomic studies on Paederinae and Xantholininae
(e.g. COIFFAIT 1972, 1982; LOHSE 1964), red data books (ZIEGLER et al. 1994) etc. It is
Manuscript accepted 30.01.1995.
962 VOLKER ASSING
rather unlikely that the reasons stated above under a), b) and c) apply to these species.
The island of Spiekeroog or the surroundings of Liibeck, their type localities, are
regions where no endemics are likely to be found. Furthermore, the externally and
presumably also ecologically most similar species (L. pallidum Nordm./L. dilutum Er.,
L. longulum Grav. and X. linearis (Ol.)/X. longiventris Heer/X. rhenanus Coiff.,
respectively) are widely distributed, mostly occur also in seminatural and agricultural
habitats and, except for the partially subterraneous L. pallidum and L. dilutum, are
epigeically active. Considering the relatively long tradition of pitfall trapping — L.
pallidum and L. dilutum, too, can be collected with pitfall traps — and other quantitative
and semiquantitative sampling methods in Northern Germany, it appeared doubtful that
such species should have been overlooked. In addition, the type locality of the two
species described by G. Benick, the Schellbruch near Liibeck, has repeatedly, yet
unsuccessfully, been visited by coleopterologists (Ziegler, pers. comm.).
Strikingly all of the three species in question have in common that their
descriptions are not only based on single specimens but they also lack references to
the genitalia, either because the type was erroneously believed to be a female (L.
patris, X. sejugatus) or because it was in fact a female (L. microps).
In order to solve the problem it was necessary to examine the types, which was
kindly made possible by Dr Ivan Löbl (Muséum d'histoire naturelle, Genève), who
arranged the loan of L. microps, and by Mr Heinrich Meybohm (Stelle) and Mr
Wolfgang Ziegler (Rondeshagen), who helped in locating and providing me with the
types of L. patris and X. sejugatus. Furthermore, I am indebted to Dr Manfred Uhlig
(Museum fiir Naturkunde, Berlin) for the loan of the types of L. dilutum and L.
longulum, and to Dr Bernhard Merz (Eidgenössische Technische Hochschule Zürich)
for those of X. longiventris.
All the types of the species described by L. and G. Benick and dealt with in
this paper are now deposited in the Muséum d'histoire naturelle, Genève.
Lathrobium patris G. Benick, 1949
Holotype d, labels: Lübeck, Schnellbruch, 12.X1.44, Uberschwemmung, Dr G. Benick leg.,
Lathrobium patris G. Ben., Typus ® [sic!].
BENICK (1949) and LOHSE (1964) state that L. patris is very similar to L.
longulum (together with which it was collected!), but can be separated from the latter
by the following characters: the more oblong and narrower head, the posteriorly
narrowed pronotum and the shorter elytra. BoRDONI (1980a) examined the type,
which turned out to be a d. Apparently Benick mistook it for a 2 because of the
shape of sternite VIII, whose hind margin is projecting (Fig. 2-4) and not incised, as
in most Lathrobium spp.. In his study of Lathrobiini BORDONI (1980a) figures the
aedeagus and the sternites VII and VIII, the latter with concave posterior margin and
conspicuous patterns of black hairs. These additional diagnostic characters are also
pointed out in LOHSE (1989).
The examination of the external morphology of the holotype showed that the
head was indeed rather narrow, whereas the shape of the pronotum and the length of
TYPES OF LATHROBIUM 963
the elytra are of little significance. I have seen some 600 specimens of L. longulum so
far, and these characters are subject to considerable variability. The study of the
aedeagus and sternites VII and VIII after microscopic preparation, with a compound
microscope at magnifications of 200-400 x held some surprises in store: the general
shape of the aedeagus as well as the characteristic structures in the internal sac were
clearly identical with those of L. longulum, both in ventral and in lateral view (Fig.
la-b). The impression of a stouter and apparently more strongly bent apex of the
median lobe is in fact an artefact caused by damage; part of the apex is missing (Fig.
2). Finally, the sternites VII and VIII show no trace of conspicuous patterns of hairs
and their shapes are completely different from those depicted in BORDONI (1980a). On
the contrary, apart from the abnormal asymmetry of sternite VIII, they were the same
as in L. longulum (Figs 2-4). Hence, since the holotype of L. patris doubtlessly repre-
sents an aberrant specimen of L. longulum, L. patris G. Benick must be considered a
junior synonym of L. longulum Grav.
Lathrobium longulum Gravenhorst, 1802
Lathrobium longulum GRAVENHORST, 1802: 53.
Lectotype d, here designated. Germania, Nr. 6346, Zool. Mus. Berlin. Paralectotypes:
236,3 9 9, here designated. Same data as lectotype.
Lathrobium patris G. BENICK, 1949: 16 ff., syn. n.
From the series of 6 syntypes of L. longulum a 3 was chosen as lectotype and
remounted. Its aedeagus (Fig. 3) and its sternite VIII (Fig. 6) were examined with a
compound microscope and subsequently glued next to the specimen. The internal sac
contained structures identical to those in Fig. la—b. Since there has been no recent
confusion regarding the identity of the species and since descriptions and diagnoses
are abundant in the literature, no detailed redescription is given here.
Lathrobium microps L. Benick, 1942
Holotype ®, labels: Spiekeroog, Diinen, VI.39, Microt. arv.-Nest, Larhrob. microps
L.B. Typus.
According to BENICK (1942) and LOHSE (1964) L. microps differs from L.
pallidum in its smaller size, uniform light colour, the coarser punctation of the
abdomen and the more distinct microsculpture on head and pronotum, and from L.
dilutum primarily in its slightly reduced length, shorter antennomeres and shorter
head. These differential characters are rather vague or even inadequate: from what the
considerably damaged type specimen had to offer, it was well within the range of
intraspecific variability of L. dilutum. In order to support this impression the female
terminalia, which have been shown to be highly useful for the identification of many
other Staphylinidae (see e.g. UHLIG 1989), of L. dilutum, L. pallidum and the holotype
of L. microps were examined (Figs 7-12). The striking differences in shape,
proportions, pubescence and number and position of long setae between L. dilutum
and pallidum indicate that the significance of the female terminalia may indeed be at
964 VOLKER ASSING
Fics 1-6
Lathrobium longulum Grav.: Aedeagus (la, b) and hind margin of & sternite VIII (4) of a
specimen from the Liineburger Heide, Northwestern Germany; apex of median lobe (2) and
hind margin (outline) of d sternite VIII (5) of the holotype of L. patris G. Benick; aedeagus in
lateral view (3) and hind margin (outline) of & sternite VIII (6) of the lectotype of L. longulum
Grav. Scale: 1, 3-6: 0.25 mm; 2: 0.10 mm.
TYPES OF LATHROBIUM 965
least as great for the identification, taxonomy and also phylogenetics of Lathrobium
species as the aedeagus. Against this background, the obvious similarity between L.
microps (Fig. 11) and L. dilutum (Figs 10, 12) leaves little doubt that they are in fact
conspecific. Hence L. microps L. Benick must be considered a junior synonym of L.
dilutum Er.
Lathrobium dilutum Erichson, 1839
Lathrobium dilutum ERICHSON, 1839: 509.
Lectotype 2, here designated. Berolin, Nr. 6348, Zool. Mus. Berlin. Paralectotypes,
here designated: 1 ©, Berolin, Nr. 6348, Zool. Mus. Berlin; 1 2 (abdomen missing!),
Pommern, Nr. 6348, Zool. Mus. Berlin; 1 2 (head missing, abdomen glued on separate
label), 6348, dilutum Er., Pomer. Er., Zool. Mus. Berlin.
Lathrobium microps L. BENICK, 1842: 168 ff., syn. n.
From the 4 syntypes of L. dilutum, a 2 in fairly good condition was selected as
lectotype and remounted. Its terminalia were examined with a compound microscope
(Fig. 12) and subsequently glued next to the specimen. The hind margin of its sternite
VIII is rounded as in Fig. 9. For further details concerning the external morphology of
the species the abundant diagnoses and keys in the literature should be consulted.
Xantholinus sejugatus G. Benick, 1953
Holotype d, labels: Lübeck, Schnellbruch, 10.XI.45, Uberschwemmung, Dr. Benick,
Xantholinus sejugatus G. Bck., Typus.
According to the original description (BENICK 1953) — which places the species
near X. linearis (Ol.) and erroneously claims the type to be a 2 — and LOHSE (1964) X.
sejugatus 1s distinguished from related species by the shape of the head with its sides
strongly diverging posteriorly. In addition, the gular sutures are completely separated,
whereas in the other species they are fused in the middle. As the aedeagus, which is of
particular taxonomic significance in this genus, was unknown, X. sejugatus has been
treated in major studies on Palaearctic Xantholinus as a species of incertae sedis (e.g.
BORDONI 1980b; COIFFAIT 1972).
Since the holotype is in fact a d, it was possible to examine the structures in
the internal sac. The aedeagus had been dissected and the internal sac glued on a
separate label prior to the present study. After microscopic preparation and slight
squeezing, the contents of the internal sac was examined with a compound micro-
scope and compared with the related species. It was clearly identical with that of X.
longiventris Heer, which means that the holotype of X. sejugatus represents an
externally aberrant specimen of X. longiventris. As far as the characters on the head
are concerned, it looks as though the gular sutures failed to fuse during ontogenesis,
which resulted in an abnormally widened head. Therefore, X. sejugatus G. Benick is
to be regarded as a junior synonym of X. longiventris Heer.
Like L. patris and L. microps the example of X. sejugatus demonstrates that
the intraspecific variability of a species is easily underestimated. In regions like
966 VOLKER ASSING
Fics 7-12
? terminalia (urites IX and X) (7) and hind margin of © sternite VII (8) of L. pallidum Nordm.;
hind margin of © sternite VIII (9) and 2 terminalia (10) of L. dilutum Er. (Hannover, Germany);
? terminalia (11) of the holotype of L. microps L. Benick (pubescence and setae omitted); 9
terminalia (12) of the lectotype of L. dilutum Er. Tergites X dotted. Scale: 0.25 mm.
TYPES OF LATHROBIUM 967
Central Europe with high collecting activity and abundance of available material
species descriptions should be based on series of specimens, and the genitalia should
always be considered. In the course of ecological field work I have seen more than
10,000 Xantholinus (ASSING 1993), among them numerous specimens with aber-
rations of various body parts. After all, malformations during ontogenesis occur in all
forms of living organisms, as the late renowned coleopterologist Dr G.A. Lohse has
repeatedly pointed out.
Xantholinus longiventris Heer, 1839
Xantholinus longiventris Heer, 1839: 247.
Lectotype d, here designated (remounted, aedeagus dissected and examined): red trian-
gular label [= Switzerland, Museum Escher] (coll. Heer, ETH Ziirich). Paralectotypes,
here designated: 19, same labels as lectotype; 12, blue label “b” [= Matt, Kanton
Glarus, leg. Heer] (coll. Heer, ETH Zürich).
Xantholinus sejugatus G. Benick, 1953: 39f., syn. n.
REFERENCES
ASSING, V. 1993. Zur Bionomie von Xantholinus rhenanus Coiff. und anderen bodenbe-
wohnenden Xantholininen (Col., Staphylinidae) in Nordwestdeutschland. Zool. Jb. Syst.
120: 13-38.
BENICK, G. 1949. Eine neue deutsche Lathrobium-Art. Entomol. Blätter 45-46 (1949-50):
16-18.
BENICK, G. 1953. Ein neuer Xantholinus (Col. Staphyl.) aus Norddeutschland. Verh. Ver.
naturwiss. Heimatforsch. Hamburg 31: 39-40.
BENICK, L. 1942. Eine neue deutsche Lathrobium-Art. Lathrobium microps n. sp.. Entomol.
Blätter 38: 168-170.
BORDONI, A. 1980a. Studi sui Paederinae — V — Intorno ad alcuni Lathrobiini poco noti,
revisione dei tipi e descrizione di una nuova specie italiana (Col. Staphylinidae).
Frustula Entomol. (N.S.) 2: 1-26.
BORDONI, A. 1980b. Stato attuale della conoscenza degli Xantholinus W-Palearctici: un
esempio significativo della necessita dello studio del sacco interno in alcuni coleotteri
stafilinidi. Atti XII Congr. Naz. Ital. Entomol., Roma 2: 37-43.
COIFFAIT, H. 1972. Coléoptères Staphylinidae de la région paléarctique occidentale. I.
Généralités; sous-fam. Xantholininae et Leptotyphlinae. Nouv. Rev. Ent. Suppl. 2(2):
1-651.
COIFFAIT, H. 1982. Coléoptères Staphylinidae de la région paléartique occidentale. IV. Sous-
famille Paederinae. Tribu Paederini 1 (Paederi, Lathrobii). Nouv. Rev. Ent., Suppl.
12(4): 1-440.
ERICHSON, W.F. 1837-39. Die Kafer der Mark Brandenbourg, I. Berlin, 740 pp; 1839: 385-740.
GRAVENHORST, J. 1802. Coleoptera Microptera Brunsvicensia. Brunsvigae: 206 pp.
HABERMAN, H. 1983. Beitrag zu “Enumeratio Coleopterorum Fennoscandiae et Daniae” (1979)
über die Staphyliniden Estlands. Notul. Entomol. 63: 97-110.
HEER, O. 1839. Fauna Coleopterorum Helvetica. Pars 1. Turici [Zürich], 145-364.
968 VOLKER ASSING
Louse, G.A. 1964. Fam. Staphylinidae I (Micropeplinae bis Tachyporinae). In: FREUDE, H.,
HARDE, K.W. & G.A. LOHSE: Die Käfer Mitteleuropas. Bd. 4, Krefeld, 1-264.
Louse, G.A. 1989. 23. Familie Staphylinidae (I) (Piestinae bisTachyporinae). In: LOHSE, G.A.
& W.H. LucHT: Die Käfer Mitteleuropas. 1. Supplementband mit Katalogteil, Krefeld,
121-183.
UHLIG, M. 1989. Zur Morphologie der weiblichen Terminalia einiger Staphylinidenarten
(Coleoptera). Verh. XI. SIEEC Gotha 1986: 227-37.
ZIEGLER, W. & R. SUIKAT. 1994. Rote Liste der in Schleswig-Holstein gefährdeten Käferarten.
Landesamt fiir Naturschutz und Landschaftspflege Schleswig Holstein: 1-96.
REVUE SUISSE DE ZOOLOGIE, 102 (4) : 969-978; décembre 1995
Espèces nouvelles ou mal connues de Cholevidae (Coleoptera)
Michel PERREAU
Laboratoire de Magnétisme des surfaces, Université Paris 7,
2, place Jussieu, F-75251 Paris cedex 05, France.
New or poorly known species of Cholevidae (Coleoptera). - New data
are given on the morphology and the distribution of Choleva gabriellae
Giachino, and Attaephilus scheuerni Perreau. The descriptions of the
following new species are given: Ptomaphaginus balazuci n. sp., Ptoma-
phaginus baliensis n. sp., Ptomaphaginus riedeli n. sp., and Choleva (Pro-
tocatops) medvedevi n. subgen., n. sp. With the latter species the distri-
bution of Choleva is extended to Vietnam. Some apomorphic characters
shared with Choleva henroti Szymezakowski, lead to the description of a
new subgenus: Protocatops.
Key-words: Coleoptera - Cholevidae - Taxonomy - Distribution.
INTRODUCTION
Nous regroupons dans cet article un certain nombre d'observations sur des
Cholevidae originaires de la région méditerranéenne et de l'Asie du sud-est. L'aire de
répartition de certaines espèces se trouve précisée, c'est le cas pour Choleva ga-
briellae Giachino, dont seul le male était jusqu'à présent connu. Attaephilus scheuerni
Perreau, n'avait fait l'objet que d'une diagnose préliminaire, et nous complétons ici sa
description. D'autres espèces sont nouvelles: Promaphaginus balazuci n. sp., Ptoma-
phaginus baliensis n. sp., Ptomaphaginus riedeli n. sp., et une espèce présentant un
intérét particulier, du point de vue phylogénétique et biogéographique: Choleva
(Protocatops) medvedevi n. subgen., n. sp., première espèce de Choleva vietna-
mienne.
Le genre Choleva, essentiellement paléarctique, n'était connu de l'Asie du sud-
est que par une seule espèce distribuée au Népal et en Assam. La présence d'une
nouvelle espece au Vietnam, d'ailleurs apparentée a l'espèce himalayenne repousse
donc encore vers le sud-est l'aire de répartition du genre Choleva, les caractères
particuliers communs à ces deux espèces sont la raison de la séparation du sous-genre
Protocatops n. subgen.
Manuscrit accepté le 23.01.1995.
970 MICHEL PERREAU
Abbréviations utilisées: MHNG: Muséum d'histoire naturelle de Genève:
MNHNP: Muséum National d'Histoire Naturelle de Paris; SMNS: Staatliches
Museum fiir Naturkunde Stuttgart; CMP: collection Michel Perreau.
DESCRIPTIONS
Le genre Ptomaphaginus est un groupe très homogène dont il est souvent bien
difficile de distinguer les espèces sur la seule morphologie externe. Certaines espèces
présentent, exceptionnellement des particularités bien marquées constituant d'inté-
ressants caractères distinctifs: échancrures entourées ou non de dents sur les ventrites
VII et VIII du male (P. franki Perreau, P. trautneri Perreau), touffe de poils a
l'extrémité des élytres (P. pilipennis Perreau, P. pilipennoides Perreau), soies parti-
culièrement longues a la base des fémurs... Toutefois, la plupart des espèces ne
peuvent être distinguées que par les caractères sexuels, édéage chez le mâle, et
spermathèque chez la femelle. Méme les mensurations ne donnent que de vagues
indications qui ne permettent plus de reconnaître le nombre croissant d'espèces. C'est
la raison pour laquelle les descriptions que nous donnons des nouvelles espèces de
Ptomaphaginus sont très courtes, et n'insistent que sur les vraies particularités.
Ptomaphaginus balazuci n. sp.
Holotype d : Indonésie, Ngalam Kamang près de Tinngo, le 20-7 1979, J. Balazuc leg.
(MNHNP).
Description: Longueur: 3,2 mm. Coloration générale brun foncé, les antennes
uniformément plus claires. Tout le corps recouvert d'une fine pubescence dorée.
Bord postérieur du pronotum rectiligne, non sinué.
Bord apical du ventrite VIII sinué (fig. 4).
Edéage presque rectangulaire en vue dorsale, (figs 1 et 2), segment IX repré-
senté sur la figure 3.
Femelle inconnue.
Cette espèce se reconnait facilement par la forme de l'édéage et la sinuosité du
bord postérieur du ventrite VIII.
Ptomaphaginus baliensis n. sp.
Holotype d: Indonésie, Bali, Mt Batukao, 500-700 m, 28-29/10/1991, I. Löbl
(MHNG). Paratypes: 1 4, même origine (MHNG), 1 ¢ et 2 ®, Bali, Mt Agung, Besahik au-
dessus du temple, 31/10—1/11/1991, I. Löbl (MHNG et CMP).
Description du male: longueur: 2,5 mm. Coloration générale brune, les
antennes brun clair. Tout le corps est recouvert d'une fine pubescence.
Bord postérieur du pronotum légèrement sinué vers l'arrière au niveau des
angles.
Ventrite VIII avec une trés faible sinuosité du bord postérieur.
ESPECES NOUVELLES DE CHOLEVIDAE 971
MÈ
Fics 1-9
1-4: Ptomaphaginus balazuci n. sp., holotype. 1. Edéage face dorsale; 2. Edéage face latérale:
3. Segment IX; 4. Ventrite VII du mâle. 5-9: Ptomaphaginus riedeli n. sp. 5. Edéage face
dorsale; 6. Edéage face latérale; 7. Segment IX; 8. Ventrite VII du male; 9. Spermathèque.
L'échelle représente 0,5 mm.
972 MICHEL PERREAU
Edéage particulièrement robuste, présentant un stylet interne très fin et très
long, près de trois fois plus long que le lobe médian et des soies sur les còtés de la
face inférieure, 6 du côté droit, et 7 du côté gauche (Figs 10 et 11). Segment IX
représenté sur la fig. 13.
Femelle sans dilatation des tarses antérieurs. Spermathèque représentée sur la
in 1125
13
10
11
Fics 10-13
Ptomaphaginus baliensis n. sp. 10. Edéage face ventrale; 11. Edéage face laterale; 12. Segment
IX; 13. Spermathèque. L'échelle représente 0,5 mm.
ESPECES NOUVELLES DE CHOLEVIDAE 973
La morphologie externe ne présente pas d'autre caractère spécial permettant la
distinction des autres espèces. P. baliensis n. sp. se reconnaît facilement au stylet
interne de l'édéage particulièrement long, le plus long connu au sein du genre
Ptomaphaginus.
Ptomaphaginus riedeli n. sp.
Holotype d: Inde, Tamil Nadu, Ooticamund, Pytara Mysore, 22-8-89, Riedel leg.
(SMNS). Paratypes: 4 d et 3 2, de même provenance (SMNS et CMP).
Description du male: longueur: 4 mm. Coloration générale brun foncé a l'ex-
ception des trois derniers articles antennaires très éclaircis, comme chez P. piraster
Szymezakowski et P. apiculatus Szymezakowski.
Elytres avec une touffe de poils épais, comme celle existant chez P. pilipennis
Perreau, et a un moindre degré chez P. pilipennoides Perreau.
Ventrite VIII du male présente une échancrure semi-elliptique le long du bord
postérieur (figure 9).
Tarses antérieurs fortement dilatés, plus larges que l'apex des tibias.
Edéage large à la base, piriforme comme celui du P. piraster, mais avec l'apex
effilé et les côtés parallèles aux extrémités, non arrondis. Stylet interne grêle (figs 5 et
6). Segment IX abdominal représenté sur la fig. 7.
Femelle différente du mâle par l'absence de la dilatation des tarses antérieurs,
et l'absence de l'échancrure sur le bord postérieur du ventrite VII. Spermathèque
représentée sur la fig. 8.
Cette espèce fait partie d'un groupe caractérisé par une forme commune de
l'édéage en poire, et qui comprend aussi P. piraster, P. apiculatus, P. pilipennis et P.
pilipennoides. P. riedeli n. sp. se distingue de ces quatre espèces par la taille nette-
ment plus grande, la présence de l'échancrure du ventrite VIII chez le mâle, et la
forme de l'édéage. Il se distingue aussi plus spécifiquement de P. pilipennis et de P.
pilipennoides par la couleur claire des trois derniers antennomeres et de P. piraster
par la présence de la touffe de poils sur l'apex des élytres.
Choleva Latreille
Protocatops n. subgen.
Espece-type: Choleva (Cholevopsis) henroti Szymczakowski
Ce sous-genre est caractérisé par sa forme large et convexe, la rugosité de la
ponctuation élytrale, le segment génital non sclérifié, la disparition des stries élytrales.
SZYMCZAKOWSKI (1961) décrivait Choleva (Cholevopsis) henroti récoltée en
Assam. Il attribua cette espèce au sous-genre Cholevopsis à cause de la rugosité de la
ponctuation élytrale, et du segment IX de la femelle non sclérifié. Il insistait toutefois
sur les différences importantes existant avec les autres espèces du sous-genre. Aujour-
d'hui, la découverte d'une seconde espèce étroitement apparentée mais bien distincte
du Vietnam nous conduit a la conclusion que Ch. henroti n'est pas une espèce isolée,
mais la première représentante d'une lignée asiatique de Choleva bien différente de
toutes les autres lignées de ce genre.
974 MICHEL PERREAU
En fait, l'habitus de ces espèces rappelle étrangement celui du genre Catops:
— les stries élytrales longitudinales ont totalement disparu (sauf la suturale),
— les élytres présentent une microréticulation, plus prononcée que celle des
Cholevopsis, mais n'allant pas toutefois jusqu'à la teinte cendrée des élytres
des Catops,
— la forme est large et très convexe,
— la coloration générale est très foncée,
— le dernier antennomère de la Ch. medvedevi est éclairci, comme chez
certains Catops.
Plusieurs caractères montrent sans ambiguité l'appartenance aux Cholevini et
non aux Catopini:
— l'édéage de la Ch. henroti présente des styles larges et épais (celui de la Ch.
medvedevi est inconnu),
— l'épistome est très nettement séparé du front par une suture clypéo-frontale
très profonde,
— le ventrite VIII des femelles ne présente pas de fossette en arrière du
spiculum ventrale, alors qu'il en existe une chez tous les Catopini.
Ces deux espèces constituent donc une lignée ayant conservé certains carac-
tères ancestraux (segment IX de la femelle non sclérifié), mais présentant aussi un
certain nombre d'apomorphies: absence de stries élytrales exceptée la suturale, rugo-
sité de la ponctuation élytrale. Toutes ces raisons nous ont conduit a créer ce nouveau
sous-genre.
En plus de ces caractéristiques morphologiques, cette lignée de Choleva pré-
sente une autre particularité. Les espèces semblent présenter une affinité particulière
pour des zones tropicales ou subtropicales, ce qui n'est le cas d'aucune espèce des
autres sous-genres.
Choleva (Protocatops) medvedevi n. sp.
Holotype 9: Vietnam, province de Vinh-Phu, Tamdao, 800-1200 m, 12/22-4-1966,
leg. Medvedev, Golovatch (SMNS).
Description: longueur: 4,3 mm. Forme large et convexe. Coloration générale
brun très foncé, presque noir, les 6 premiers antennomères et l'extrémité du dernier
plus clairs. Tout le corps couvert d'une fine pubescence dorée dressée. Téte, pronotum
et élytres entièrement et très finement microréticulés.
Antennes très élancées, les longueurs de chaque antennomère en microns (+
10) sont les suivantes: 200, 140, 120, 130, 150, 130, 120, 120, 170, 170, 200.
Pronotum ovale, 1,8 fois plus large que long, avec tous les angles très large-
ment arrondis, et nettement plus étroit que les élytres. Ponctuation beaucoup plus fine,
mais la microréticulation semblable à celle des élytres.
Elytres très larges et très convexes, seulement 1,23 fois plus longs que larges,
et 1,5 fois plus larges que les pronotum. Stries longitudinales totalement absentes
ESPECES NOUVELLES DE CHOLEVIDAE 975
excepté la strie suturale. La ponctuation est rugueuse entremélée d'une fine micro-
réticulation.
Tibias intermédiaires légèrement arqués.
Segment IX abdominal non sclérifié.
Spiculum ventrale étroit, les côtés parallèles, arrondi en avant (fig. 19).
Male inconnu.
Cette espèce est très semblable a Choleva henroti Szymezakowski décrite
d'Assam (SZYMCZAKOWSKI 1961, 1964), et retrouvée par la suite au Népal (PERREAU,
1988). Elle s'en distingue par les caractères suivants:
— taille plus grande,
— huitième antennomere moins allongé, à peu près carré au lieu de 1,4 fois
plus long que large,
— tibias postérieurs moins arqués,
— le caractère le plus important est la forme très différente du spiculum
ventrale, étroit, parallèle et arrondi a l'extrémité chez Ch. medvedevi, triangulaire a
base large, et anguleux a l'extrémité chez Ch. henroti (Figs 18 et 19).
Choleva gabriellae Giachino
Algérie: Atlas de Blida, Chréa sur Blida, 1400 m, 3-5-1988, Besuchet, Löbl et Burck-
hardt, 1 d et 1 2 (MHNG).
La femelle était inconnue jusqu'alors. Le ventrite VIII et le spiculum ventrale
sont représentés sur la fig. 20 et le segment IX sur la fig. 21.
Lors de sa description, Giachino n'a pas tranché entre l'appartenance au groupe
“cisteloides” ou au groupe “reitteri”. Cette espèce présente en effet des caractères
intermédiaires, la structure du sac interne de l'édéage la rapproche du groupe
“cisteloides” et la forme générale du corps, la forme des trochanters postérieurs mâles
et la présence de soies dressées sur les élytres la rapprochent du groupe “reitteri”. Les
deux groupes sont étroitement apparentés, et les caractères utilisés par Jeannel pour
les séparer ne sont pas clairement des apomorphies.
Attaephilus scheuerni Perreau
Seule avait été publiée une diagnose préliminaire de cette espèce (PERREAU,
1994). Nous en donnons ici une description plus détaillée. La série typique ne se
trouve plus dans la collection Scheuern comme indiqué dans la diagnose. Le matériel
qui a servi a la description est le suivant: holotype d: Paxos, Corfu (Kerkira), 2 km
NW Gaios, 17-4-81, J. Scheuer (SMNS); paratype: 1 4 même provenance (CMP).
Description: longueur: 3,2 mm. Coloration générale brun-jaune clair. Tout le
corps recouvert d'une double sétulation dorée, l'une composée de soies courtes et
couchées, l'autre de soies longues et dressées.
Pronotum 1,8 fois plus large que long, les angles postérieurs très étroitement
arrondis, la base saillante en arrière, arrondie en son milieu. Côtés régulièrement
rétrécis de l'arrière vers l'avant, la plus grande largeur à la base. Soies dressées du
976 MICHEL PERREAU
14
Miti i
18 19
Si Sons 20 jai, 9 21
Han
Fics 14-21
14-18: Attaephilus scheuerni Perreau. 14. Edéage face dorsale; 15. Ventrite V du mâle; 16.
Tibia intermédiaire mâle: 17. Trochanter intermédiaire droit du mâle, face ventrale. 18.
Choleva (Protocatops) henroti Szymczakowski, ventrite VII femelle et spiculum ventrale. 19:
Choleva (Protocatops) medvedevi n. sp. holotype, ventrite VIII femelle et spiculum ventrale. 20
et 21: Choleva (s. str.) gabriellae Giachino. 20. ventrite VIII femelle et spiculum ventrale; 21.
Segment IX face dorsale. L'échelle représente 0,5 mm. pour les figs 14, 16, 17, 19, 20, 21, et 1
mm pour les figs 15, 18, 19, 20, 21.
ESPECES NOUVELLES DE CHOLEVIDAE 977
pronotum particulièrement bien visibles à la base le long de laquelle un certain
nombre d'entre elles s'alignent.
Elytres à ponctuation rugueuse et irrégulière. Pubescence dressée alignée
longitudinalement.
Ventrite V avec une très forte dent médiane (fig. 15). Trochanters inter-
médiaires aplatis en une palette épaisse perpendiculaire au plan du corps (fig. 17).
Face interne des tibias intermédiaires très échancrée. Face supérieure ornée de
deux carènes, l'une sur le bord interne s'étendant sur toute la longueur du tibia excepté
une très courte distance à la base, l'autre sur le bord externe, s’étendant sur le tiers
médian. Tibia représenté sur la figure 16. Tarses antérieurs peu dilatés, à peine plus
larges que le tibia.
Edéage large, rétréci très brusquement à l'apex, avec un petit bouton apical.
Sac interne armé de deux phanères latérales constituées de dents fines et d'une rangée
centrale de dents nettement plus épaisses. Paramères épais sauf à l'apex, et divergents
aux extrémités (fig. 14).
La conformation des tibias intermédiaires rapproche cette espèce de A.
arenarius Hampe. Par contre, l'édéage ressemble à celui de A. funebris Reitter.
Le genre Artaephilus peut être divisé en trois groupes d'espèces. L'un caractérisé
par un édéage effilé et pointu à l'apex, et avec des antennes courtes à massue compacte.
Il comprend Attaephilus laticornis Frank & Perreau, A. weisi Reitter, et A. servati
Coiffait. Ce groupe ressemble aux Catopomorphus sauf la pubescence beaucoup plus
longue. Le second groupe présente aussi un édéage du méme type que le groupe
précédent, mais les antennes sont plus longues et plus déliées. Il comprend Attaephilus
arenarius Hampe, punctipennis Jeannel, paradoxus (Motschoulsky) et weiratheri
Jeannel. Le troisième groupe est caractérisé par un édéage quadrangulaire en vision
dorsale, rétréci très brusquement a l'apex, terminé ou non par un petit bouton apical.
Les antennes sont également longues et grèles. Il comprend les Aftaephilus funebris
Reitter, rambouseki Jeannel, illyricus Jeannel, angustus (Reitter), persicus Szymeza-
kowski, niger Karaman. Les males d'A. molitori Scheerpeltz et reitteri Jeannel sont
inconnus et ne peuvent étre situés actuellement dans ces groupes. A. scheuerni n. sp. se
place dans le troisième groupe et se distingue des autres espèces par la forme des tibias
intermédiaires males.
Les deux exemplaires connus ont été récoltés dans une fourmilière de Messor
structor.
REMERCIEMENTS
Je tiens a remercier toutes les personnes qui m'ont confié l'étude du matériel
figurant dans cet article ainsi que celles qui m'ont permis d'accéder aux collections
pour effectuer les comparaisons nécessaires aux déterminations et aux descriptions: J.
Balazuc (Nogent sur Marne), N. Berti (Muséum National d'Histoire Naturelle, Paris),
I. Löbl (Muséum d'histoire naturelle de Genève), W. Schawaller (Staatliches Museum
für Naturkunde, Stuttgart), J. Scheuern (Sinzig-Westum). Je remercie aussi J.
978 MICHEL PERREAU
Weulersse (Muséum National d'Histoire Naturelle, Paris), pour la détermination de la
fourmi hôte de l'Attaephilus scheuerni, et I. Ruzicka (Prague) pour ses intéressantes
remarques.
REFERENCES
GIACHINO, P.M. 1985. Choleva (s. str.) gabriellae e Sciodrepoides casalei nuove specie della
Kabylia e note corologiche sui Catopidi d'Algeria (Coleoptera, Catopidae). Boll. Mus.
Reg. Sc. Nat., Torino, 3: 337-348.
PERREAU, M. 1988. Les Cholevidae himalayens du Muséum d'Histoire naturelle de Genève.
Revue Suisse Zool., 95: 1005-1018.
PERREAU, M. 1994. Diagnose préliminaire d'une nouvelle espèce d'Attaephilus de Grèce (Colé-
optères, Cholevidae). Bull. Soc. ent. France, 99: 384.
SZYMCZAKOWSKI, W. 1961. Eléments paléarctiques dans la faune orientale des Catopidae
(Coleoptera). Acta zool. cracov., 6: 123-136.
SZYMCZAKOWSKI, W. 1974. Nouvelles remarques sur les Catopidae (Coleoptera) de la région
orientale. Acta zool. cracov., 19: 197-216.
SZYMCZAKOWSKI, W. 1975. Catopidae (Coleoptera) récoltés dans le sud de l'Inde par l'expé-
dition du Muséum de Genève en 1972. Acta zool. cracov., 20: 121-149.
REVUE SUISSE DE ZOOLOGIE, 102 (4) : 979-988; décembre 1995
Two remarkable afromontane Theridiidae:
Proboscidula milleri n. sp. and Robertus calidus n. sp.
(Arachnida, Araneae)
Barbara KNOFLACH
Institute of Zoology, University of Innsbruck,
Technikerstraße 25, A-6020 Innsbruck, Austria.
Two remarkable afromontane Theridiidae: Proboscidula milleri n. sp.
and Robertus calidus n. sp. (Arachnida, Araneae). - Two new minute
theridiid species are described from montane forests in Rwanda and Zaire
and their genital morphology and relationships are discussed. P. milleri is the
second species in this genus after the type species from Angola. R. calidus is
the first afrotropical species in an otherwise holarctic genus. It is closely
related to the western palaearctic species R. neglectus (O.P.-Cambridge).
Key-words: Araneae - Theridiidae - Taxonomy - Afromontane - Probosci-
dula - Robertus.
INTRODUCTION
In the linyphiid spider fauna of the African mountains both afrotropical ele-
ments as well as species with palaearctic relatives are present (HOLM 1962, SCHARFF
1992). Two minute theridiid species collected by E. Heiss in montane forests in
Rwanda and Zaire which are described in the following pages show analogous
relationships. The first one belongs clearly to the attractive genus Proboscidula,
which was previously known only from Angola. The second species, Robertus cali-
dus, is the first African representative of a hitherto holarctic genus.
ABBREVIATIONS
C conductor, E embolus, f tegular fold, Pc paracymbium, ST subtegulum, T
tegulum, TA tegular apophysis. — MHNG Muséum d'histoire naturelle, Genève.
DESCRIPTIONS
Proboscidula milleri n. sp. (Figs 1-16)
Types: Rwanda, Nyakabuye, montane rain forest Cyamudongo, ca. 2000 m, 1 d
holotype, 1 9 paratype, early February 1986; 1 ® paratype, 20.-25. January 1984, leg. E. Heiss
(MHNG).
Manuscript accepted 16.05.1995.
980 BARBARA KNOFLACH
Diagnosis: d carapace with characteristic clypeal knob (Figs 1, 3, 5), legs
I with conspicuous ventral spines (Fig. 4), abdomen with dorsal and ventral scutum,
sternum covered with numerous glandular hairs (Fig. 2). 4 palp with projecting tegular
apophysis (Fig. 14). Females can be recognised by their epigyne/vulva (Figs 11, 12). P.
milleri is separated clearly from the type species P. loricata Miller, 1970.
Etymology: This striking species is named in honour of Prof. F. Miller,
Brno (1902-1983).
d: Measurements (mm): carapace 0.53 long, 0.46 wide. Length of abdomen
0.71. Posterior end of sternum 0.14 wide.
Colour, pattern: Carapace brown, sides reticulate, lateral margins with dark
seam, median area dark grey. Legs brown, trochanters, tibiae and distal part of femora
light brown. Dorsum of abdomen dusky brown, light median band continuous with
anterior light belt (Fig. 1). Sides of abdomen pale with a dark longitudinal stripe (Fig.
10). Venter pale, epigaster darkened. Spinnerets light brown.
Carapace with conspicuous curved clypeal knob covered with stout spines (Figs
1-3, 5). Lateral margins of carapace with a row of very short hairs, median area with 5
long bristles. Eye region (Fig. 1) as in P. loricata (MILLER 1970). Chelicerae typically
theridiid, with basal extension, anterior margin with two teeth (Fig. 3). Carapace ele-
vated, posterior declivity steep (Fig. 5). Stridulatory organ well developed, not divided
in midline, similar to Robertus. Anterior to stridulatory ridges at each side a row of 5-6
warts which form the bases of short hairs. Sternum vaulted, broadly truncate behind,
twice as wide as diameter of coxa IV. Numerous modified hairs present, which may be
glandular, for their bases show conspicuous tubes (Fig. 2). Labium not separated.
Abdomen with large dorsal scutum covering > 2/3 of its length and with
ventral scutum from pedicel to epigastric furrow (Fig. 10). Dorsum smooth, with 3
pairs of impressed dots (Fig. 1). Branchial opercula and stigmata sclerotised.
Spinnerets (Fig. 9) surrounded by small sclerotised ridges. Colulus large, 3 setae
present. The two modified spigots of the posterior lateral spinnerets of the females
could not be recognised.
Leg measurements (mm):
Fe Pa Ti Mt Ta Total
Palp 0.20 0.11 0.09 = 0.19 0.59
I 0.40 0.18 025 0.19 0.24 125
II 0.34 0.16 0.21 0.16 0.24 1.10
III 0.30 0.14 0.16 0.14 0.22 0.95
IV 0.44 0.17 0.26 0.20 0.27 11:93)
Legs: 4123. Femora, tibiae and metatarsi of first leg ventrally with two rows of
conspicuous spines (Fig. 4), their bases conical. Numbers of prodorsal/retrodorsal
trichobothria of tibiae I-III 1/2, IV 2/2, their position on I 0.18/0.24;0.42, on IV
0.16:0.36/0.29:0.47. Metatarsi HI with 1 trichobothrium (0.39; 0.32). Tarsal organ
I-IV 0.25. Tarsi I-IV 1.3-1.5 times longer than metatarsi. Tarsal claws with ca. 3
minute teeth.
TWO AFROMONTANE THERIDIIDAE 981
Fics 1-5
Proboscidula milleri n. sp. Male. Dorsal view (1). Sternum and mouthparts (2). Carapace and
chelicerae, frontal (3) and lateral view (5). Leg I (4). Scale lines 0.2 mm.
982 BARBARA KNOFLACH
Fics 6-12
Proboscidula milleri n. sp. Female (6-8, 11, 12). Male (9, 10). Carapace, dorsal (6) and frontal
view with chelicerae (8). Right chelicera, frontal view (7). Spinnerets (9). Abdomen, lateral view
(10). Epigyne/vulva, ventral (11) and dorsal view (12). Scale lines 0.2 mm (6, 8, 10), 0.05 mm (7,
OED):
TWO AFROMONTANE THERIDIIDAE 983
6 Palp: Figs 13-16. Patella slightly widened. Tibia with 1 retrolateral tricho-
bothrium. Cymbium plate-shaped, distally broad, not tapering (Figs 14, 16). Para-
cymbium in retrolateral-distal position, hook-like, locking with tegular apophysis
(Figs 14, 15). Subtegulum large, at dorsal side of bulb. Tegulum in retrolateral-ventral
position, with wide retrolateral concavity which holds a projection of embolar base
and is hidden by the cymbium (arrow, Fig. 13). Conductor and tegular apophysis
present. Conductor: dorsal, transparent, bandlike, supporting tip of embolus (Figs
13-15). Tegular apophysis elongate, attached to tegulum on ventral side (M, Fig. 14)
and enormously developed. Its characteristic prolateral angle extends beyond the
cymbium (Figs 14, 16), retrolateral end with concavity into which fits the para-
cymbium. At the base of the tegular apophysis a further small process is discernible
beside the embolus (arrow, Fig. 14). Embolus: Fig. 14. Embolar base broad, rounded,
attached to tegulum retrolaterally. Distal part very short, sperm duct with subterminal
coil.
2: Measurements (mm): Carapace 0.56 long, 0.52 wide. Length of abdomen
0.94. Posterior end of sternum 0.15 wide.
Colour, pattern: Coloration of carapace slightly stronger than in d, legs as in
3d. Abdomen dark, median light band clear, no lateral markings. Venter pale except
one median greyish patch behind epigastric furrow.
Carapace without knob (Figs 6, 8), stridulatory ridges on posterior declivity
reduced. Chelicerae as in d (Figs 7, 8). Sternum broadly truncate as in d , but without
glandular hairs. Abdomen evenly and strongly sclerotised, regularly covered with
small indentations, but without definite scutum. Branchial opercula and stigmata scle-
rotised, sclerotised ridges also present around spinnerets. Colulus as in d, posterior
lateral spinnerets with two modified cylindrical spigots as is typical for theridiids.
Leg measurements (mm):
Fe Pa Ti Mt Ta Total
Palp 0.17 0.09 0.08 = 0.19 0.52
I 0.40 0.18 0.24 0.19 027 127
II 0.36 0.16 022 0.18 0.26 137,
II 0.30 0.15 0.18 Ons 023 1.02
IV 0.41 0.18 0.30 0.19 0.28 ISS
Legs: 4123. Leg I without ventral spines. Trichobothria of tibiae as in d,
position on I 0.18/0.24;0.43, on IV 0.19;0.36/0.38;0.58. Trichobothrium of metatarsi
HI 0.37;0.42. Tarsal organ I-IV 0.27-0.31. Leg IV with tarsal comb of 5 serrate
bristles. Tarsal claws as in d.
Epigyne/vulva: Figs 11, 12. Epigyne with transverse, sclerotised edge, in front
of which are the orifices of the copulatory ducts (Fig. 11). Receptacula seminis
posterior to epigynal edge. Copulatory ducts short, leading straight backwards into the
receptacula (Fig. 12). Fertilization ducts start on ventral side of receptacula and turn
to the midline. Glandular pores present in receptaculum and copulatory duct (Fig. 12).
984 BARBARA KNOFLACH
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Fics 13-16
Proboscidula milleri n. sp. Male palp, retrolateral (13), ventral (14), frontal (15) and dorsal
view (16). Scale line 0.1 mm.
TWO AFROMONTANE THERIDIIDAE 985
A ffinifies: The generic placement of P. milleri seems to be unam-
biguous as most characters mentioned in the diagnosis of MILLER (1970) are present,
especially the peculiar clypeal projection of the 4. Even the “glandular” hairs of the
Sd sternum were seen by MILLER (1970, “Doppeldellen”). Only the following hardly
discernible characters differ: In P. loricata the colulus is said to be replaced by two
setae, the chelicerae are not armed and the tarsal claws smooth.
P. milleri is smaller than P. loricata, its clypeal knob directed ventrally and
bearing short setae. In P. /oricata the knob is bent forwards and bears long hairs. The
strong ventral spines of leg I of P. milleri were not mentioned for P. loricata. The
tegular apophysis is long also in P. loricata, but extends laterally. Distal part of
embolus and conductor are much longer. Therefore the copulatory ducts in the female
of P. loricata, which is still unknown, can be expected to be longer than in P. milleri.
Distribution, habitat: Known only from the montane rain forest at
Rwanda, Nyakabuye at about 2000 m. All specimens came from sieving bark and
litter. The type species was likewise found under bark of Brachistegia manga in
gallery forest of the river Luachimo, Dundo, Angola, at the end of February.
Discussion: P. milleri belongs to the group of theridiids with locking
system A in the male palp (SAARISTO 1978), with hook-like paracymbium which
anchors the tegular apophysis, the latter contains the sperm duct. Its colulus is
relatively large and the stridulatory organ well developed as in most genera of this
group which includes the former “Asageneae”.
As already pointed out by MILLER (1970) and LEVI (1972) there are affinities
to the South American genus Wirada. However, Wirada lacks a proboscis (LEVI
1963) and shows differences in the male palp: embolus strongly developed, conductor
absent, two tegular apophyses discernible. The female's receptacula are anterior to the
epigynal ridge. Similar modifications of the 4 carapace are present also in Craspe-
disia (LEVI & LEVI 1962, LEVI 1963) and Carniella (THALER & STEINBERGER 1988),
both of which have a large colulus, but their palpal structures are quite different. LEVI
(1972) supports the generic rank of Proboscidula.
Robertus calidus n. sp. (Figs 17-20)
é Holotype: Zaire, Prov. Kivu, Mt. Kahuzi, sieving bark and litter in bamboo
zone (Arundinaria alpina) ca. 2300 m, early February 1986, leg. Heiss (MHNG).
Diagnosis: R. calidus is closely related to R. neglectus (O.P.-Cambridge).
It differs by the following palpal structures: tegular apophysis distinct, embular base
large, distal part of embolus long and slender, its tip narrow. 2 unknown.
Etymology: Latin calidus, a, um = hot (german: heiß). The specific
name refers to both the occurrence of the species in a warm region as well as to its
collector.
3: Measurements (mm): Carapace 0.94 long, 0.73 wide. Length of abdomen
0.89.
Colour: Carapace and legs light brown, abdomen grey. Ocular area as in its
congeners, fig. 260 in WIEHLE (1937). Stridulatory organ well developed, not divided
in midline. Anterior margin of chelicerae with 3 teeth. Colulus large, 2 setae present.
986 BARBARA KNOFLACH
Leg measurements (mm):
Fe Pa da Mt Ta Total
Palp 0.44 0.19 0.11 ~ 0.32 1.06
I 0.75 0.31 0.57 0.42 0.38 DAN
II 0.57 0.30 0.40 0.36 0.35 1.97
III 0.50 0.24 0.35 0.31 0.31 1.70
IV 0.78 0.30 0.58 0.40 0.38 2.44
Legs: 4123. Legs IV and I almost equal. Numbers of prodorsal/retrodorsal
trichobothria of tibiae HIV: 1/3, 1/2, 2/2, 2/3, their respective positions (0.34/0.26;
0.43;0.66), (0.29/0.30;0.58), (0.16:0.33/0.33:0.53), (0.25;0.42/0.41;0.54;0.73). Meta-
tarsi HI with trichobothrium, 0.45/0.42/0.44. Tarsal organ I (0.31), IV (0.26). Tarsi
almost as long as metatarsi, I 0.90, IV 0.95. Tarsal claws I with 6 teeth.
d Palp: Figs 17-20, similar to R. neglectus (Figs 21, 22). Tibia with 1
retrolateral trichobothrium. Cymbium distally tapering, paracymbium retrolateral-
subterminal and hook-like (Fig. 20). Tegulum with conductor, tegular fold and one
tegular apophysis only. Dorsal part of tegulum bulging beside conductor (Fig. 17),
which is a retrolateral projection of the tegulum (Fig. 20), membranous and lobe-like.
Tegular fold (f): Figs 17, 18, transparent, lamelliform, as in R. neglectus (Fig. 21).
Tegular apophysis situated prolaterally, with three branches diverging in dorsal,
ventral and retrolateral directions (Figs 17, 18, 19; D, V, R). Dorsal branch with
concavity which tilts over the paracymbium in the expanded palp. Ventral branch
taxonomically important, long and curved, not straight as in R. neglectus. Retrolateral
branch short and blunt. Sperm duct runs in a loop through tegular apophysis. Embolar
base larger than in R. neglectus (Figs 18 vs. 21), distal part of embolus long and
slender, its tip narrow (Figs 17 vs. 22). Sperm duct orifice terminal. In R. neglectus
the embolar tip broadens and the sperm duct runs along one side.
Discussion: Robertus species occur mainly in Europe and North
America and centres of endemism are in Europe, North America and East Siberia
(Eskov 1987). R. calidus is the first African representative. In palpal structure and
trichobothriotaxy it is closely allied to the western palaearctic R. neglectus. They both
belong, together with R. scoticus Jackson and R. kuehnae Bauchhenss & Uhlenhaut,
to a species group with only | tegular apophysis which is strongly branched, and with
a small and hyaline conductor.
Comparative characters of R. neglectus: Carapace 0.94 mm long, 0.69 mm
wide. Legs 1423. Numbers of prodorsal/retrodorsal trichobothria of tibiae HIV: 1/3,
1/2, 2/2, 2/3, their respective positions (0.38/0.25;0.46;0.67), (0.34/0.40;0.68), (0.17;
0.40/0.40;0.63), (0.24;0.43/0.45;0.62;0.82). Metatarsi I-III with trichobothrium (0.50/
0.49/0.42). Tarsi almost as long as metatarsi, I 0.88, IV 0.92. Tarsal claws I with 6
teeth. d palp: Figs 21, 22.
TWO AFROMONTANE THERIDIIDAE 987
Fics 17-22
Robertus calidus n. sp. (17-20). Robertus neglectus (O.P.-Cambridge) (21, 22). Left male palp,
ventral (17), ventral-retrolateral (18), retrolateral (20, 21) and prolateral view (19, 22). Scale
lines 0.1 mm.
988 BARBARA KNOFLACH
ACKNOWLEDGEMENTS
I am grateful to Dr E. Heiss for collecting the material and for information.
Many thanks are due to Dr K. Thaler for discussion and literature and to Dr P. Merrett
for linguistic revision.
LITERATURE
Eskov, K.Y. 1987. The spider genus Robertus O. Pickard-Cambridge in the USSR, with an
analysis of its distribution (Arachnida: Araneae: Theridiidae). Senckenbergiana biol.
67: 279-296.
Hom, À. 1962. The spider fauna of the east African mountains. Zool. Bidr. Uppsala 35: 19-
204, pl. 1-6.
Levi, H.W. 1963. The spider genera Cerocida, Hetschkia, Wirada and Craspedisia (Araneae:
Theridiidae). Psyche 70: 170-179.
Levi, H.W. 1972. Taxonomic-nomenclatural notes on misplaced theridiid spiders (Araneae:
Theridiidae), with observations on Anelosimus. Trans. Am. Micros. Soc. 91: 533-538.
Levi, H.W. & L.R. Levi. 1962. The genera of the spider family Theridiidae. Bull. Mus. comp.
Zool. 127: 1-71, Figs 1-334.
MILLER, F. 1970. Spinnenarten der Unterfamilie Micryphantinae und der Familie Theridiidae
aus Angola. Publcoes cult. Co. Diam. Angola 82: 75-166.
SAARISTO, M.I. 1978. Spiders (Arachnida, Araneae) from the Seychelle Islands, with notes on
taxonomy. Ann. Zool. Fennici 15: 99-126.
SCHARFF, N. 1992. The linyphiid fauna of eastern Africa (Araneae: Linyphiidae) — distribution
patterns, diversity and endemism. Biol. J. Linn. Soc. 45: 117-154.
THALER, K. & K.H. STEINBERGER. 1988. Zwei neue Zwerg-Kugelspinnen aus Osterreich
(Arachnida: Aranei, Theridiidae). Revue suisse Zool. 95: 997-1004.
WIEHLE, H. 1937. Spinnentiere oder Arachnoidae VIII. 26. Familie: Theridiidae oder Hauben-
netzspinnen (Kugelspinnen). - Tierwelt Dtl. 33: 119-222.
REVUE SUISSE DE ZOOLOGIE, 102 (4) : 989-994; décembre 1995
Redescription and lectotype designation
of Aploparaxis pseudofilum (Clerc, 1902)
(Cestoda: Hymenolepididae)
Josef K. MACKO* & Claude VAUCHER**
* Parasitological Institute of the Slovak Academy of Sciences, Hlinkova 3,
040 01 Kosice, Slovak Republic.
** Muséum d’histoire naturelle, Case postale 6434, CH-1211 Genève 6, Switzerland.
Redescription and lectotype designation of Aploparaxis pseudofilum
(Clerc, 1902) (Cestoda: Hymenolepididae. - A. pseudofilum (Clerc, 1902)
is redescribed from original material stored in the Geneva Natural History
Museum and a lectotype is designated. Various cestodes previously named
A. pseudofilum are not conspecific with this species, characterized by a short
cirrus pouch.
Key-words: Aploparaxis pseudofilum - Cestodes - Hymenolepididae -
Taxonomy - Bird parasite.
INTRODUCTION
Aploparaxis pseudofilum (Clerc, 1902) is a relatively common parasite of the
woodcock Scolapax rusticola. Various authors (SPASSKIJ, 1963; SPASSKAYA, 1968;
DEBLOCK & RAUSCH, 1968) have treated under this name different hymenolepidids
from a wide range of hosts. According to BONDARENKO (1990a, 1990b), A. pseudo-
filum may represent a species complex. A. pseudofilum has been thoroughly redes-
cribed from new material which was compared with syntypes (BONDARENKO, 1990b).
CLERC’s (1902) description was based on several specimens, a single slide conserved
in the Geneva Museum contains two or three species. CLERC’s (1902) drawing of a
transverse section suggests that this is only a part of the original type series. It is
therefore important to designate a lectotype.
MATERIAL
Some of the syntypes are preserved in MHNG, the rest is probably lost.
Following information is available: slide No 13/78 “Haploparaxis filum (Goeze,
1782), Scolopax rusticola, Oural, M.G.V 5/8”. The geographic origin (Ural) of the
label authenticates this slide as original material of Clerc, who was at that time a
Manuscript accepted 13.04.1995.
990 JOSEF K. MACKO & CLAUDE VAUCHER
student of O. Fuhrmann at the Zoological Institute of Neuchatel University (see
CLERC 1902, 1903), whose collection is deposited in the Geneva Museum. The slide
bears the following identifications made by Dr. S. Bondarenko in 1980: A. pseudo-
filum, A. filum and A. ? parafilum.
The lectotype of A. pseudofilum designated berein is a specimen 48 mm long
and max. 660 um wide. It lacks the scolex and is not fully gravid. The pregavid
extremity has been slightly damaged when Clerc mounted the slide. The other
specimens are 58 mm long and max. 980 Mm wide (A. parafilum ?) and 13 mm long
and max. 290 Mm wide (A. filum), respectively. The slide contains two additionnal
fragments of A. pseudofilum, and two fragments which are unidentifiable.
Measurement are given in um unless otherwise stated.
DESCRIPTION OF THE LECTOTYPE
Coiled strobila without scolex and without gravid segments, ca 48 mm long. In
posterior portion, anteriorly to slightly deformed end, it is up to 660 wide. Young
proglottides show stained primordia of the reproductive organs. In the next portion of
strobila, 180 wide, testis and distal genital ducts start to differentiate. Testis 38-
61x103-128, transversely elongate, more or less constricted. In some segments
separating into 2 (Fig. 1). Cirrus pouch (CP) 17-24x72-113, exceptionally 121 long,
in male segments reaching or slightly exceeding beyond poral excretory ducts. In
hermaphroditic and postmature proglottids, CP not reaching excretory ducts (Figs 2,
3). Dorsal excretory ducts ca 4 in diameter, ventral ca 16. Vesicula seminalis interna
51-55 in longer diameter. Vesicula seminalis externa pyriform.
Genityl atrium situated on the margin in mid-portion of proglottis or at the
beginning of the second half of segment margin.
Cirrus cylindrical to club-shaped, 16-21x39-42, attenuated at base to 9-11.
Maximum width of not fully protuberant cirri ranging between 14.5 and 21. Distal
portion of cirrus ca 13 long, without observable ornamentation. Middle portion of
cirrus (ca 12 long) covered with distinct spines, 2-3 long. Spines at the basal portion
of cirrus very fine and poorly visible (Fig. 4).
Ovary more or less lobate, transversely elongate up to 159. Vitellarium 25-
28x34-41 (Fig. 2). Distal female ducts not observable. Uterus initially a transversely
elongate tube irregularly thick, saccular in the last segments (Fig. 5). Eggs unde-
veloped in the examined specimen.
Other two fragments of A. pseudofilum are no more than 5 mm long. Bursa
cirri as in lectotype. Width of cirrus up to 23. Uterus saccular, as in lectotype.
Embryophore indistinctly outlined, ca 51x43. Lateral embryonic hooks 16.
DISCUSSION
CLERC (1902) differenciated Aploparaxis pseudofilum from A. filum (Goeze,
1782) by the shorter cirrus pouch. CLERC (1903) considered A. pseudofilum a variety
of A. filum but pointed out the cirrus itself is longer in A. filum. Clerc did not mention
APLOPARAXIS PSEUDOFILUM (CLERC), 1902 991
Fic. 1-2
Aploparaksis pseudofilum (Clerc, 1902) lectotype. 1 - fragment of strobila with one and two
testes. Scale bar : 100 um. 2 - hermaphroditic segments showing variable structure of the ovary.
Scale bar : 100 um.
992 JOSEF K. MACKO & CLAUDE VAUCHER
Fic. 3-5
Aploparaksis pseudofilum (Clerc, 1902) lectotype. 3 - opening of the cirrus pouch with
protuberant cirrus. Scale bar: 40 um. 4 - cirrus. Scale bar: 20 um. 5 - portion of saccular uterus.
Scale bar : 100 um.
a host species but noted that the parasite was found together with A. filum. There is
no subsequent information on type-host.
Until recently, the name A. pseudofilum has been attributed to different species
(BONDARENKO, 1990b). According to this author, the taxon of Clerc was known in the
Soviet literature as A. sanjuanensis Tubangui & Masilungan, 1937, described from
Gallinago megala in the Philippines. Unfortunately, the type material of A. sanjua-
nensis is lost and the synonymy proposed by BONDARENKO (loc. cit.) cannot be
checked. Moreover, this author states that A. clerci Yamaguti, 1935, from Europe and
A. speudofilum are identical though without having examined material from A. clerci
due to unavailability of Japanese specimens.
From CLERC’s (1902) type material, probably only specimens from Scolopax
rusticola have been preserved; we select as lectotype of A. pseudofilum the 48 mm
long specimen on slide No 13/78. The specimen has the characteristic short cirrus
pouch, which extends slightly beyond the poral excretora ducts only in young
proglottides.
APLOPARAXIS PSEUDOFILUM (CLERC), 1902 993
ACKNOWLEDGEMENTS
This research was financed by the Slovak Grant Agency No 2/1364/95.
REFERENCES
BONDARENKO, S.K. 1990a. Type species of the genus Aploparaksis (Cestoda, Hymeno-
lepididae), A. filum, and its life cycle. Parazitologia 24: 379-389. (In Russian).
BONDARENKO, S.K. 1990b. Aploparaksis pseudofilum (Clerc, 1902) non Gasowska, 1931 and
its postembryonal development. Parazitologia 24: 509-517. (In Russian).
CLERC, W. 1902. Contribution à l’étude de la faune helminthologique de l’Oural. Zool. Anz. 25:
569-575.
CLERC, W. 1903. Contribution à l’étude de la faune helminthologique de l’Oural. Rev. suisse
Zool. 11: 241-368.
DEBLOCK, S. & R. L. RAUSCH. 1968. Dix Aploparaksis (Cestoda) de Charadriiformes d’ Alaska,
et quelques autres d’ailleurs. Ann/s Paras. hum. comp. 43: 429-448.
SPASSKAYA, L.P. 1966. Cestodes of birds in the USSR (Hymenolepididae). Moscow Izdalstvo
Akademya Nauk SSSR, 98 pp. (In Russian).
Spassky, A.A. 1963. Hymenolepdidae - tapeworms of free living domestic birds (Part 1). In
“Skrjabin, K.I. (Ed.): Essentials of Cestodology. Moscou, Izdavatelstvo Akademya
Nauk SSSR 2, 417 pp. (In Russian).
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7
REVUE SUISSE DE ZOOLOGIE, 102 (4) : 995-1044; décembre 1995
A review of the Passandridae of the world
(Coleoptera, Cucujoidea). IV. Genus Ancistria
Daniel BURCKHARDT* & Stanislaw Adam SLIPINSKI**
* Muséum d'histoire naturelle, Case postale 6434, CH-1211 Genève 6, Switzerland.
** Museum and Institute of Zoology, Wilcza 64, 00-679 Warszawa, Poland.
A review of the Passandridae of the world (Coleoptera, Cucujoidea). IV.
Genus Ancistria. - The genus Ancistria 1s revised to include 32 valid
species, 14 of which are described as new. One genus, 3 subgenera, 15
species and | variety are synonymised, and 1 primary junior homonym at
species group level is replaced by a new name. The species are diagnosed
with external morphological, genital and meristic characters, keyed and
illustrated. Ancistria is restricted to the Old World where it is most speciose
in the tropics. Ancistria retusa (Fabricius), the only common species, has a
distribution almost congruent with that of the genus; the other species are
much more restricted. The biogeographical relationships are briefly
discussed.
Key-words: Coleoptera - Passandridae - Ancistria - taxonomy - Old World
tropics.
INTRODUCTION
The Passandridae or Passandrinae (of Cucujidae) constitute a small taxon of
some 100 species placed in 9 genera. The present paper is the ultimate in a series
revising its taxonomy (SLIPINSKI, 1987, 1989; BURCKHARDT & SLIPINSKI, 1991, 1995). It
deals with the Old World genus Ancistria which is predominantly tropical. GROUVELLE
(1913a) revised the genus recognising 29 species and 1 variety. He described each
species in detail, listed distributional data and provided a key to species. Based on
differences on the head, he subdivided the genus into the four subgenera Obancistria,
Anancistria, Ancistria s. str. and Parancistria. Little has been added since. GROUVELLE
(1913b, 1919) published the descriptions of A. sauteri, A. emarginata and A. africana,
KESSEL (1921) that of A. bakeri and HELLER & GUNTHER (1936) that of A. tenera.
HETSCHKO (1930) summarised published information up to that date. Not mentioned,
neither by GROUVELLE (1913a) nor by HETSCHKO (1930), is Bostrichoides angulatus
described by MONTROUZIER (1855) from Woodlark Island; the species belongs, as the
Manuscript accepted 16.05.1995.
996 DANIEL BURCKHARDT & ADAM SLIPINSKI
study of type material showed, to Ancistria and is conspecific with Ancistria retusa
(Fabricius).
Despite this relatively good base, it remains currently difficult, if not impossible,
to identify Ancistria species. Grouvelle had only a very limited material at hand which
led him to overestimate the taxonomic value of some external morphological
characters. His species concept is typological and does not take into account the
considerable morphological variability which, as in related genera, is to expect in
Ancistria.
The biology of Ancistria spp. is largely unknown. They are probably parasitic,
as other passandrids, and may be specialised on bark and wood boring Coleoptera.
Several adults have been extracted from tunnels in various trees. HAWKESWOOD (1987)
mentions that, in Australia, adults of A. retusa may be attracted to light in summer.
The present study is based on extensive material, including types of all
described taxa. For the first time, the genitalia are examined and it is concluded that, as
in other passandrid genera, they provide good characters for defining species. Other
important characters are the relative lengths of head and pronotum, the shapes of the
lateral frontal processes and frontoclypeal depression, the relative length of the
mesofemur and details on the antennae and the elytral apex. The last is sometimes
subjected to sexual dimorphism. A key for species identification and illustrations are
provided to supplement the descriptions which diagnose the species.
MATERIAL AND METHODS
Material from following institutions and collections was examined and/or is
mentioned in the text:
ANIC Australian National Insect Collection, CSIRO, Canberra (J. F. Lawrence);
BMNH Natural History Museum, London (R.J.W. Aldridge, M. Kerley, R.D. Pope,
S. L. Shute);
BPBM _ Bernice P. Bishop Museum, Honolulu (J.A. Samuelson);
DEIC Deutsches Entomologisches Institut, Eberswalde (L. Zerche);
HNHM Hungarian Natural History Museum, Budapest (O. Merkl);
ISNB Institut Royal des Sciences Naturelles de Belgique, Bruxelles (R. Damoiseau);
KYUN Kyushu University, Kyushu (T. Ueno);
MCSN Museo Civico di Storia Naturale "Giacomo Doria", Genova (R. Poggi);
MHNG Muséum d'histoire naturelle, Genève;
MNHN Muséum National d'Histoire Naturelle, Paris (N. Berti);
MRAC Musée royal de l'Afrique centrale, Tervuren (J. Decelle);
NHMB Naturhistorisches Museum, Basel (M. Brancucci, R. Heinertz);
QMBA Queensland Museum, Brisbane (G. Monteith);
RMNH _ Rijksmuseum van Natuurlijke Historie, Leiden (J. Krikken);
SAMC South African Museum, Cape Town (R. Whitehead);
SMTD Staatliches Museum für Tierkunde, Dresden (R. Krause, O. Jäger);
USNM _ National Museum of Natural History, Smithsonian Institution, Washington,
D.C. (J. Pakaluk);
ZMHB Zoologisches Museum, Humboldt-Universitàt, Berlin (M. Uhlig);
ZMPA Museum and Institute of Zoology, Polish Academy of Sciences, Warszawa;
ZMUC Zoological Museum, University of Copenhagen (O. Martin).
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 997
Measurements were made using a filar micrometer as follows: body length, from
apical margin of clypeus to elytral apex, excluding the mandibles; head length, from
apical margin of clypeus to occipital groove; head width, maximum width across the
eyes; joint width of lateral frontal processes, cf. figs 96, 150: B + B'; width of
frontoclypeal depression, cf. figs 96, 150: A; temple length, maximum length, in
profile, in a line parallel to the dorsal head margin, from occipital groove to posterior
eye margin; eye diameter, maximum length along same axis as temple length; pronotal
length, from base to apex along the middle; pronotal width, across maximum width;
elytral length, along suture including scutellum; elytral width, across maximum width
of both elytra together. Where many specimens were available measurements were
taken from a selection of specimens representing both sexes and exhibiting maximum
range of size and form.
Ancistria Erichson
Ancistria ERICHSON, 1845: 305. Type species Colydium retusum Fabricius, by monotypy.
Prionophora WESTWOOD, 1848: 85. Type species Prionophora cylindrica Westwood, by
monotypy. Synonymised with Ancistria by GEMMINGER & VON HAROLD, 1868: 870.
Bostrichoides MONTROUZIER, 1855: 17. Type species Bostrichoides angustatus Montrouzier, by
monotypy. Syn. n.
Ancistria (Anancistria) GROUVELLE, 1913a: 436. Type species Ancistria beccarii Grouvelle, here
designated. Syn. n.
Ancistria (Obancistria) GROUVELLE, 1913a: 436. Type species Ancistria (Obancistria) longior
Grouvelle, by monotypy. Syn. n.
Ancistria (Parancistria) GROUVELLE, 1913a: 437. Ancistria (Parancistria) bouchardi Grouvelle,
here designated. Syn. n.
Diagnosis. Small to medium sized, cylindrical, elongate Passandridae (3.4-14.6
mm long). Head with distinct median and admedian grooves and more or less depressed
frontoclypeus. Jugular plates widely separated, mentum large, triangular. Antenna | 1-
segmented, segment | bearing | to several dorsal grooves; apical 5 or 6 segments
flattened, asymmetrical, forming a saw-shaped, sometimes indistinct club. Basal tarsal
segment very long, sometimes longer than tibia; tibiae bearing a long hooked apical
spur. Elytra with costa 3 more or less strongly inflated subapically; interval 1 present,
grooved, complete, intervals 2-6 grooved when present, often incomplete.
Description. Colour light, reddish or chestnut brown to almost black, sometimes
bicoloured; surface shiny or mat. Usually glabrous, in A. pilosa head with very short,
pronotum and elytra with fine erect setae.
Head weakly convex or almost flat, bearing a shallow or deep frontoclypeal
depression which is delimited laterally by the lateral frontal processes which can be
careniform or flattened; mid line partly or completely grooved; admedian lines short to
moderately long, almost straight to strongly curved or bent; admedian lobes short to
moderately long, narrowly triangular to almost lamellar, parallel-sided; occipital
transverse groove wide and deep; punctation fine to coarse, laterally often strigose. Eye
moderate-sized to large. Antenna 11-segmented; segment 1 usually flattened dorsally
998 DANIEL BURCKHARDT & ADAM SLIPINSKI
Fics 1-6
Ancistria spp. 1, A. apicalis; 2, A. brancuccii; 3, 4, A. retusa; 5, 6, A. bakeri. 1-3, head, frontal
view; 4, bases of pronotum and elytra; 5, elytra, lateral view; 6, elytra, oblique rear view. (Scale
bars 1, 3-6 = 0.2 mm; 2 = 0.1 mm).
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 999
Fics 7-12
Ancistria spp., oblique rear view of elytra. 7, A. apicalis 2; 8, A. brancuccii 3 ; 9, A. nicolettae 2;
10, A. tarsalis 9,11, A. retusa 3; 12, A. retusa 2. (Scale bars = 0.2 mm).
1000 DANIEL BURCKHARDT & ADAM SLIPINSKI
with | to several dorsal grooves which can be more or less distinct; segment 2 normally
subglobular, sometimes oval; segment 3 oval or clavate; segments 4 and 5 symme-
trically or asymmetrically oval; segments 6 or 7 to 11 flattened, asymmetrical, forming
saw-shaped club, in some species club very indistinct; segments 3-11 irregularly setose,
club segments in addition apico-ventrally with well-defined region bearing very short
setae producing felt-like appearance; segment 11 subrectangular, irregularly rounded
apically.
Pronotum longer than wide, distinctly narrowed basally, widening towards the
middle or almost the apex, always margined laterally; border at base absent, or partly or
fully developed. Prosternum with sternopleural sutures fully developed; prosternal
process parallel-sided, strongly curved towards mesosternum, flat.
Elytra with costa 3 very weakly to very strongly inflated subapically, sometimes
in females more than in males, costae 7 and 8 merging subapically forming a lateral rim
which fuses with the apical margin; interval 1 completey grooved, 2-6 completely or
partly grooved, 6 sometimes obsolete, intervals 2-6 usually reduced apically; apex
truncate or emarginate, sometimes with V or U-shaped incision along suture, with or
without sutural tooth.
Ventral side of head with widely separated jugular plates, mentum large, trian-
gular. Punctation on ventral side usually coarse. Mesosternum small, foveolate or flat,
smooth or punctate. Metasternum with long median groove apically, irregularly
punctate. Abdominal ventrites simple, punctate; last ventrite without apical groove.
The male genitalia provide useful means of species identification, even though
there is a certain amount of variation. The aedeagus has an elongate median lobe with
ventral strut and flagellum, and the tegmen which consists of a moderately sclerotised
dorsal plate and ventral, setose parameres. The apical piece (in ventral view) of the
median lobe, the parameres, and sternite and tergite VIII are figured for all species
where males are known. Females have a moderately sclerotised, strongly convoluted
spermatheca which is figured were known.
Comments. Ancistria shares with Passandrella, Taphroscelidia and Scalidiopsis
the presence of a grooved median line on the head and the basal metatarsomere which
is longer than tarsomere 2. Without a phylogenetic analysis of the whole family (in
prep.) it is, however, not possible to say whether these characters express close
phylogenetic relationship. Ancistria differs from the 3 other genera in the absence of a
pair of setal brushes on the lower head surface, the absence of deep pits anterior to the
procoxae, the absence of a transverse groove on the last abdominal ventrite, and the
presence of transversely rugose costae on the elytra.
Based on the structure of the anterior portion of the head GROUVELLE (1913a)
divided Ancistria into the four subgenera Obancistria, Anancistria, Ancistria s. str. and
Parancistria. These characters are, however, not as clear-cut and stable as indicated by
Grouvelle. The subdivision is therefore not adopted in the following and the subgenera
are synonymised.
At present, 32 species are recognised within Ancistria.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1001
KEY TO SPECIES OF Ancistria ERICHSON
Bronotalllensthn/widthiratio)lesssthanelkA ern RR e CAEN CE 2
Bronotallencth/vidthrationmore than 1A PE er ee en. 3
Frontoclypeus flattened, admedian lines on head widened anteriorly to
form circular to oval depressions (fig. 43). Genitalia as ın figs 44, 45, 47-
JO NewiGumea Sumatra Malaysia. nen. nn. ee men we beccarii
Frontoclypeus horn-shaped, admedian lines on head not markedly
widened anteriorly (Fig. 74). Genitalia as in figs 75-77. Laos. ........ cornuta
Mesotibia about 0.8 times as long as or shorter than first tarsomere of
MG SLES SHE. BE he MA PERRET RAS Facto ives RT Ante Re ee 4
Mesotibia as long as or longer than first tarsomere of mid-legs. ............ 6
Frontoclypeal depression deep, well-delimited, admedian lines short and
deep, admedian lobes short, acutely triangular (figs 90, 96). Genitalia as
mets OI 94 /ava Sumatra New Guinea ri 4s wre: foraminifrons
Frontoclypeal depression shallower, admedian lines longer and shal-
lowergadmedianalobesilonsi(i12S83 4175) ARE Beeren RARE 5
Lateral margin of elytra notched in the middle (figs 5, 6). Genitalia as in
HUSSHS4=9 Oh BomeosSumatra:M3 nr i der, Mawes RRs SS tee bakeri
Lateral elytral margin not notched in the middle. Genitalia as in figs 173,
IW GeSumatray Borneo Mal HE MON Se A BEE tarsalis
Pronotum and elytra covered in erect, fine setae (clearly visible at 10 x
magnification). Genitalia as in figs 140, 141. Taiwan. ................ pilosa
Bronetumyandıelytra withoutsetaen. rn. sr oe ce A n 7
Admedian lines strongly curved or angular, admedian lobes, therefore,
lamellar and suddenly narrowing towards apex (figs 2, 55). ............... 8
Admedian lines straight or weakly curved, admedian lobes, therefore,
regularly or irregularly cuneate gradually narrowing towards apex (figs
SSD IERI ROOTS AGE O O E MER od. A _ 12
Large species, body length 8.0 mm. Pronotum less than 1.5 times as long
asiwide Genitaliafassimifi 990997 Sika e Re EURE bostrychoides
Small species, body shorter than 7.0 mm. Pronotum more than 1.7 times
asl onesasiwi desti TEE a TEE TER EHI AMC Be EC 9
Mesotibial apex angular externally lacking a spiniform process. Head
more than 1.7, and pronotum more than 2.2 times as long as wide.
Geritaliaa sinti gs MS MIS SNeW Guineate sn longicapitata
Mesotibial apex with distinct external spine. Head less than 1.6, and
pronotumilessithani2:0 times along as wide BRR PE EE ee eee 10
Pronotum bordered at base, less than 1.8 times as long as wide. Genitalia
as impress 35=l 37 New Guinea. te 5 ar aie ein. oe een papuana
Pronotum not or incompletely bordered at base, more than 1.8 times as
IOMEFASIWA CS Shire: RITI A N SO af
Body black. Elytral length/width ratio less than 3.8. Lateral frontal
processes on head flattened (figs 2, 63). Genitalia as in figs 64-66. Nepal.
DO D D AE CARINA OS N ATP TE Le brancuccit
1002
DANIEL BURCKHARDT & ADAM SLIPINSKI
Body chestnut brown. Elytral length/width ratio more than 4.0. Lateral
frontal processes on head careniform (fig. 107). Genitalia as in figs 108-
LUO. ER Niet ies aac Eee PRO D M ic a 0 410 0 0 co © lewisi
Lateral frontal processes of head careniform, subcareniform or rarely
flattened, narrow, their joint width less than 0.9 times that of fronto-
elypealldepression digs 3, 143, 178) MN en ee ce ee eee 13
Lateral frontal processes of head flattened, wide, their largest joint
width, in dorsal view, more than 0.9 times that of frontoclypeal de-
pression: (He M) EU ET data a Baar SAN ARRETE 18
Admedian lobes subcarinate anteriorly, ending level with clypeal base
and lateral frontal processes (fig. 51). Elytra reddish with bases and
median transverse band black. Genitalia as in figs 52, 53. Vietnam. . . bicolorata
Admedian lobes sometimes raised but not carinate anteriorly, always
much shorter than lateral frontal processes anteriorly. Elytral colour different. 14
Elyitraswithinterval 4 Shorterthan 5) Ge. 1 12) Re Re 15
Elytra with apical length of intervals decreasing gradually from 3 to 6 (fig. 9). 17
Lateral frontal process of head careniform (figs 3, 146, 148, 150). Costa
3 on elytra strongly inflated subapically (figs 11, 12). Genitalia as in figs
151-164. Widely distributed from Australia, through tropical Asia to
ATC E La RELAIS EIA retusa
Lateral frontal process of head flattened (figs 143, 179). Costa 3 on elytra
vem»weakly,inflatedisubapically 2.2.2.0. sr. sr Der ee 16
Antennae 6-segmented (fig. 142). Pronotum not bordered basally.
Genitaliafas inte 144 Japan... 2 cae 0 Ba reitteri
Antennae 5-segmented (fig. 178). Pronotum bordered basally. Genitalia
as. ın-fes 180-1825 Tavast mem seine cate eine ee tenera
First tarsomere of mid-leg long, as long as tarsomeres 2-5. Large, body
longer than 9.0 mm, black. Elytra never emarginate apically. Genitalia as
in fig. 172. New Guinea, Ambon, Ceram, Solomon Island............ strigosa
First tarsomere of mid-leg short, as long as tarsomeres 2-3. Small, body
shorter than 8.5 mm, brown. Elytra with apical tooth in females. Geni-
talia as in figs 129-132. Malaysia, Palawan, Borneo, Sumatra. ...... nicolettae
Lateral frontal processes with large anterior tubercle (fig. 98). Genitalia
asset, 99 sNews Guinea: + str er. Ar grouvellei
Lateral frontal processes without large anterior tubercle (fig. 87)........... 19
Pronotum more than 1.9 times as longias wide> tt. 555 were 20
Bronotum less’than! 1:8 times ‘as long’asiwider 22.2. 22.2 se ern eee 22
Antennal club 6-segmented (fig. 86). Genitalia as in fig. 83. "Southeast
Saas cedar oe. wirt see addi ee c 6 0 © o à fabricii
Antennal'club:S-seemented (fig. 183). Sees ger il
Body colour brown. Body length shorter than 4.0 mm. Genitalia as in
fie: 185. New Guinean he. 2. TE tenuis
Body colour almost black. Body length more than 5.0 mm. Genitalia as
in. 11081255126: Nepal... > OR See. Ree oo nepalensis
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1003
22 Antennal segment 1 with 2 wide, subequal, dorsal grooves, delimited by
3 subparallel ridges (figs 58, 59). Genitalia as in figs 60, 61. Sumatra.
eg ee ire hte: se ansehe bouchardi
- Antennalisesmentsldifferen@ ne here soe eerie ae er ioe oer 23
23 iliemplezlensth/eyefdiameteriratio moretthanilM/ Reni ae 24
- temple length/eyeidiametenmatiogess than) es eae ee 25
24 Large species, body longer than 12.0 mm. Pronotum slender, 1.8 times as
long as wide, with narrow puncture-free median longitudinal stripe.
Genmitaliavassmyhiessis=1 Ss ehilippines: ere na EE ieee ee alternans
- Small species, body shorter than 6.0 mm. Pronotum wide, 1.5 times as
long as wide, evenly covered in punctures. Genitalia as in figs 167-169.
ES ORME OOM SAPOLCs pos an go seine dali nie le ENS furs stricta
25 Antennal club 5-segmented (fig. 100). Pronotum bordered at base,
bordeuneidistnet laterally.andimediallyao ao... 0.0 men... 26
- Antennal club 6-segmented (fig. 67). Pronotum not, indistinctly or in-
completelyaborderedaarbaserr vr ee NO IN TI 29
26 Mandibles strongly transversely rugose dorsally (fig. 101). Genitalia as
HDL 10s OPEN SAINTE ESS RTE e indica
= Mandibles not rugose but punctured dorsally (fig. 79). .................. 27
24] Body longer than 6.0 mm. Genitalia as in figs 80, 81. "British Bootang". costata
- Bodyzshorterthan 3:03mm so I N 28
28 Frontoclypeal depression with small tubercle (fig. 28). Elytra bicoloured.
Genttalrarassnat 8529-31 Indian ee o assamensis
- Frontoclypeal depression without tubercle (fig. 40). Elytra completely
black. Genitalia as in figs 40, 41. New Guinea. ..................... basseti
29 Antennal segment 11 distinctly longer than wide (fig. 67). ............... 30
- Antennal segment 11 about as long as wide (fig. 24)..................... 3
30 Elytral apex emarginate. Genitalia as in figs 69-72. Philippines, Sumatra.
PAH: Stia pre BW EMME) ioni atk SEE ER concava
- Elytral apex truncate. Genitalia as in figs 118-122. Malaysia, Borneo. . . . micros
31 Mandibles punctate (fig. 19). Elytra entirely or apically dark chestnut
brown, weakly emarginate apically. Genitalia as in figs 20-23, 25, 26.
Japan Gna Vic tna Ren oa apicalis
- Mandible strongly transversely rugose (fig. 83). Elytra uniformly dark
brown or black, strongly emarginate apically. Genitalia as in figs 84, 85.
AVATAR © nt a ree EE aad arson e E cg es AN emarginata
Ancistria alternans Grouvelle (Figs 13-18)
Ancistria (Ancistria) alternans GROUVELLE, 1913a: 472. Holotype d, without locality data
(Castelnau) (MNHN); (examined).
Description. Length 12.6-14.6 mm. Dark brown to almost black, apical half of
elytra in d dark chestnut brown, in £ almost black; surface mat. Head (fig. 14) 1.3-1.4
1004 DANIEL BURCKHARDT & ADAM SLIPINSKI
times as long as wide, coarsely, irregularly punctured; median line completely or almost
impressed to base; admedian lines moderately short, only weakly curved; admedian
lobes narrowly triangular; lateral frontal lobes flattened, their joint width 2.3-2.8 times
that of frontoclypeal depression. Eye relatively small; temple length/eye diameter ratio
as 1.8-2.4. Antenna (fig. 13) with segment | bearing 1 deep and 1 or 2 shallow grooves;
segment 2 subglobular; segments 3-5 oval; segments 6-11 strongly flattened, forming a
distinct club; segment 11 wider than long. Pronotum 1.7-1.8 times as long as wide, not
bordered at base, widening in basal third, with subparallel margins in apical third;
punctation coarser than on head, leaving a puncture-free longitudinal median stripe.
Elytra 3.2-3.5 times as long as wide; 2.0-2.1 times as long as pronotum; costa 3 more,
costa 5 less inflated subapically; intervals gradually shortened from 3 to 6; apex in d
weakly, in 2 strongly emarginate, bearing a smaller or larger sutural tooth. Genitalia as
in figs 15-18.
Distribution. The only recorded specimen has no locality data (GROUVELLE, 1913a);
HETSCHKO (1930) mentions the species questionably from Australia for which is no evidence.
Material examined. Holotype 4, without locality data (MNHN). Philippines: 1 2, Masbate Island,
Aroroy, 28.vili.1917, Böttcher (ZMHB).
Fics 13-18
Ancistria alternans. 13, antenna; 14, head, dorsal view; 15, parameres; 16, median lobe, apical
piece; 17, spermatheca; 18, ostium bursae.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1005
Ancistria apicalis Reitter (Figs 1, 7, 19-26)
Ancistria apicalis REITTER, 1889: 314. Lectotype d, Japan: Yuyama in Higo, 1883, G. Lewis,
labelled "Nagasaki, 22.v.-3.vi.1881, G. Lewis" (BMNH); here designated (examined).
Ancistria (Parancistria) apicalis Reitter; GROUVELLE, 1913a: 479.
Ancistria (Parancistria) fouqueti GROUVELLE, 1913a: 481. Holotype 9, Vietnam: Tonkin, Pha-
Vi, Fouquet (MNHN); (examined). Syn. n.
Description. Length 5.3-7.2 mm. Black, legs dark brown, elytra entirely or
partly chestnut brown towards the apex; surface mostly mat. Head (figs 1, 19) 1.2-1.3
times as long as wide; finely, laterally strigosely punctured; median line impressed
almost to base; admedian lobes narrowly triangular; lateral frontal processes flattened,
their joint width 2.6-3.4 times that of frontoclypeal depression. Dorsal surface of
mandibles punctate. Eye moderately large; temple length/eye diameter ratio as 0.9-1.3.
Antenna (fig. 24) with segment | bearing 2 distinct grooves; segment 2 subglobular;
segment 3 clavate; segments 4 and 5 asymmetrically oval; segments 6-11 flattened,
forming indistinct club; segment 11 as long as wide. Pronotum 1.7 times as long as
wide, incompletely or indistinctly bordered at base, strongly widened in basal half, with
subparallel margins in apical half; punctation coarser than on head, density uneven
leaving a puncture-free longitudinal band in basal two thirds. Elytra 2.8-3.0 times as
long as wide, 1.8-2.0 times as long as pronotum; costa 3 weakly inflated subapically;
interval 3 longer than 4 which is shorter than 5; weakly emarginate apically with small
sutural tooth (fig. 7). Genitalia as in figs 20-23, 25, 26.
Distribution. Recorded as A. apicalis from Japan (REITTER, 1889; GROUVELLE, 1913a;
HETSCHCKO, 1930) and as A. fouqueti from Vietnam (GROUVELLE, 1913a; HETSCHKO, 1930).
Material examined. Japan: lectotype and paralectotype 14, 19 (BMNH, HNHM) of A. apicalis;
16, Koike, Nakayama-chö; Ehime Prefecture, 10.v.1964, S. Hisamatu (ANIC); 1 specimen,
Kyushu, Kagoshima Pref., Mount Hoyoshidake, 1-2.v.1991, T. Ueno (KYUN). Vietnam:
holotype 9 of A. fouqueti (MNHN); 16, 19, Tonkin, area of Hoa-Binh, 1928, A. de Cooman
(BMNH, MNHN), 2 specimens, Tonkin, Thanh Moi, M. Perrot (MHNG). China: 29, Sichuan,
Mount Emei, 600-1050 m, 5-19.v.1889, L. Bocàk (ZMPA).
Comments. According to REITTER's (1889) description of A. apicalis the type
material was collected at Yuyama in Higo. The lectotype in the BMNH is labelled
"Nagasaki"; this label has been added subsequently and is certainly wrong (cf.
comments to A. reitteri). A paralectotype in HNHM bears the same data as mentioned
in the original description (REITTER, 1889).
Ancistria assamensis sp. n. (Figs 27-31)
Description. Length 4.9-6.2 mm. Dark chestnut brown to black with apical half
of elytra light brown to brown; surface mostly mat. Head (fig. 28) 1.2 times as long as
wide, finely punctate; median line impressed almost to base; admedian lines long,
impressed, weakly curved, admedian lobes long, narrowly triangular; lateral frontal
processes flattened, their joint width 1.4-1.7 times that of frontoclypeal impression
which bears a small tubercle in the middle. Eye moderately large, temple length/eye
diameter ratio as 1.0-1.2. Antenna (fig. 27) with segment 1 bearing 2 fine dorsal
1006 DANIEL BURCKHARDT & ADAM SLIPINSKI
Fics 19-26
Ancistria apicalis. 19, head, dorsal view; 20, parameres; 21, median lobe, apical piece; 22, 23,
25, spermatheca; 22, holotype of A. fouqueti; 23, with developed gland, which is usually missing
in KOH treated specimens; 24, antenna; 26, sternite and tergite VIII.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1007
grooves; segment 2 subglobular; segments 3 and 4 oval; segments 5 and 6 asym-
metrically widened towards apex; segments 7-11 flattened, forming club; segment 11
slightly longer than wide. Pronotum 1.7 times as long as wide, entirely bordered at
base, strongly widened to middle, apical half with subparallel margins; punctation
coarser than on head, sparser on disk. Elytra 3.3-4.1 times as long as wide, 2.1-2.2
times as long as pronotum; costa 3 inflated subapically, costae 7 and 8 fused and
inflated subapically; intervals 4 and 6 subequal, both shorter than intervals 3 and 5;
apex emarginate with small sutural tooth. Genitalia as in figs 29-31.
Material examined. Holotype d, India: Assam, Sadiya Division, 10.vi.1932, ex Termi-
nalia myriocarpa, Dehra Dun collection (BMNH). Paratype d, India: same data as holotype but
10.ix.1932 (MHNG).
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Fics 27-31
Ancistria assamensis. 27, antenna; 28, head, dorsal view; 29, sternite and tergite VIII; 30,
parameres; 31, median lobe, apical piece.
1008 DANIEL BURCKHARDT & ADAM SLIPINSKI
Ancistria bakeri Kessel (Figs 5, 6, 32-38)
Ancistria (Obancistria) bakeri KESSEL, 1921: 32. Holotype 9, East Malaysia: Borneo, Sabah,
Sandakan, F. Baker (USNM); (examined).
Description. Length 4.9-5.4 mm. Dark brown to almost black, legs lighter
brown, abdominal venter sometimes light brown; medio-lateral notch on elytra semi-
transparent, light brown, strongly contrasted from remainder of elytra; surface mostly
mat. Head (fig. 33) 1.0-1.1 times as long as wide, densely covered in fine strigose
punctures; basal line indistinct at base, well-defined at apex; admedian lines moderately
long, shallow, almost straight, admedian lobes large, triangular; lateral frontal processes
flattened, their joint width 0.5-0.8 times that of frontoclypeal depression, which is very
shallow. Eye large, temple length/eye diameter ratio as 0.3-0.4. Antenna (fig. 32) with
segment | bearing 2 fine dorsal grooves; segment 2 oval; segment 3 clavate; segment 4
oval; segments 5-11 flattened, forming club; segment 11 distinctly longer than wide.
Fics 32-38
Ancistria bakeri. 32, antenna; 33, head, dorsal view; 34, parameres; 35, median lobe, apical
piece; 36, sternite and tergite VIII; 37, ostrium bursae; 38, spermatheca.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1009
Pronotum 1.8-2.0 times as long as wide, not bordered at base, relatively narrow at base,
widening towards apical third, from there to apex narrowing; punctation coarser and
more spaced than on head, not markedly sparser on disk. Elytra 3.9-4.5 times as long as
wide, 2.2-2.3 times as long as pronotum; costal margin notched in the middle (fig. 5),
with a circular area membranous; costae 3 and 4 fused subapically, inflated, fused
costae 7 and 8 ending in a tooth; intervals 3-6 increasing in length apically; strongly
emarginate apically with a sutural tooth (fig. 6). Genitalia as in figs 34-38.
Distribution. Reported from Borneo (KESSEL, 1921; HETSCHKO, 1930). Material exa-
mined. Borneo: 16, 59, including holotype, Sabah, Sandakan (USNM, MHNG). Sumatra: 5d,
Sinabang, Sima, 11.1913, E. Jacobson (RMNH, MHNG, ZMPA).
Ancistria basseti sp. n. (Figs 39-42)
Description. Length 4.2-4.9 mm. Dark brown, tibiae and tarsi brown; surface
mostly mat. Head (fig. 40) 1.2 times as long as wide; relatively densely somewhat
strigosely punctate; median line impressed becoming indistinct towards base; admedian
lines moderately short, weakly curved, admedian lobes triangular, pointed apically;
lateral frontal processes flattened, their joint width 1.9-2.4 times that of frontoclypeal
depression. Eye large, temple length/eye diameter ratio as 0.8-1.0. Antenna (fig. 39)
with segment 1 bearing 2 distinct dorsal grooves; segment 2 subglobular; segments 3-6
oval; segments 7-11 flattened, forming a well-defined club; segment 11 distinctly
longer than wide. Pronotum 1.7 times as long as wide, distinctly bordered at base,
strongly widened towards the middle from there to apex weakly narrowing; coarsely,
slightly irregularly punctured. Elytra 3.4 times as long as wide, 2.2 times as long as
pronotum; costa 2 longer than 3 apically, costa 3 weakly swollen subapically; intervals
42
Fics 39-42
Ancistria basseti. 39, antenna; 40, head, dorsal view; 41, spermatheca; 42, ostium bursae.
1010 DANIEL BURCKHARDT & ADAM SLIPINSKI
5 and 6 subequal, longer than 4 but shorter than 5; apex not emarginate, bearing an
indistinct sutural tooth. Genitalia as in figs 41, 42.
Material examined. Holotype 2, Papua New Guinea: Morobe Province, Wau, Mount
Kaindi, 1150-2300 m, 11-25.1.1993, flight intercept trap in Caldcluvia, Y. Basset (MHNG).
Paratype 9, Papua New Guinea: Morobe District, Kunai Creek, 1250 m, 26.viii.1963, Malaise
trap, J. Sedlacek (BPBM).
Ancistria beccarii Grouvelle (Figs 43-49)
Ancistria beccarii GROUVELLE, 1883: 277. Lectotype ©, Aru Island: Wokan, 1873, O. Beccari
(MCSN); here designated (examined).
Ancistria (Anancistria) beccarii Grouvelle; GROUVELLE, 1913a: 439.
Ancistria (Anancistria) arrowi GROUVELLE, 1913a: 437. Holotype, Moluccas: Makian and Kajoa
Islands, J. Wallace, Kai, Pascoe collection 93-60 (BMNH); (examined). Syn. n.
Ancistria (Anancistria) castelnaui GROUVELLE, 1913a: 441. Holotype d, Malacca, Castelnau
(MNHN); (examined). Syn. n.
Description. Length 8.2-11.4 mm. Black, elytra sometimes chestnut brown
apically; surface mostly shiny. Head (fig. 43) 0.8-0.9 times as long as wide, finely to
coarsely punctate; median line basally weakly, apically strongly impressed, admedian
lines moderately long, anteriorly suddenly widened to form circular or oval
impressions; admedian lobes narrowly triangular anteriorly; lateral frontal processes
flattened, their joint width 1.1-1.3 times that of frontoclypeal depression. Eye large,
temple length/eye diameter ratio as 0.5-0.8. Antenna (fig. 46) with segment 1 bearing 2
large dorsal grooves; segment 2 small, subglobular; segments 3-5 elongate, widest
subapically, each segment wider than preceding one; segments 6-11 strongly flattened,
forming an indistinct club; segment 11 slightly longer than wide. Pronotum 1.3 times as
long as wide, distinctly bordered at base, strongly widening towards apex; punctation
coarse, sparser on disk with punctation-free longitudinal band in the middle. Elytra 2.5-
2.8 times as long as wide, 2.1-2.2 times as long as pronotum; in male costa 3 forming
large ridge subapically, female similar but ridge of costa 3 more inflated and oblique
subapically; interval 3 longer than intervals 5 and 6 which are both longer than 4
apically; apex not emarginate. Genitalia as in figs 44, 45, 47-49.
Distribution. Recorded as A. beccarii from Aru Island and New Guinea: Ramoi
(GROUVELLE, 1883), as A. arrowi from the Moluccas: Makian and Kajoa Islands, and as A.
castelnaui from Malaysia (GROUVELLE, 1913a; HETSCHKO, 1930). Material examined. Malaysia:
holotype d of A. castelnaui, Malacca; 19, Perat, Chenderoh, 16.11.1935, ex Myrtaceae (BMNH).
Sumatra: 16, Tjinta Radja, T. Nevermann (USNM). Aru Island: lectotype 2 of A. beccarii
(MCSN); 19, same, Unreiuning, 1884, C. Ribbe (ZMHB). New Guinea: paralectotypes of A.
beccarii 22, Ramoi, ii. O. Beccari (MCSN, MNHN); 22, Papua New Guinea, Stony L. A.,
17.iv.1969, 24,v11.1976, ex Toona and unknown host, H. Roberts (BMNH); 1d, same, Fly River,
Kiunga, viii.1969, J. Sedlacek (BPBM); 1 specimen, same, Lae, 11.x1.1978, N-311, L. Deharveng
(MHNG).
Ancistria bicolorata sp. n. (Figs 50-53)
Description. Length 7.5 mm. Black, tibial spurs and apical tarsal segments
reddish brown, apical 4 abdominal sternites brown, elytra reddish brown with narrow
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1011
black transverse bands at base and in the the middle; surface mostly mat. Head (fig. 51)
1.2 times as long as wide, coarsely punctate laterally, slightly finer on disk; midline
impressed: admedian lines moderately long, almost straight, admedian lobes narrowly
triangular, ending anteriorly almost level with lateral frontal processes which are
careniform and whose combined width is 0.4 times that of frontoclypeal depression.
Eye large, temple length/eye diameter ratio as 0.9. Antenna (fig. 50) with segment 1
bearing 2 to 3 indistinct dorsal grooves; segment 2 subglobular; segment 3 oval; seg-
ments 4 and 5 asymmetrically oval; segments 6-11 flattened, forming an indistinct club;
segment 11 as long as wide. Pronotum 1.6 times as long as wide, bordered at base,
Fics 43-49
Ancistria beccarii. 43, head, dorsal view; 44, parameres; 45, median lobe, apical piece; 46,
antenna; 47, ostium bursae; 48, sternite and tergite VIII; 49, spermatheca.
1012 DANIEL BURCKHARDT & ADAM SLIPINSKI
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Ancistria bicolorata. 50, antenna; 51, head, dorsal view; 52, spermatheca; 53, ostium bursae.
strongly widened in basal third, subparallel in apical two thirds; punctation coarser than
on head, sparser on disk. Elytra 3.5 times as long as wide, 2.3 times as long as prono-
tum; costa 3 strongly, costa 5 weakly inflated subapically; interval 4 shortest followed
by 6, 5 and 3 which is longest; apex with V-shaped emargination, without tooth at
suture. Genitalia as in figs 52, 53.
Material examined. Holotype 9, Vietnam: East Tonkin, region of Hoa Binh, 1918, R. P.
A. de Cooman (MNHN).
Ancistria bostrychoides Grouvelle (Figs 54-57)
Ancistria bostrychoides GROUVELLE, 1908: 454. Lectotype d, Sikkim: (MNHN); here designated
(examined).
Ancistria (Ancistria) bostrychoides Grouvelle; GROUVELLE, 191 3a: 444.
Description. Length 8.0 mm. Black, venter of abdomen dark brown; surface
mostly shiny. Head (fig. 55) 1.2 times as long as wide, strigosely punctate; median line
impressed, except for base; admedian lines short, strongly curved, admedian lobes
short, their largest width near apex which is obliquely truncate; lateral frontal processes
flattened, their joint width 0.5 times that of frontoclypeal depression. Eye small, temple
length/eye diameter ratio as 1.7. Antenna (fig. 54) with segment 1 bearing 1 distinct and
1-2 indistinct dorsal grooves; segment 2 subglobular; segment 3 clavate; segments 4-6
oval; segments 7-11 flattened, forming distinct club; segment 11 distinctly longer than
wide. Pronotum 1.5 times as long as wide, bordered at base, evenly widening almost to
apex; punctation somewhat coarser than on head, leaving indistinct longitudinal
puncture-free stripe in the middle. Elytra 2.8 times as long as their joint width, 2.2
times as long as pronotum; costa 3 strongly swollen and oblique subapically; interval 3
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1013
Fics 54-57
Ancistria bostrychoides. 54, antenna; 55, head, dorsal view; 56, parameres; 57, median lobe,
apical piece.
longest, 4 shortest and 5 intermediate; apex with V-shaped emargination, without
sutural tooth. Genitalia as in figs 56, 57.
Distribution. Recorded from Sikkim (GROUVELLE, 1908, 1913a; HETSCHKO, 1930).
Material examined. India: lectotype d , Sikkim (MNHN).
Ancistria bouchardi Grouvelle (Figs 58-61)
Ancistria (Parancistria) bouchardi GROUVELLE, 1913a: 474. Lectotype 9, Sumatra: Palembang,
Bouchard (MNHN); here designated (examined).
Description. Length 5.7-8.9 mm. Dark brown, apical half of elytra light or
chestnut brown except for apex which has a dark spot along the suture; surface mostly
mat. Head (fig. 59) 1.2 times as long as wide, finely, laterally strigosely punctate;
median line impressed, admedian lines moderately short, almost straight, admedian
lobes narrowly triangular; lateral frontal processes flat, their joint width 1.8-2.3 times
that of frontoclypeal depression. Eye moderately large, temple length/eye diameter ratio
as 1.2-1.6. Antenna (fig. 58) with segment | bearing 2 distinct wide dorsal grooves,
delimited by 3 subparallel ridges; segment 2 subglobular; segment 3 clavate, segments
4 and 5 irregularly oval; segments 6-11 strongly flattened, forming club; segment 11
longer than wide. Pronotum 1.6-1.7 times as long as wide, incompletely bordered at
base, widening in basal half, with subparallel margins in apical half; punctation much
coarser than on head, somewhat irregularly spaced, leaving narrow longitudinal median
puncture-free band in basal three quarters. Elytra 3.0-3.2 times as long as wide, 2.0-2.1
times as long as pronotum; costa 3 weakly inflated subapically, weakly oblique, costae
4 and 6 merging subapically; intervals 3 and 6 subequal, longer than subequal intervals
1014 DANIEL BURCKHARDT & ADAM SLIPINSKI
Fics 58-61
Ancistria bouchardi. 58, antenna; 59, head, dorsal view; 60, 61, spermatheca; 60, lectotype.
4 and 5; apex truncate, without or with indistinct tooth at suture. Genitalia as in figs 60,
61.
Distribution. Recorded from Sumatra (GROUVELLE, 1913a; HETSCHKO, 1930). Material
examined. Indonesia: lectotype 2, Sumatra (MNHN). Without locality data: 19 (ZMPA).
Comments. The type series includes, apart from the lectotype, 16 paralectotype
from the same locality which is similar to A. micros (cf. comments on A. micros).
Ancistria brancuccii sp. n. (Figs 2, 8, 62-66)
Description. Length 5.1-6.6 mm. Dark brown to black, legs chestnut brown;
surface shiny. Head (figs 2, 63) 1.4-1.6 times as long as wide, finely punctate; median
line impressed, weak in basal third, evanescent at base, admedian line moderately long,
strongly curved or bent, admedian lobes short, their largest width near apex, apex
obliquely truncate; lateral frontal processes flattened, their joint width 0.9-1.2 times that
of frontoclypeal depression. Eye small, temple length/eye diameter ratio as 1.7-1.9.
Antenna (fig. 62) with segment 1 bearing 2 shallow dorsal grooves; segment 2 sub-
globular; segment 3 clavate; segments 4-6 irregularly oval; segments 7-11 moderately
flattened forming a distinct club; segment 11 distinctly longer than wide. Pronotum 1.8-
2.0 times as long as wide, not bordered at base, widening in basal half, with subparallel
margins in apical half; punctation coarser than on head, in the middle leaving puncture-
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1015
free longitudinal band. Elytra 3.5-3.8 times as long as their joint width, 1.9-2.1 times as
long as pronotum; costa 3 weakly inflated subapically, costae 4 and 5 merging
subapically; interval 3 longer than 4 which is shorter than 5, interval 6 not impressed;
apex truncate with V-shaped incision at suture, with small sutural tooth (fig. 8).
Genitalia as in figs 64-66.
Material examined. Holotype d, East Nepal: Arun Valley, Mure - Num, 1550-2000 m, 4-
7.vi.1983, M. Brancucci (NHMB). Paratypes. Nepal: 184, same data as holotype (NHMB,
MHNG, ZMPA); 26, same but Chichila, 1950 m, 31.v.1983 (NHMB); 14, same but Num -
Hedanga, 800- 1500 m, 7.vi.1983; 16, Num, 1550 m, 5-6.vi.1983; 26, East Nepal, Koshi,
Lumbughat - Saiseghat, 450 m, 15.vi.1985, M. Brancucci (NMHB).
FIGS 62-66
Ancistria brancuccii. 62, antenna; 63, head, dorsal view; 64, parameres; 65, sternite and tergite
VIII; 66, median lobe, apical piece.
Ancistria concava sp. n. (Figs 67-72)
Description. Length 4.6-5.4 mm. Almost black, legs and abdominal venter dark
brown, elytra black at base, irregularly dark brown in apical two thirds; head mat,
pronotum and elytra shiny. Head (fig. 68) 1.2 times as long as wide; punctation fine,
1016 DANIEL BURCKHARDT & ADAM SLIPINSKI
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Fics 67-72
Ancistria concava. 67, antenna; 68, head, dorsal view; 69, parameres; 70, median lobe, apical
piece; 71, sternite and tergite VIII; 72, spermatheca.
slightly strigose; median line strongly impressed in apical half gradually getting weaker
towards base, absent from basal part; admedian lines moderately long, weakly curved,
admedian lobes narrowly triangular; lateral frontal processes flattened, their joint width
2.4-3.4 times that of frontoclypeal depression. Eye moderate in size, temple length/eye
diameter ratio as 0.8-1.3. Antenna (fig. 67) with segment 1 bearing 2-3 grooves, |
distinct, the other ones fine; segment 2 subglobular; segments 3-5 asymmetrically oval
to clavate; segments 6-11 strongly flattened, forming distinct club; segment 11
distinctly longer than wide. Pronotum 1.7-1.8 times as long as wide, not bordered
basally, widening in apical half, with subparallel margins in most of apical half;
punctures as on head, slightly coarser at base, sparser on disk leaving puncture-free area
in the middle. Eyltra 2.8-3.1 times as long as wide, 1.8 times as long as pronotum;
costae 2 and 3 subequal, 4 shorter; intervals 3 and 5 subequal, 4 shorter; apex weakly
emarginate with indistinct sutural tooth. Genitalia as in figs 69-72.
Material examined. Holotype 2, Sumatra: Babahrot, 100 m, 15.viii.1983, J. Klapperich
(MHNG). Paratype d, Philippines: Luzon, Laguna, Mount Makiling, 400 feet elevation,
26.vill. 1930, ex branches of Toona calentas, F. C. Hadden (BPBM).
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1017
Ancistria cornuta sp. n. (Figs 73-77)
Description. Length 5.7 mm. Dark brown to black; surface mostly mat. Head
(fig. 74) 1.0 times as long as wide, moderately punctate; median line impressed except
for base; admedian lines long, impressed, weakly curved; admedian lobes narrowly
triangular; lateral frontal processes careniform, their joint width 0.5 times that of
frontoclypeal depression which bears a small horn in the middle. Eye large, temple
width/eye diameter ratio as 0.6. Antenna (fig. 73) with segment | bearing 2 shallow
dorsal grooves; segment 2 subglobular; segment 3 clavate; segments 4-6 asymmetri-
cally widening towards apex, gradually enlarged from segment 4 to 6; segments 7-11
flattened, forming an indistinct club; segment 11 sightly longer than wide. Pronotum
1.3 times as long as wide, distinctly bordered at base, strongly widening in basal half,
: |
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VEN, No
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Fics 73-77
Ancistria cornuta. 73, antenna; 74, head, dorsal view; 75, parameres; 76, median lobe, apical
piece; 77, sternite and tergite VIII.
1018 DANIEL BURCKHARDT & ADAM SLIPINSKI
with subparallel margins in apical half; punctation coarser than on head. Elytra 3.0
times as joint width, 2.3 times as long as pronotum; costa 3 swollen and oblique
subapically; intervals getting gradually longer from 3 to 6; apex with V-shaped incision
at suture, without sutural tooth. Genitalia as in figs 75-77.
Material examined. Holotype d, Laos: Monhot, G. Lewis collection B.M. 1910-248
(BMNH).
Ancistria costata sp. n. (Figs 78-81)
Description. Length 6.9 mm. Head and thorax black, elytra very dark brown;
surface moderately shiny. Head (fig. 79) 1.1 times as long as wide, relatively convex
dorsally, with strigose punctation; median line impressed; admedian lines moderately
long, straight, admedian lobes narrowly triangular; lateral frontal processes flattened,
their joint width 1.1 times that of frontoclypeal depression. Eye large, temple length/eye
width ratio as 0.9. Antenna (fig. 78) with segment | bearing 2 distinct, unequal dorsal
grooves; segment 2 subglobular; segments 3-6 oval; segments 7-11 flattened, forming a
distinct club; segment 11 distinctly longer than wide. Pronotum 1.6 times as long as
wide, bordered at base, widening in basal quarter from there to apex with subparallel
margins; punctation slightly coarser than on head, denser at base, leaving longitudinal
median puncture-free stripe. Elytra 3.1 times as long as wide, 2.1 times as long as
79
81
Fics 78-81
Ancistria costata. 78, antenna; 79, head, dorsal view; 80, spermatheca; 81, ostium bursae.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1019
pronotum; costa 3 strongly raised forming oblique ridge which reaches subapically the
fused costae 7 and 8 which form a ridge, costa 5 merging with costa 3; intervals 3 and 4
merging subapically, shorter than merging intervals 5 and 6; with V-shaped incision at
suture, without sutural tooth. Genitalia as in figs 80, 81.
Material examined. Holotype ®, "British Bootang": 1899, M. Basti (MNHN).
Comments. We were unable to locate "British Bootang". In the MNHN are other
passandrids with the same locality labels, or with "British Bootan, Padong".
Ancistria emarginata Grouvelle (Figs 82-85)
Ancistria emarginata GROUVELLE, 1913b: 56. Holotype ?, Taiwan: Hoozan, 1908-1910, H.
Sauter (DEIC); (examined).
Description. Length 5.7-7.9 mm. Black, legs dark brown; surface mostly mat.
Head (fig. 83) 1.2 times as long as wide; finely, laterally strigosely punctured; median
line impressed almost to base; admedian lobes narrowly triangular; lateral frontal
processes flattened, their joint width 4.5-5.0 times that of frontoclypeal depression.
Dorsal surface of mandibles transversely rugose (fig. 83). Eye moderately large; temple
length/eye diameter ratio as 1.1-1.2. Antenna (fig. 82) with segment 1 bearing 2 distinct
grooves; segment 2 subglobular; segment 3 clavate; segments 4 and 5 asymmetrically
oval; segments 6-11 flattened, forming club; segment 11 as long as wide. Pronotum 1.6-
1.7 times as long as wide, base not distinctly bordered, strongly widened in basal half,
FIGs 82-85
Ancistria emarginata. 82, antenna; 83, head, dorsal view; 84, 85, spermatheca.
1020 DANIEL BURCKHARDT & ADAM SLIPINSKI
with subparallel margins in apical half; punctation coarser than on head, unevenly
spaced leaving a puncture-free longitudinal band in basal two thirds. Elytra 3.2-3.4
times as long as wide, 2.1-2.3 times as long as pronotum; costa 3 weakly inflated
subapically; interval 3 longer than 4 which is shorter than 5; apex distinctly emarginate,
sutural apex with tooth. Genitalia as in figs 84, 85.
Distribution. Recorded from Taiwan (GROUVELLE, 1913b; HETSCHCKO, 1930). Material
examined. Taiwan: holotype © (DEIC). China: 32, Sichuan, Mount Emei, 600-1050 m, 5-
19.v.1889, L. Bocak (ZMPA, MHNG).
Ancistria fabricii Reitter (Figs 86-88)
Ancistria fabricii REITTER, 1877: 134. Lectotype 2, Southeast Asia [India orientalis] (MNHN);
here designated (examined).
Ancistria (Ancistria) fabricii Reitter; GROUVELLE, 1913a: 494.
Description. Length 4.5 mm. Head and thorax dark chestnut brown, elytra and
legs somewhat lighter; surface mostly mat. Head (fig. 87) 1.2 times as long as wide,
with very fine, laterally slightly strigose punctation; median line evenly impressed from
base to apex; admedian lines moderately long, almost straight, admedian lobes
narrowly triangular; lateral frontal processes flattened, their joint width 3.0 times that of
frontoclypeal depression. Eye large, temple length/eye diameter ratio as 1.0. Antenna
(fig. 86) with segment | bearing each 1 distinct and 1 indistinct dorsal groove; segment
2 subglobular; segment 3 clavate; segments 4-5 oval; segments 6-11 flattened, forming
distinct club; segment 11 longer than wide. Pronotum 2.0 times as long as wide, not
Fics 86-88
Ancistria fabricii. 86, antenna; 87, head, dorsal view; 88, spermatheca.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1021
bordered at base, widening towards the middle, apical half with subparallel margins;
punctation much coarser than on head, covering whole surface, slightly denser
anteriorly. Elytra 3.2 times as long as wide, 2.0 times as long as pronotum; costa 3
slightly thickened and weakly oblique subapically, costae 4 and 5 merging apically:
interval 3 longer than 5, which is longer than 4; apex truncate with V-shaped incision at
suture, without sutural tooth. Genitalia as in fig. 88.
Distribution. Reported from Southeast Asia (REITTER, 1877; GROUVELLE, 1913a;
HETSCHKO, 1930). Material examined. Southeast Asia: lectotype 2 (MNHN).
Ancistria foraminifrons sp. n. (Figs 89-96)
Ancistria tarsalis sensu GROUVELLE, 1913a: 453, nec WATERHOUSE, 1876: 121.
Description. Length 6.9-8.9 mm. Dark brown to black; surface shiny. Head (figs
90, 96) 1.2-1.5 times as long as wide, covered in moderately fine strigose punctation;
median line indistinct in basal two thirds, grooved in apical third; admedian lines very
short and deep, admedian lobes short, triangular; lateral frontal processes careniform,
their joint width 0.7-1.2 times that of frontoclypeal depression. Eye small, temple
length/eye diameter ratio as 1.7-2.0. Antenna (figs 89, 95) with segment 1 bearing 2
shallow dorsal grooves; segment 2 oval; segments 3-6 oval to shortly clavate; segments
7-11 flattened, forming a distinct club; segment 11 distinctly longer than wide.
Pronotum 1.8-2.0 times as long as wide, entirely bordered at base, strongly narrowed in
basal third; punctation coarser than on head, sparser medially in basal two thirds. Elytra
4.2-4.6 times as long as wide, 2.3-2.5 times as long as pronotum; costa 3 inflated
subapically, costae 4-6 merging together at level of inflated portion of costa 3; interval
3 longer than intervals 4 to 6 which are gradually increasing in length apically; apex
with V-shaped incision at suture, lacking a sutural tooth. Genitalia as in figs 91-94.
Distribution. Recorded as A. tarsalis from Sumatra (GROUVELLE, 1913a). Material
examined. Holotype d, Papua New Guinea: EH Province, region of Kainantu Onerunka,
18.x11.1979, W. G. Ullrich (MHNG). Paratypes. Papua New Guinea: 14, same as holotype but
4.11.1980; NE New Guinea, S Garaina, 900 m, 15-21.1.1968, J. & M. Sedlacek (BPBM): 14,
West New Guinea, Star Mountains, Sibil Valley, 1245 m, 18.x.-8.x1.1961, S. Quate (BPBM).
Java: 16, Mount Preanger, 1400 m, 24.vi.1927, L. G. E. Kalshoven (RMNH). Sumatra: 46, Si-
Rambé, x11.1890-111.1891, E. Modigliani (MCSN, MNHN) (specimens identified as A. tarsalis by
Grouvelle).
Ancistria grouvellei nomen nov. (Figs 97-99)
Ancistria (Parancistria) filum GROUVELLE, 1913a: 484. Holotype 9, New Guinea: Dorey, Pascoe
collection 93-60 (BMNH); (examined). Junior primary homonym of Ancistria filum
REITTER, 1876: 39 (= Taphroscelidia filum (Reitter); BURCKHARDT & SLIPINSKI, 1991).
Description. Length 5.1 mm. Dark brown to almost black, antennae and legs
somewhat lighter, apical half of elytra reddish brown, abdominal venter brown; surface
mostly mat. Head (fig. 98) 1.2 times as long as wide; punctation strigose laterally;
median line distinct in apical half, very fine in basal half; admedian lines moderately
1022 DANIEL BURCKHARDT & ADAM SLIPINSKI
Fics 89-96
Ancistria foraminifrons. 89, 95, antenna; 90, 96, head, dorsal view; 91, parameres; 92, 93, median
lobe, apical piece; 94, sternite and tergite VIII.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1023
we
004, TR
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RS
Fics 97-99
Ancistria grouvellei. 97, antenna; 98, head, dorsal view; 99, spermatheca.
long, straight, admedian lobes narrowly triangular; lateral frontal processes flattened,
bearing a blunt, foreward directed tubercle antero-medially, their joint width 9.5 times
that of frontoclypeal depression. Eye large, temple length/eye diameter ratio as 0.9.
Antenna (fig. 97) with segment | bearing 2 fine dorsal grooves; segment 2 subglobular;
segments 3-6 oval; segments 7-11 flattened, forming a distinct club; segment 11 longer
than wide. Pronotum 1.9 times as long as wide, not bordered at base, widening in basal
half, apical half mostly with subparallel margins; sparsely punctate, sparser at base and
on disk. Elytra 3.4 times as long as wide, 1.9 times as long as pronotum; costa 3 not
markedly inflated subapically, costae 4-6 merging subapically; interval 3 long, intervals
4 and 5 much shorter; apex weakly emarginate, hardly incised at suture, with indistinct
sutural tooth. Genitalia as in fig. 99.
Distribution. Recorded from New Guinea (GROUVELLE, 1913a; HETSCHKO, 1930). Ma-
terial examined. New Guinea: holotype 2 (BMNH); 12, Koitaki, 1500 feet, xi-xii.1928, Pem-
berton (BPBM).
Ancistria indica sp. n. (Figs 100-105)
Description. Length 6.0-6.6 mm. Black; surface shiny. Head (fig. 101) 1.1 times
as long as wide, with coarse strigose punctation; median line impressed from base to
apex; admedian lines straight, long, admedian lobes narrowly triangular, lateral frontal
processes flattened, their combined width 1.7-2.0 times that of frontoclypeal
1024 DANIEL BURCKHARDT & ADAM SLIPINSKI
Fics 100-105
Ancistria indica. 100, antenna; 101, head, dorsal view; 102, parameres; 103, median lobe, apical
piece; 104, sternite and tergite VIII: 105, spermatheca.
depression. Mandibles transversely rugose dorsally. Eye large, temple length/eye
diameter ratio as 0.6-0.7. Antenna (fig. 100) with segment | bearing 1-2 indistinct
dorsal grooves; segment 2 subglobular; segment 3 clavate; segments 4-6 short, asym-
metrically widening towards apex; segments 7-11 flattened, forming an indistinct club;
segment 11 slightly longer than wide. Pronotum 1.6-1.7 times as long as wide, bordered
at base, strongly widening in basal half, weakly narrowing toward apex in apical half;
anteriorly finely, otherwise coarsely punctured as on head, a narrow longitudinal,
median band without punctures. Elytra 3.1 times as long as wide, 2.0-2.2 times as long
as pronotum: costa 3 distinctly inflated subapically, costae 4 merging with 5 and both
together with 6; intervals increasing in length apically from 3 to 6; apex with U-shaped
incision at suture, sutural tooth indistinct. Genitalia as in figs 102-105.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1025
Material examined. Holotype d, India: Dehra Dun Coll. B. M. 1932-26 (BMNH).
Paratypes. India: 24, 19, same data as holotype, 14 in addition with following information:
Tamil Nadu, Anamalai hills, 2400, Madras, 15.51.1930, J. C. M. Gardner (BMNH, MHNG).
Comments. Even though only one specimen bears a label, indicating that it was
collected in Southern India, it is likely that the whole type series has the same origin.
The series came probably via the Dehra Dun collection to the BMNH were the "Dehra
Dun coll." labels were added .
Ancistria lewisi Reitter (Figs 106-110)
Ancistria lewisi REITTER, 1889: 315. Lectotype, Japan: Yuyama in Higo, 1.vi.1881, G. Lewis,
labelled "Nagasaki, 22.v.-3.v1.1881, G. Lewis" (BMNH); here designated (examined).
Ancistria (Parancistria) lewisi Reitter; GROUVELLE, 1913a: 488.
Description. Length 4.8 mm. Dark chestnut brown, elytra in subapical quarter
lighter; surface mostly mat. Head (fig. 107) 1.5 times as long as wide, with fine strigose
punctation; median line basally weakly, apically distinctly impressed; admedian lines
moderately long, strongly curved, admedian lobes widest subapically and then suddenly
narrowing towards apex; lateral frontal processes careniform, their joint width 0.9 times
that of frontoclypeal depression. Eye small, temple length/eye diameter ratio as 1.9.
Antenna (fig. 106) with segment 1 bearing 2 weak dorsal grooves; segment 2
subglobular; segment 3 clavate; segments 4-6 irregularly oval; segments 7-11 flattened,
forming a distinct club; segment 11 distinctly longer than wide. Pronotum 2.0 times as
long as wide, not bordered at base, strongly widening in basal two thirds, with almost
subparallel margins in apical third; punctation coarser than on head, slightly sparser on
disk. Elytra 4.1 times as long as wide, 2.0 times as long as pronotum; costa 3 hardly
109
Fics 106-110
Ancistria lewisi. 106, antenna; 107, head, dorsal view; 108, sternite and tergite VIII; 109,
parameres; 110, median lobe, apical piece.
1026 DANIEL BURCKHARDT & ADAM SLIPINSKI
inflated subapically; interval 5 shorter than 3 but longer than interval 4, interval 6 indis-
tinct; apex weakly emarginate, small sutural tooth present. Genitalia as in figs 108-110.
Distribution. Recorded from Japan (REITTER, 1889; GROUVELLE, 1913a; HETSCHKO,
1930). Material examined. Japan: holotype, Nagasaki (BMNH); 14 paralectotype, Higo
(HNMH).
Comments. According to REITTER’s (1889) indications the type material was
collected at Yuyama in Higo. The lectotype in the BMNH is labelled “Nagasaki”; this
label has been added later and is certainly wrong. A paralectotype in the HNHM bears
the same data as mentioned in the original description (REITTER, 1889) (cf. comments to
A. reitteri).
Ancistria longicapitata sp. n. (Figs 111-115)
Description. Length 5.8 mm. Black, apical half of elytra chestnut brown; surface
mostly mat. Head (fig. 112) 1.7 times as long as wide, with strigose punctation
laterally; median line finely impressed in apical third, indistinct in basal two thirds;
ae
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115
Fics 111-115
Ancistria longicapitata. 111, antenna; 112, head, dorsal view; 113, parameres; 114, median
lobe, apical piece; 115, sternite and tergite VIII.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1027
admedian lines short, strongly angular, admedian lobes widest near apex, obliquely
truncate apically; lateral frontal processes flattened, their joint width 1.1 times that of
frontoclypeal impression. Eye small, temple length/eye diameter ratio as 1.9. Antenna
(fig. 111) with segment | bearing 2-3 shallow dorsal grooves; segment 2 subglobular;
segment 3 shortly clavate; segments 4-6 oval; segments 7-11 strongly flattened, for-
ming a well-defined club; segment 11 slightly longer than wide. Pronotum 2.3 times as
long as wide, not bordered at base, widened in basal half, with subparallel margins in
apical half. Elytra 3.7 times as long as their joint width, 1.9 times as long as pronotum;
costa 3 weakly thickened subapically, costae 4, 5 and 6 merging apically; intervals 4
and 5 subequal, shorter than 3 and 6; apex truncate, without sutural tooth. Genitalia as
in figs 113-115.
Material examined. Holotype d, Papua New Guinea: Morobe Province, Wau, Mount
Kaindi, 1150-2300 m, 20.1.-1.11.1993, flight intercept trap in Cordia, Y. Basset (MHNG).
Ancistria micros Grouvelle (Figs 116-122)
Ancistria (Parancistria) micros GROUVELLE, 1913a: 482. Holotype 2, Malaysia: Perak, Doherty
(MNHN); (examined).
Description. Length 3.4-3.9 mm. Brown to dark chestnut brown, elytra some-
what lighter; surface mostly mat. Head (fig. 117) 1.2 times as long as wide; finely to
coarsely punctate, sometimes strigose; median line impressed, fine; admedian lines
moderately long, more or less straight, admedian lobes narrowly triangular; lateral
frontal lobes flattened, their joint width 2.8-4.0 times that of frontoclypeal depression.
Eye more or less convex, temple length/eye diameter ratio as 1.0-1.1. Antenna (fig.
116) with segment | bearing | distinct dorsal groove; segment 2 subglobular; segment
3 clavate; segments 4 and 5 oval; segments 6-11 flattened, forming a club; segment 11
longer than wide. Pronotum 1.6-1.8 times as long as wide, not bordered at base,
widening in basal half, weakly narrowing towards the apex or with subparallel margins
in apical half; punctation coarser than on head, sparser on disc, leaving puncture-free
longitudinal band in the middle. Eyltra 3.1-3.2 times as long as wide, 2.0-2.1 times as
long as pronotum; costa 3 weakly inflated subapically, costae 4 to 6 merging apically;
intervals 3 and 6 longer than 4 and 5; apex truncate, without sutural tooth. Genitalia as
in figs 118-122.
Distribution. Recorded from Malaysia (GROUVELLE, 1913a; HETSCHKO, 1930). Material
examined. Malaysia: Holotype 9, Perak (MNHN). Borneo: 16, Sabah, Tenompok, 10-19.ii.
1959, T. C. Maa (BPBM); 1 specimen, same, near Keningau, 20.iii.1989, M. Ito (KYUN); 16,
Sarawak, Bau, Lake area, 29-30.viii.1958, T. C. Maa (BPBM).
Comments. A d specimen from Sumatra: Palembang (MNHN), paralectotype of
A. bouchardi Grouvelle) keys out with A. micros, but differs in following taxono-
mically relevant characters: the relatively wider and more impressed frontoclypeal
depression, the narrower lateral frontal processes and the genitalia. The specimen may
represent a new species but more material is necessary to confirm this.
1028 DANIEL BURCKHARDT & ADAM SLIPINSKI
À GO ER lou x
117 118
120
Fics 116-122
Ancistria micros. 116, antenna; 117, head, dorsal view; 118, parameres; 119, median lobe, apical
piece: 120, ostium bursae; 121, sternite and tergite VIII; 122, spermatheca.
Ancistria nepalensis sp. n. (Figs 123-126)
Description. Length 5.7-6.0 mm. Dark brown to black, legs brown; surface
mostly shiny, head mat. Head (fig. 124) 1.3-1.4 times as long as wide, with strigose
punctation; median line impressed apically getting weaker towards base; admedian
lines moderately long, gently curved, admedian lobes narrowly triangular; lateral
frontal processes flattened, their joint width 2.5-3.2 that of frontoclypeal depression.
Eye large, temple length/eye diameter ratio as 1.2-1.6. Antenna (fig. 123) with segment
1 bearing 1 distinct and 1 indistinct dorsal grooves; segment 2 subglobular; segment 3
clavate; segments 4-6 oval, segments 7-11 flattened, forming distinct club; segment 11
longer than wide. Pronotum 2.0-2.1 times as long as wide, not or indistinctly bordered
at base, widening towards middle, apical half with subparallel margins; punctation
somewhat coarser than on head, absent from a longitudinal patch in the middle towards
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1029
126
124
Fics 123-126
Ancistria nepalensis. 123, antenna; 124, head, dorsal view; 125, spermatheca; 126, ostium bursae.
the base. Elytra 3.6-4.1 times as long as wide, 2.0-2.1 times as long as pronotum; costa
3 slightly thickened subapically, costae 4 and 5 merging apically; interval 3 longer than
6, intervals increasing in length from 4 to 6; apex truncate, with V-shaped incision at
suture, without sutural tooth. Genitalia as in figs 125, 126.
Material examined. Holotype 2, Nepal: East Nepal, Arun Valley, Mure - Num, 1550-
2000 m, 4-7.vi.1983, M. Brancucci (NHMB). Paratypes 42, same data as holotype (NHMB,
MHNG).
Ancistria nicolettae sp. n. (Figs 9, 127-132)
Description. Length 6.9-7.4 mm. Dark chestnut brown to almost black, distal
third of elytra sometimes lighter; surface mostly shiny. Head (fig. 128) 1.3 times as
long as wide; moderately strigosely punctate; median line impressed except for apical
quarter; admedian lines moderately long, weakly curved, admedian lobes narrowly
triangular; lateral frontal processes careniform, their joint width 0.3-0.6 times that of
frontoclypeal depression. Eye large, temple length/eye diameter ratio as 0.6-1.0.
Antenna (fig. 127) with segment 1 bearing 2 indistinct dorsal grooves; segment 2
subglobular; segment 3 shortly clavate; segments 4 and 5 asymmetrically oval; seg-
ments 6 to 11 weakly flattened, forming an indistinct club; segment 11 slightly longer
than wide. Pronotum 1.9-2.0 times as long as wide, distinctly bordered at base, strongly
widening from base almost to apex; relatively evenly punctured, coarser than on head.
Elytra 3.8-4.1 times as long as wide, 2.2 times as long as pronotum; costa 3 weakly
inflated subapically, slightly oblique; intervals getting gradually shorter from 3 to 5; in
males apex emarginate with small sutural tooth, in females apex truncate or weakly
convex without sutural tooth (fig. 9). Genitalia as in figs 129-132.
1030 DANIEL BURCKHARDT & ADAM SLIPINSKI
128
130
Vy
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Zus Er I \
131
Fics 127-132
Ancistria nicolettae. 127 antenna; 128, head, dorsal view; 129, parameres; 130, median lobe,
apical piece; 131, sternite and tergite VIII; 132, spermatheca.
Material examined. Holotype ©, Philippines: N Palawan, Binaluan, xi-xii.1913, G.
Boettcher (ZMHB). Paratypes. Philippines: 16, 19, same data as holotype (ZMHB, MHNG).
Malaysia: 19, Selangor, Sg, Baloh F. R., 22.vi.1936, ex gallery of Xyleborus in Dipterocarpus
baudii, F. G. Browne (BMNH). Borneo: 16, Sabah, Kalabakan, 10-19.xi.1958, T. C. Maa
(BPBM). Sumatra: 1d, North Brastagi, G. Sibayak, 1450-1900 m, 19-23.11.1991, Bocäk &
Bocäkova (NHMB).
Ancistria papuana sp. n. (Figs 133-137)
Description. Length 6.0 mm. Black, legs, apical 2-3 abdominal sternites and
apical half of eyltra brown; surface shiny. Head (fig. 134) 1.3 times as long as wide,
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1031
135
Fics 133-137
Ancistria papuana. 133, antenna; 134, head, dorsal view; 135, parameres; 136, median lobe,
apical piece; 137, sternite and tergite VIII.
punctate; median line deeply impressed in apical half, indistinct in basal half; admedian
lines relatively short, impressed and strongly curved, admedian lobes broadest near
apex, obliquely trunctae apically; lateral frontal processes flattened dorsally, their
combined width 1.1 times that of frontoclypeal depression. Eye small, temple
length/eye diameter ratio as 1.6. Antenna (fig. 133) with segment | bearing 2 distinct
dorsal grooves; segment 2 subglobular; segments 3-6 increasing in size, oval; segments
7-11 strongly flattened, forming a dinstinct club; segment 11 longer than wide.
Pronotum 1.7 times as long as wide, bordered at base, widening to the middle, margins
of apical half subparallel; punctation somewhat coarser than on head, partly absent
from longitudinal stripe in the middle. Elytra 3.5 times as long as wide, 2.2 times as
1032 DANIEL BURCKHARDT & ADAM SLIPINSKI
long as pronotum; costa 3 inflated; intervals getting shorter from 3 to 6; apex with V-
shaped incision at suture, with small sutural tooth. Genitalia as in figs 135-137.
Material examined. Holotype d, Papua New Guinea: Wau, 15.ix., J. & M. Sedlacek
(BPBM).
Ancistria pilosa sp. n. (Figs 138-141)
Description. Length 4.9 mm. Dark brown, legs slightly lighter; surface shiny;
head covered in very short setae, pronotum and elytra in light, 0.1 mm long erect setae.
Head (fig. 139) 1.0 times as long as wide, with fine sparse, strigose punctation; median
line fine, slightly more impressed apically; admedian lines moderately long, weakly
curved, admedian lobes indistinctly delimited anteriorly, blunt; lateral frontal processes
flattened, their joint width 1.0 times that of frontoclypeal depression. Eye large, temple
length/eye diameter ratio as 0.7. Antenna (fig. 138) with segment 1 bearing 2 dorsal
grooves; segment 2 subglobular; segments 3-6 oval; segments 7-11 weakly flattened,
forming club; segment 11 slightly longer than wide. Pronotum 1.6 times as long as
wide, bordered at base, weakly widening towards apex; punctation sparser and coarser
than on head, with punctation-free longitudinal band in the middle. Elytra 3.4 times as
long as wide, 2.3 times as long as pronotum; costa 3 strongly raised and oblique
subapically; intervals 3 and 4 merging apically, 5 slightly longer but shorter than
interval 6; apex with large V-shaped incision at suture, without sutural tooth. Genitalia
as in figs 140, 141.
Distribution. Holotype 2, Taiwan: Fenchihu, 1400 m, iv-vi.1977, J. & S. Klapperich
(MHNG).
141
138 139
Fics 138-141
Ancistria pilosa. 138, antenna; 139, head, dorsal view; 140, spermatheca; 141, ostium bursae.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1033
Ancistria reitteri Lewis (Figs 142-144)
Ancistria reitteri LEWIS, 1893: 83. Lectotype ®, Japan: Yuyama in Higo, labelled "Nagasaki,
22.v.-3.v1.1881", G. Lewis (BMNH); here designated (examined).
Description. Length 4.5 mm. Dark brown, tarsi brown; surface shiny. Head (fig.
143) 1.1 times as long as wide; finely, laterally strigosely punctured; median line
impressed almost to base; admedian lobes narrowly triangular, blunt anteriorly; lateral
frontal processes flattened, their joint width 0.8 times that of frontoclypeal depression.
Dorsal surface of mandibles punctate. Eye moderately large; temple length/eye
diameter ratio as 1.0. Antenna (fig. 142) with segment 1 bearing 2 distinct grooves;
segment 2 subglobular; segment 3 shortly clavate; segments 4 and 5 asymmetrically
oval; segments 6-11 flattened, forming indistinct club; segment 11 as long as wide.
Pronotum 1.6 times as long as wide, not bordered at base, strongly widened in basal
two thirds, with subparallel margins in apical third; punctation coarser than on head,
density uneven leaving a puncture-free median longitudinal band in the middle. Elytra
2.6 times as long as wide, 1.8 times as long as pronotum; costa 3 hardly inflated
subapically; interval 3 longer than 4 which is shorter than 5; straight apically without
sutural tooth. Genitalia as in fig. 144.
NS
TE
11
QU
x
STONE
Figs 142-144
Ancistria reitteri. 142, antenna; 143, head, dorsal view; 144, spermatheca.
Distribution. Recorded from Japan (Lewis, 1893). Material examined. Japan: lectotype 9
of A. reitteri (BMNH).
Comments. According to LEWIS' (1893) description of A. reitteri the type
material was collected at Yuyama in Higo. The lectotype in the BMNH is labelled
"Nagasaki". The specimen at hand is the only representative known of the species and
is without any doubt the same specimen examined by Lewis. The label with "Nagasaki"
has been added subsequently and is wrong.
1034 DANIEL BURCKHARDT & ADAM SLIPINSKI
Ancistria retusa (Fabricius) (Figs 3, 4, 11, 12, 145-164)
Colydium retusa Fabricius, 1801: 555. Lectotype d, Sumatra, Daldorff (ZMUC); here designated
(examined).
Bostrichoides angulatus MONTROUZIER, 1855: 17. Lectotype, Woodlark Island, X. Montrousier
(ISNB); here designated (examined). Syn. n.
Ancistria retusa (Fabricius); GEMMINGER & VON HAROLD, 1868; GROUVELLE, 1882.
Prionophora cylindrica WESTWOOD, 1848: 85. Lectotype, India (BMNH); here designated
(examined); synonymised with A. retusa by GEMMINGER & VON HAROLD, 1868: 870.
Ancistria (Ancistria) albertisi GROUVELLE, 1913a: 446: Lectotype 2, New Guinea, Ramoi,
vi.1892, L. M. D'Albertis (MCSN); here designated (examined). Syn. n.
Ancistria (Ancistria) atra GROUVELLE, 1913a: 455. Lectotype 2, New Guinea, Allem? (MNHN);
here designated (examined). Syn. n.
Ancistria (Ancistria) brevis GROUVELLE, 1913a: 469. Lectotype, Southeast Borneo (BMNH); here
designated (examined). Syn. n.
Ancistria (Ancistria) excavata GROUVELLE, 1913a: 464. Lectotype ©, East Java: Sukabumi,
2000’, 1893, H. Fruhstorfer (MNHN); here designated (examined). Syn. n.
Ancistria (Ancistria) filiformis GROUVELLE, 1913a: 458. Holotype d, Australia: Cook Town
(MNHN); (examined). Syn. n.
Ancistria (Ancistria) gestroi GROUVELLE, 1913a: 449. Lectotype d, Sulawesi: Kandari, vii. 1874,
O. Beccari (MCSN); here designated (examined). Syn. n.
Ancistria (Ancistria) leai GROUVELLE, 1913a: 462. Lectotype ©, Australia: Morton Bay, H.
Deyrolle (MNHN); here designated (examined). Syn. n.
Ancistria (Ancistria) magna GROUVELLE, 1913a: 470. Lectotype ®, Singapore, Wallace
(BMNH); here designated (examined). Syn. n.
Ancistria (Ancistria) retusa (Fabricius); GROUVELLE, 1913a: 467.
Ancistria (Ancistria) retusa var. propingua GROUVELLE, 1913a: 469. Lectotype d, Philippines:
Palawan (MNHN); here designated (examined). Syn. n.
Ancistria sauteri GROUVELLE, 1913b: 54. Lectotype ©, Taiwan: Taihorin, Fuhosho, H. Sauter
(DEIC), here designated (examined). Syn. n.
Ancistria (Ancistria) africana GROUVELLE, 1919; 56. Holotype, Rhodesia: Sebakove, D. Dods
(SAMC); (examined). Syn. n.
Description. Length 4.8-8.6 mm. Dark brown to black with elytra sometimes
partly lighter, ranging from light brown to dark chestnut brown; surface mostly mat on
head and shiny on pronotum and elytra. Head (figs 3, 146, 148, 150) 1.2-1.4 times as
long as wide; finely, laterally strigosely punctate; median line impressed except for
base; admedian lines moderately short, weakly curved, admedian lobes narrowly
triangular; lateral frontal processes careniform, their joint width 0.5-0.7 times that of
frontoclypeal depression. Eye large, temple length/eye diameter ratio as 1.2-1.3.
Antenna (figs 145, 147, 149) with segment | bearing 2 dorsal grooves; segment 2
subglobular; segment 3 clavate; segments 4 and 5 asymmetrically oval; segments 6 to
11 flattened forming club; segment 11 slightly longer than wide. Pronotum 1.7-1.8
times as long as wide, bordered at base (fig. 4); widening in basal half, apical half with
subparallel margins or narrowing weakly to apex; punctation coarser than on head
leaving a puncture-free median stripe. Elytra 3.1-3.5 times as long as wide, 2.0-2.4
times as long as pronotum; carina 3 strongly inflated and oblique, in 2 more than in d;
interval 4 shorter than 3 and 5 which merge apically; apex truncate or weakly emar-
ginate with V-shaped incision at suture, without sutural tooth (figs 11, 12). Genitalia as
in figs 151-164.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1035
DA
4
i
N
148
ee 60 0p tly
VICHONMU EVI EBENE,
«C0
152
153 154
Fics 145-154
Ancistria retusa. 145, 147, 149, antenna; 146, 148, 150, head, dorsal view; 151, 152, sper-
matheca; 153, 154, ostium bursae.
Distribution. Reported from Sumatra, India, New Guinea, Borneo, Java, Australia,
Sulawesi, Singapore, Palawan, Taiwan and Zimbabwe (HETSCHKO, 1930, and literature cited
therein) as A. retusa, A. cylindrica, A. albertisi, A. atra, A. brevis, A. excavata, A. filiformis, A.
gestroi, A. leai, A. magna, A. retusa var. propinqua and A. africana, from Woodlark Island
(MONTROUZIER, 1855) as Bostrichoides angulatus, and as A. retusa from Australia (Queensland,
New South Wales) (HAWKESWOOD, 1987). Material examined: 236 specimens from Cameroon
(Biyan), Zaire, Zimbabwe, South Africa, India (Assam, Chambaganor, Madura), Sri Lanka,
1036 DANIEL BURCKHARDT & ADAM SLIPINSKI
x
---
7
/
=
Fics 155-164
Ancistria retusa. 155-157, 160-162, median lobe, apical piece; 158, 159, parameres; 163, 164,
sternite and tergite VIII.
China (Hainan), Taiwan, Thailand, Laos, Vietnam (Tonkin), Philippines (Mindoro, Negros,
Mindanao, Luzon, Calapan, Palawan), Borneo (Sabah, Brunei, Sarawak), Singapore, Sumatra,
Java, Sulawesi, Kapulauan Tukanbesi, Moluccas, Aru Island, New Guinea, Australia (Queens-
land, Victoria, possibly Tasmania) (ANIC, BMNH, BPBM, DEIC, ISNB, MCSN, MHNG,
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1037
MNHN, MRAC, NHMB, QMBA, SMTD, USNM, ZMHB, ZMPA, ZMUC). A single specimens
bears following data: Brazil (Coll. L. W. Schaufuss, ZMHB).
Comment. Some specimens bear the indication "Somerset" which may refer to a
locality in Queensland or in Tasmania. The occurence of A. retusa in Tasmania should
be confirmed with additional material. The single specimen from Brazil may have been
mislabelled or may be an introduction.
Ancistria stricta Grouvelle (Figs 165-169)
Ancistria (Parancistria) stricta GROUVELLE, 1913a: 477. Lectotype, Singapore (BMNH); here
designated (examined).
Description. Length 5.7 mm. Brown, apical half of elytra light brown; surface
mostly mat. Head (fig. 166) 1.3 times as long as wide; punctation fine, laterally slightly
strigose; median line distinctly impressed in apical half, finer in basal half fading
Fics 165-169
Ancistria stricta. 165, antenna; 166, head, dorsal view; 167, parameres; 168, sternite and
tergite VIII; 169, median lobe, apical piece.
1038 DANIEL BURCKHARDT & ADAM SLIPINSKI
towards base; admedian lines weakly curved to almost straight, admedian lobes
narrowly triangular; lateral frontal processes flattened, their joint width 1.6 times that of
frontoclypeal depression. Eye small, temple length/eye diameter ratio as 1.7. Antenna
(fig. 165) with segment | bearing 1 distinct dorsal groove; segment 2 subglobular;
segments 3-5 oval; segments 6-11 flattened, forming well-defined club; segment 11
slightly longer than wide. Pronotum 1.5 times as long as wide, bordered at base,
widening from base to apical third, from there weakly narrowing to apex; punctation
slightly coarser than on head, relatively evenly spaced. Elytra 2.6 times as long as wide,
1.9 times as long as pronotum; costa 3 hardly inflated subapically, costae 4 and 6
merging subapically; intervals 3 and 6 subequal, longer than subequal intervals 4 and 5;
apex truncate, no distinct sutural tooth developed. Genitalia as in figs 167-169.
Distribution. Singapore (GROUVELLE, 1913a; HETSCHKO, 1930). Material examined.
Singapore: Lectotype and paralectotype 2 (BMNH, MNHN). Borneo: 1d, West Sarawak, Quop,
Ancistria strigosa Grouvelle (Figs 170-172)
Ancistria (Ancistria) strigosa GROUVELLE, 1913a: 451. Lectotype ©, New Guinea: Amberbaki
(MCSN); here designated (examined).
Description. Length 9.4-14.6 mm. Chestnut brown to almost entirely black;
surface mostly shiny. Head (fig. 171) 1.2 times as long as wide, relatively strongly
convex; punctation coarse, strigose laterally; median line in basal quarter reduced,
impressed otherwise; admedian lines moderately long, weakly curved, admedian lobes
narrowly triangular; lateral frontal processes careniform, their joint width 0.3-0.8 times
that of frontoclypeal depression. Eye large, temple length/eye diameter ratio as 0.8-1.3.
172
Fics 170-172
Ancistria strigosa. 170, antenna; 171, head, dorsal view; 172, spermatheca.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1039
Antenna (fig. 170) with segment 1 bearing 2 dorsal grooves; segment 2 subglobular;
segments 3 oval; segments 4-11 gradually enlarged from 4 to 11, flattened; segment 11
about as long as wide. Pronotum 1.7-1.8 times as long as wide, distinctly bordered at
base, strongly widening from base to the middle, from there weakly narrowing to apex;
punctures much coarser than on head, absent from a longitudinal median stripe. Elytra
3.5-4.1 times as long as wide, 2.3-2.5 times as long as pronotum; costa 3 moderately
thickened subapically, fused costae 7 and 8 in 6 weakly, in 9 strongly inflated
apically; intervals getting gradually shorter from 3 to 6; apical margin incised at suture,
3 with small, 2 with large sutural tooth. Genitalia as in fig. 172.
Distribution. Recorded from New Guinea (Grouvelle, 1913a; Hetschko, 1930). Material
examined. New Guinea: Lectotype 2, paralectotypes 14, 19 (MCSN, MNHN); 19, Papua New
Guinea, Klunga, Fly River, 17.viii.1957, W. W. Brandt (BPBM). Solomon Islands: 12, Santa
Ysabel, Hageulu, 600-650 m, 11.ix.1964, R. Straatman (BPBM). Indonesia: 1 2, Ceram, Jilo,
1884, C. Ribbe (MNHN); 12, Vanikoro, François (MNHN); 22, Amboina, Doll. (MNHN).
Southeast Asia: 12 (MNHN).
Ancistria tarsalis Waterhouse (Figs 10, 173-177)
Ancistria tarsalis WATERHOUSE, 1876: 121. Lectotype, Java: Bowring (BMNH); here designated
(examined).
Ancistria (Obancistria) longior GROUVELLE, 1913a: 442. Lectotype 2, Sumatra, Si-Rambe,
xii. 1890-111.1891, E. Modigliani (MCSN); here designated (examined). Syn. n.
Fics 173-177
Ancistria tarsalis. 173, 177, spermatheca; 174, antenna; 175, head, dorsal view; 176, ostium
bursae.
1040 DANIEL BURCKHARDT & ADAM SLIPINSKI
Description. Length 8.1 mm. Dark brown; surface shiny. Head (fig. 175) 1.4
times as long as wide, strigosely punctate; admedian line fine in basal fifth, strongly
impressed in apical four fifth; admedian lines moderately long, relatively straight,
admedian lobes narrowly triangular; lateral frontal lobes flattened, their joint width 1.9
times that of frontoclypeal depression. Eye moderate in size, temple length/eye
diameter ratio as 1.3. Antenna (fig. 174) with segment | bearing 2 distinct dorsal
grooves; segment 2 subglobular; segment 3 clavate; segments 4-6 asymmetrically oval;
segments 7-11 strongly flattened, forming a well-defined club; segment 11 slightly
longer than wide. Pronotum 2.1 times as long as wide, distinctly bordered at base,
widened in basal third, with subparallel margins in apical two thirds; punctation slightly
coarser on head, iess dense on disk. Elytra 3.1 times as long as wide, 2.1 times as long
as pronotum; costa 3 strongly inflated and oblique subapically, fused costae 7 and 8
strongly thickened apically; intervals 3 and 6 long, merging apically, interval 4 shorter,
interval 5 intermediate; apex with V-shaped incision at apex, without sutural tooth (fig.
10). Genitalia as in figs 173, 176, 177.
Distribution. Recorded as A. tarsalis from Java (WATERHOUSE, 1876), and as A. longior
from Sumatra (GROUVELLE, 1913a; HETSCHKO, 1930). Material examined. Java: lectotype of A.
tarsalis (BMNH). Sumatra, lectotype and paralectotype of A. longior, Si-Rambe, xii.1890-
i11.1891, E. Modigliani (MCSN, MNHN). Borneo: 1%, Sabah, Mount Kinabalu National Park,
head quarters, Liwagu river, 1500 m, 21.v.1987, A. Smetana (MHNG).
Comment. Ancistria (Ancistria) tarsalis sensu GROUVELLE, 1913a, nec Water-
house, is the species described here as A. foraminifrons sp. n.
Ancistria tenera Giinther (Figs 178-182)
Ancistria (Obancistria) tenera Günther, in HELLER & GÜNTHER, 1936: 65. Holotype d, Indo-
nesia: West Java, Kamodjan, Southern slope of Mount Guntur, 1350 m, Garoet, 111.1934,
H. Overbeck (SMTD); (examined).
Description. Length 6.0 mm. Almost black, legs brown, elytra irregularly dark
chestnut brown; surface mostly shiny. Head (fig. 179) 1.3 times as long as wide; finely
punctured; median line impressed in apical half, indistinct basally; admedian lines rela-
tively short, weakly curved, admedian lobes narrowly triangular; lateral frontal pro-
cesses narrow, flattened, their joint width 0.7 times that of frontoclypeal depression.
Eye relatively small; temple length/eye diameter ratio as 1.7. Antenna (fig. 178) with
segment | weakly flattened dorsally, bearing 1 distinct and 1 indistinct grooves; seg-
ment 2 subglobular; segments 3-6 oval to clavate; segments 7-11 strongly flattened,
forming distinct club; segment 11 distinctly longer than wide. Pronotum 1.7 times as
long as wide, indistinctly bordered at base, strongly widened in basal two thirds and
weakly narrowing in apical third; punctation slightly coarser than on head, leaving a
puncture-free longitudinal band in basal two thirds. Elytra 3.1 times as long as wide, 2.1
times as long as pronotum; costa 3 very weakly inflated subapically; interval 3 longer
than 4 which is shorter than 5; weakly emarginate apically with small sutural tooth.
Genitalia as in figs 180-182.
Distribution. Recorded from Java (HELLER & GUNTHER, 1936). Material examined. Java:
holotype ¢, Kamodjan, Southern slope of Mount Guntur, 1350 m, Garoet, 11.1934, H. Overbeck
(SMTD).
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1041
180
Fics 178-182
Ancistria tenera. 178, antenna; 179, head, dorsal view; 180, parameres; 181, median lobe, apical
piece; 182, sternite and tergite VIII.
Ancistria tenuis Grouvelle (Figs 183-185)
Ancistria (Parancistria) tenuis GROUVELLE, 1913a: 486. Holotype 9, Southeast New Guinea:
Mount Astrolabe, 11.1893, Loria (MCSN); (examined).
Description. Length 3.7 mm. Head and thorax chestnut brown, antennae legs,
elytra and apex of abdominal venter lighter brown; surface on head mat, on pronotum
and elytra mostly shiny. Head (fig. 184) 1.3 times as long as wide, with fine, laterally
strigose punctation; median line impressed apically getting weaker towards base;
admedian lines moderately long, gently curved, admedian lobes narrowly triangular;
lateral frontal processes flattened, their joint width 2.5 times that of frontoclypeal
depression. Eye large, temple length/eye diameter ratio as 1.8. Antenna (fig. 183) with
segment | bearing 1 distinct and 1 indistinct dorsal grooves; segment 2 subglobular;
segment 3 clavate; segments 4-6 oval; segments 7-11 flattened, forming distinct club;
segment 11 longer than wide. Pronotum 2.0 times as long as wide, not bordered at base,
widening towards the middle, from there weakly narrowing to the apex; punctation
1042 DANIEL BURCKHARDT & ADAM SLIPINSKI
185
Fics 183-185
Ancistria tenuis. 183, antenna; 184, head, dorsal view; 185, spermatheca.
somewhat coarser than on head, irregular. Elytra 4.2 times as long as wide, 2.1 times as
long as pronotum; costa 3 slightly thickened and weakly oblique subapically, costae 4
and 5 merging apically; interval 3 longer than intervals 4 and 5; apex truncate, without
sutural tooth. Genitalia as in fig. 185.
Distribution. Reported from New Guinea (GROUVELLE, 1913a; HETSCHKO, 1930).
Material examined. New Guinea: holotype 2 (MCSN).
BIOGEOGRAPHY
Most of the 32 currently recognised species of Ancistria occur in the Old World
tropics. One species, A. retusa, has a very large range and has been recorded also from
Africa and Australia in addition to many localities in Asia. One specimen of A. retusa
may even originate from Brazil, a record which needs confirmation. The other 31
species, on the contrary, are restricted in their distribution and may be of use in
examining historical relationships among areas of endemism. For this, however, formal
hypotheses of the phylogenetic relationships of Ancistria are required.
The number of species with restricted distribution is relatively high (18 spp.),
suggesting that the available faunistic information is very incomplete. Particularly, with
the exception of A. retusa, the absence of species shared between the North Indian/
Himalayan region and continental Southeast Asia (excluding the Malayan Peninsula) on
one hand and the isolated position of the Philippines (excluding Palawan) on the other
hand indicate this lack of material. The most species rich area is New Guinea (9 spp.),
followed by Sumatra (7 spp.) and Borneo (6 spp.). The endemism is seemingly high on
New Guinea but virtually absent from Sumatra, Borneo and the Malayan Peninsula.
REVIEW OF PASSANDRIDAE IV: ANCISTRIA 1043
ACKNOWLEDGEMENTS
We thank all the persons who kindly provided material for our study or were
helpful in various ways during our visits to their institutions (cf. material and methods).
For useful comments on the manuscript draft we are very grateful to J. Pakaluk
(USNM) and I. Löbl (MHNG). The drawings of the heads and antennae were executed
by N. Lavoyer, Geneva, the other figures were inked by G. Roth, Geneva, and the SEM
pictures prepared by J. Wiiest, Geneva. To all we extend our sincerest thanks.
NOTE ADDED IN PROOF
SASAJ (1993) described Ancistria kurosawai and gave additional records of A.
apicalis from Japan. This publication came to our attention only when the present paper
was already submitted for publication. It was, therefore, too late to include this
information in the present revision. According to the description of SASAM (1993)
Ancistria kurosawai keys out with A. costata from which it can be differentiated as
follows:
27bis Body length 6.9 mm. Head 1.1 times as long as wide. “British Bootang”
ERDE IRR PER ED ERRO I A costata
- Body length 7.1-8.0 mm. Head 1.4 times as long as wide. Japan. . . . kurosawai
REFERENCES
BURCKHARDT, D. & S. A. SLIPINSKI. 1991. A review of the Passandridae (Coleoptera, Cucujoidea)
of the world. III. Genera Anisocerus, Aulonosoma, Passandrella, Passandrina, Scali-
diopsis and Taphroscelidia. Revue suisse Zool. 98: 453-497.
BURCKHARDT, D. & S. A. SLIPINSKI. 1995. Catogenus klapperichorum sp. n. and nomenclatoral
remarks on Catogenus carinatus Newman (Coleoptera, Passandridae). Ent. Probl. 26:
79-84.
ERICHSON, W. F. 1845. Naturgeschichte der Insecten Deutschlands. Erste Abteilung. Coleoptera,
vol. 3, Berlin, Verlag der Nicolaischen Buchhandlung.
Fabricius, I. C. 1801. Systema Eleutheratorum, 2. Kiel, 687 pp.
GEMMINGER, M. & E. VON HAROLD. 1868. Catalogus Coleopterorum, 3: 753-978.
GROUVELLE, A., 1883. Cucujides nouveaux du Musée civique de Génes. Annali Mus. civ. Stor.
nat. Genova 18: 275-296.
GROUVELLE, A., 1908. Coléopteres de la région indienne. Rhysodidae, Trogositidae, Nitidulidae,
Colydiidae, Cucujidae. Annls Soc. ent. Fr. 77: 315-495.
GROUVELLE, A., 1913a. Etude sur les Ancistria (Col. Passandridae). Annls Soc. ent. Fr. 81: 433-
494.
GROUVELLE, A., 1913b. H. Sauter's Formosa-Ausbeute. Rhysodidae, Nitidulidae, Ostomidae,
Colydiidae, Passandridae, Cucujidae, Cryptophagidae, Diphyllidae, Lathridiidae, Myce-
tophagidae, Dermestidae. Arch. Naturgesch. 79(11): 33-76.
GROUVELLE, A. 1919. Descriptions d'espèces nouvelles de Clavicornes de l'Afrique australe.
Mém. ent. 2: 47-60.
1044 DANIEL BURCKHARDT & ADAM SLIPINSKI
HAWKESWOOD, T. 1987. Beetles of Australia. Angus & Robertson Publishers, North Ryde, 248
PP.
HELLER, K. M. & K. GUNTHER. 1936. Ueber einige von H. Overbeck 1933/34 auf Java
gesammelte Insekten. Tijdschr. Ent. 79: 64-76.
HETSCHKO, A. 1930. Cucujidae, Thorictidae (Suppl.), Cossyphodidae (Suppl.), in Junk, W. & S.
Schenkling. Coleopterorum Catalogus, 109. Berlin, Junk, 124 pp.
KESSEL, F. 1921. Neue Monotomiden, Cucujiden u. Passandriden aus der Sammlung des
Deutschen Entomologischen Museums in Berlin. Arch. Naturgesch. 87 (6): 24-33.
LEWIS, G. 1893. Japanese Coleoptera - two new species. Entomologist’s mon. Mag. 29: 83.
MONTROUZIER, X. 1855. Essai sur la faune de l'île de Woodlark ou Moiou. Annis Soc. Agric. Lyon
(2) 7 (1): 1-114.
REITTER, E. 1876. Neue Gattungen und Arten aus der Familie der Cucujidae. Coleopterologische
Hefte 15: 37-64.
REITTER, E. 1877. Beiträge zur Kenntisse aussereuropäischer Coleopteren. Mitt. miinch. ent. Ver.
1: 126-140.
REITTER, E. 1889. Verzeichnis der Cucujiden Japans mit Beschreibung neuer Arten. Wien. ent.
Ztg. 8: 313-320.
SASAJI, H. 1993. Contribution to the taxonomy of the superfamily Cucujoidea (Coleoptera) of
Japan and adjacent districts, VII. Mem. Fac. Educ. Fukui University, series II (Nat. Sci.)
44(2): 17-25.
SLIPINSKI, S. A. 1987. A review of the Passandridae of the world (Coleoptera, Cucujoidea). I.
Genus Passandra Dalman. Annali Mus. civ. Stor. nat. Genova 86: 553-603.
SLIPINSKI, S. A. 1989. A review of the Passandridae (Coleoptera, Cucujoidea) of the world. II.
Genus Catogenus Westwood. Polskie Pismo ent. 59: 85-129.
WATERHOUSE, C. O. 1876. Descriptions of new Cucujidae and Cleridae. Entomologist's mon.
Mag. 13: 118-126.
WESTWOOD, J. O. 1848. The Cabinet of Oriental Entomology, 86 pp.
REVUE SUISSE DE ZOOLOGIE, 102 (4) : 1045-1064; décembre 1995
Review of the Pseudoscorpion Fauna of China
(Arachnida: Pseudoscorpionida)
Wolfgang SCHAWALLER
Staatliches Museum fiir Naturkunde
Rosenstein 1
D-70191 Stuttgart, Germany.
Review of the Pseudoscorpion Fauna of China (Arachnida: Pseudo-
scorpionida). - The known pseudoscorpions from China (including Tibet,
excluding Mongolia and Taiwan) and new material are compiled. 47 species
are listed, remarks on taxonomy and distribution are added. New species:
Centrochthonius sichuanensis n.sp., Stenohya chinacavernicola n.sp. New
records for China: Lagynochthonius tonkinensis, Tyrannochthonius japo-
nicus, Tyrannochthonius pachythorax, Tyrannochthonius robustus, Ditha
proxima, Geogarypus javanus, Bisetocreagris annamensis, Bisetocreagris
indochinensis, Bisetocreagris thailandica, Bisetocreagris cf. ussuriensis
(only a single tritonymph), Microbisium brevifemoratum, Anatemnus orites,
Chernes hahni, Allochernes tropicus n.comb., Lamprochernes savignyi,
Megachernes cf. titanius, Megachernes cf. vietnamensis, Verrucachernes
oca, Hyperwithius tonkinensis.
Key-words: Arachnida - Pseudoscorpiones - China - Review - Taxonomy.
INTRODUCTION
Although China covers a huge area within the Palaearctic and Oriental region,
only few records of pseudoscorpions are scattered in different papers (see citations in
the catalogue of HARVEY 1990). New collections at my disposal from that country,
mostly from the southern provinces, induced me to prepare a review of all known
pseudoscorpions (47 species) from China. It is not the purpose of this paper to figure
all morphological details of all species and/or to provide an identification key. Of
course this compilation is only a first step and far away from completeness. Some
series (for example in Bisetocreagris, Megachernes) remained unidentified or can be
determined only with some doubts. Further knowledge of pseudoscorpion taxonomy
and further investigations with special collecting techniques in vast and remote areas
of China may enlarge this first review in many aspects.
Manuscript accepted 10.06.1995.
1046 WOLFGANG SCHAWALLER
The Chinese pseudoscorpion fauna is composed of different elements. Records
from southern tropical and subtropical regions show a closer relation to the fauna of
Indochina (REDIKORZEV 1938, BEIER 1951), Thailand (SCHAWALLER 1994) and Nepal
(SCHAWALLER 1991). The fauna of the northeastern provinces has a palaearctic charac-
ter similar to eastern Siberia (SCHAWALLER 1995). The species from the northwestern
arid regions of China might have affinities with the Mongolian fauna. The pseudoscor-
pions from Korea have been treated recently (HONG & KIM 1993, Kim & HONG 1994).
In this review, "China" covers the area of the Peoples Republic of China
including Tibet, but without Mongolia and Taiwan.
Not included in this review are: Microcreagris japonica Ellingsen, 1907 from
Japan, which was recorded by CHAMBERLIN (1930) from China: Chofu; this seems to be
an error, because Chofu is a city near Tokyo. CHAMBERLIN (1930) described 3 further
species of "Microcreagris" with doubtful origin, which are treated here as nomina
dubia. Geogarypus irrugatus (Simon, 1899) was recorded from China (Amoy =
Xiamen) by CHAMBERLIN (1930); the genus needs a revision and probably the herein
recorded species javanus (Tullgren, 1905) is a synonym of irrugatus (Simon, 1899).
MATERIAL
CGG Collection G. Gardini, Genova.
MHNG Muséum d'Histoire Naturelle, Genève.
MSF Museo La Specola, Firenze.
MSNV Museo Civico di Storia Naturale, Verona.
SMNS Staatliches Museum für Naturkunde, Stuttgart.
ZISB Zoological Institute of the Academia Sinica, Beijing.
SPECIES LIST
Chthoniidae (7 species)
1. Centrochthonius kozlovi (Redikorzev, 1918) (Fig. 6)
Material: Not seen.
Remarks: The single specimen published from Nepal is not a tritonymph, but
an adult male, thus in the figures given in SCHAWALLER (1991) two additional tricho-
bothria are missing.
Distribution: Tibet (locus typicus), Nepal Himalayas.
2. Centrochthonius sichuanensis n.sp. (Figs 1-5)
Holotype (6 ): China, Sichuan Prov., Wolong Nature Reserve, 1500 m, 21.V.1994,
MHNG.
Paratypes: Together with holotype, 1 d 3 2 MHNG. Sichuan Prov., Wolong Nature
Reserve, 1700 m, 18.V.1994, 2 ¢ MHNG. Sichuan Prov., Wolong Nature Reserve, 1000 m,
24.V.1994, 2 3 1 2 SMNS 3509. Sichuan Prov., Mt. Emei, 1500 m, 21.IX.1994, 1 2 MHNG.
PSEUDOSCORPION FAUNA OF CHINA 1047
Description (6): Carapace (Fig. 1) (0.57/0.59 mm) with the basal margin
somewhat narrower than the anterior margin, anterior margin without epistome, each
side with 2 distinct lense eyes; carapace with 26 setae, anterior margin with 8 (4 long
and on each side with 2 shorter) setae, basal margin with 4 setae. Setation on tergites
4-8-9-8-10-10-10-8-8-6-4, setation around the genital opercula see Fig. 4. Palpal coxa
with 3 setae, coxa I with 1+4 setae and with 9-10 serrate coxal spines on a single
socle, coxa II with 2+3 setae, coxa III with 3+3 setae, coxa IV with 4+3 setae.
Fics 1-7
Centrochthonius sichuanensis n.sp. holotype male (1-5), C. kozlovi from Nepal (6) and C.
ussuriensis from Siberia (7); scale 0.5 mm. - 1: Carapace; 2: Palp from dorsal; 3: Chelicera; 4:
Setation around genital opercula; 5-7: Palpal chela from lateral.
1048 WOLFGANG SCHAWALLER
Chelicera (Fig. 3): 6 setae on the hand, movable finger with 1 seta; flagellum with 8
feathered blades; serrula exterior with 22, serrula interior with 17 blades; fixed finger
with a striking long tooth and distally with additional 3 teeth, movable finger with a
row of indistinct teeth. Palp (Figs 2, 5): Femur (1.00/0.18 mm) 5.6x, patella
(0.45/0.18 mm) 2.5x, chela without pedicel (1.55/0.33 mm) 4.7x longer than wide;
finger 1.6x longer than hand; trichobothriotaxie see Fig. 5; fixed finger with 13 errect
and acute teeth, movable finger with 18 errect and acute teeth. Leg IV: Tibia 0.66
mm, basitarsus 0.34 mm, telotarsus 0.64 mm long; basitarsus and telotarsus each with
a long seta distally. Body length 2.6 mm.
Description (9): No distinct sexual dimorphism. Palp: Femur (1.10/0.20 mm)
5.5x, patella (0.50/0.20 mm) 2.5x, chela without pedicel (1.74/0.42 mm) 4.15x longer
than wide; finger 1.3x longer than hand. Body length 3.0 mm.
Discussion: The new material from China distinctly differs from the 3 known
species of that genus (kozlovi (Redikorzev, 1918), schnitnikovi (Redikorzev, 1934)
and ussuriensis Beier, 1979) by the combination of the following characters: striking
body size, the absence of an epistome, the high number of setae on the carapace, few
and distant teeth on the palpal chela (Figs 5-7) and the striking tooth on the fixed
cheliceral finger.
3. Lagynochthonius sinensis (Beier, 1967)
Material: Not seen.
Distribution: China (Hupeh).
4. Lagynochthonius tonkinensis (Beier, 1951)
Material: Zhejiang Prov., Mt. Tianmu, 5.[X.1989, 8 ex. ZISB, 3 ex. SMNS 2898.
Yunnan Prov., Xishuangbanna, IV.-VII.1993, 110 ex. ZISB, 5 ex. SMNS 3427, 6 ex. SMNS
3429. Yunnan Prov., Mengleng Tropical Garden, 9.1V.1992, 22 ex. ZISB, 4 ex. SMNS 3437.
Yunnan Prov., Mengla-Mengleng, 10.1V.1992, 6 ex. ZISB. Yunnan Prov., Mengyang Nature
Reserve, 500 m, 10.-14.IX.1994, 4 ex. MHNG.
Remarks: These specimens possess the same chelal proportions and dentations
as material from Thailand (SCHAWALLER 1994); the chelal hand has about the same
length as the fingers. In the previous congener sinensis the hand is significantly
shorter.
Distribution: Vietnam (locus typicus), Thailand, China (new records).
5. Tyrannochthonius japonicus (Ellingsen, 1907) (Fig. 8)
Material: Yunnan Prov., Xishuangbanna, IV.-VII.1993, 1 ex. ZISB, 1 ex. SMNS 3431.
Yunnan Prov., Mengleng Tropical Garden, 9.1V.1992, 2 ex. SMNS 3439.
Remarks: Both males and females have a very distinct epistome and a den-
tation on the palpal fingers (Fig. 8), which is characteristic for japonicus (SATO 1979).
The genus 7yrannochthonius urgently needs revision to clarify, which characters are
PSEUDOSCORPION FAUNA OF CHINA 1049
species-specific and which are not. The species differs significantly from the both
following congeners, which have no distinct epistomes and a different chelal den-
tation (Figs 9-10).
Distribution: Japan (locus typicus), Taiwan, China (new records).
Fics 8-10
Palpal chela from lateral, scale 0.2 mm. - 8: Tyrannochthonius japonicus, SMNS 3431; 9:
Tyrannochthonius pachythorax, SMNS 3428; 10: Tyrannochthonius robustus, SMNS 2897.
6. Tyrannochthonius pachythorax Redikorzev, 1938 (Fig. 9)
Material: Yunnan Prov., Xishuangbanna, IV.-VII.1993, 19 ex. ZISB, 3 ex. SMNS 3428,
3 ex. SMNS 3430. Yunnan Prov., Kunming, 4.1V.1992, 2 ex. ZISB. Yunnan Prov., Mengleng
Tropical Garden, 9.1V.1992, 2 ex. ZISB. Yunnan Prov., Mengla-Mengleng, 10.1V.1992, 1 ex.
ZISB. Yunnan Prov., Mengla, 10.1V.1992, 1 ex. SMNS 3440. Yunnan Prov., Mengyang Nature
Reserve, 500 m, 10.-14.IX.1994, 2 ex. MHNG. Yunnan Prov., Kunming, montagne de l'ouest
(Xi Shan), 17.1V.1982, leg. M. Freiburghaus, 3 ex. MHNG. Sichuan Prov., Wolong Nature
Reserve, 1500 m, 21.V.1994, 2 ex. MHNG. Sichuan Prov., Wolong Nature Reserve, 1700 m,
18.V.1994, 1 ex. MHNG. Fujian Prov., Mt. Wuyi, 31.VII.-13.VII.1983, leg. H.F. Wang, 5 ex.
ZISB, 2 ex. SMNS 3444.
Remarks: Dentation on palpal fingers of the Xishuangbanna specimens see
Fig. 9. The length of the palpal fingers and the number of the teeth are variable to a
certain extent but without distinct differences, thus I hope not to fail in assigning all
series to a single species.
Distribution: Vietnam (locus typicus), Cambodia, Thailand (SCHAWALLER
1994), China (new records).
1050 WOLFGANG SCHAWALLER
7. Tyrannochthonius robustus Beier, 1951 (Fig. 10)
Material: Zhejiang Prov., Mt. Tianmu, V.-IX.1987, 36 ex. ZISB, 6 ex. SMNS 2896.
Hunan Prov., Mt. Hengshan, 1988, 1 ex. SMNS 2897. Sichuan Prov., Wolong Nature Reserve,
1000 m, 24.V.1994, 2 ex. MHNG. Sichuan Prov., Wolong Nature Reserve, 1500 m, 21.V.1994,
6 ex. MHNG, 3 ex. SMNS 3502. Sichuan Prov., Wolong Nature Reserve, 1700 m, 18.V.1994,
9 ex. MHNG. Sichuan Prov., Mt. Emei, 1500 m, 21.1X.1994, 5 ex. MHNG, 2 ex. SMNS 3508.
Sichuan Prov., Mt. Emei, 1800 m, 24.IX.1994, 6 ex. MHNG. Sichuan Prov., Dege County,
3500 m, 14.VIIL.1983, leg. H.F. Wang, | ex. ZISB. Shaanxi Prov., E Xian, Mt. Huashan, 1000
m, 9.-11.V.1994, 2 ex. MHNG. Beijing, no date, leg. H.F. Wang, | ex. ZISB, 1 ex. SMNS
3442. Beijing, 18.IX.1985, leg. H.F. Wang, 2 ex. ZISB.
Remarks: The pedipalp chela has a heterodentate dentation on both fingers
(Fig. 10), which is characteristic for this species. An epistome is lacking, in contrary
to the type material which is said to have a small epistome. The above cited speci-
mens possess on the medial side of the chelal hand a spine-like seta, which is also
figured by HARVEY (1988) for other congeners from the Krakatau Islands. Such a
thicker seta is lacking (or broken ?) in the preceding 2 species from China. Probably
this is a character which could help in a natural classification of the genus Tyran-
nochthonius.
Distribution: Vietnam (locus typicus), China (new records).
Tridenchthoniidae (I species)
8. Ditha proxima (Beier, 1951)
Material: Yunnan Prov., Mengyang Nature Reserve, 500 m, 10.-14.IX.1994, 3 ex.
MHNG, 1 ex. SMNS 3504.
Remarks: These specimens have the palpal chela somewhat more slender than
Himalayan populations (nearly as in /aosana Beier, 1951), but all other characters
show no differences (for example: trichobothrium st closer to sb and not just in the
middle between sb and t as in /aosana).
Distribution: Vietnam, Thailand, Bhutan, Nepal, China (new record).
Geogarypidae (1 species)
9. Geogarypus javanus (Tullgren, 1905)
Material: Yunnan Prov., Kunming, montagne de l'ouest (Xi Shan), 17.1V.1982, leg. M.
Freiburghaus, 2 ex. MHNG. Yunnan Prov., Mengyang Nature Reserve, 500 m, 10.-14.IX.1994,
1 ex. MHNG.
Remarks: This species has been redescribed by HARVEY (1988). Probably this
species is a synonym of irrugatus (Simon, 1899), which was recorded from China by
CHAMBERLIN (1930).
Distribution: Southeastern Asia, China (new record), Taiwan, New Guinea,
Salomon Islands.
PSEUDOSCORPION FAUNA OF CHINA 1051
Olpiidae (1 species)
10. Euryolpium agniae Redikorzev, 1938
Material: Not seen.
Distribution: Vietnam (locus typicus), China (BEIER 1967).
Neobisiidae (13 species)
11. Bisetocreagris sp. A (Fig. 16)
Material: Shaanxi Prov., E Xian, Mt. Huashan, 1000 m, 9.-11.V.1994, 3 ex. MHNG.
Remarks: In general, the identification of Bisetocreagris species is quite
doubtful and nearly impossible without a revision. So I avoid in creating new names,
even if it seems quite sure that the herein not named material represents undescribed
species. A lot of species are described only under a typological point of view without
the knowledge of morphological variability or sexual dimorphic pattern. Even the
splitting of the old paraphyletic Microcreagris in some genera (CURCIC 1983) should
be revised. Furthermore, CHAMBERLIN (1930) described 3 species from China (exact
localisations doubtful, even China is given with a question mark), which cannot be
recognized and which are treated herein as nomina dubia: Microcreagris lampra
Chamberlin, 1930, Microcreagris orientalis Chamberlin, 1930 and Microcreagris
silvestrii Chamberlin, 1930. See also remarks under 20. Chinacreagris nankingensis.
These specimens from the Huashan mountains have the palpal proportions
(Fig. 16) similar to annamensis (Figs. 14-15), but the patella is significantly shorter.
The granulation on the palps is quite weak and can easily be overlooked.
12. Bisetocreagris sp. B (Figs 17-18)
Material: Sichuan Prov., Mt. Emei, 1500 m, 21.IX.1994, 6 ex. MHNG. Sichuan Prov.,
Mt. Emei, 1800 m, 24.IX.1994, 4 ex. MHNG.
Remarks: These specimens have quite sexual dimorphic palps with a long
patella (Figs 17-18), all segments have no granulation.
13. Bisetocreagris annamensis (Beier, 1951) (Figs 14-15)
Material: Yunnan Prov., Xishuangbanna, IV.-VII.1993, 2 ex. ZISB, 2 ex. SMNS 3433.
Yunnan Prov., Mengyang Nature Reserve, 500 m, 10.-14.1X.1994, 5 ex. MHNG, 2 ex. SMNS
3505.
Remarks: Both series shows no significant differences to material just
published from Thailand (SCHAWALLER 1994). Proportions of the sexually dimorphic
palps of the Xishuangbanna specimens see Figs 14-15.
1052 WOLFGANG SCHAWALLER
Fics 11-20
Palpal proportions from dorsal, scale 1.0 mm. - 11: Microcreagris gigas, SMNS 2893; 12:
Chinacreagris chinensis, female, SMNS 2895; 13: Chinacreagris chinensis, male, SMNS
2894; 14: Bisetocreagris annamensis, female, SMNS 3433; 15: Bisetocreagris annamensis,
male, SMNS 3433; 16: Bisetocreagris sp. A, MHNG; 17: Bisetocreagris sp. B, male, SMNS
3511; 18: Bisetocreagris sp. B., female, MHNG; 19: Bisetocreagris thailandica, SMNS
3432; 20: Bisetocreagris indochinensis, male, SMNS 3434.
Distribution: Vietnam (locus typicus), Thailand (SCHAWALLER 1994), China
(new record).
14. Bisetocreagris indochinensis (Redikorzev, 1938) (Fig. 20)
Material: Yunnan Prov., Xishuangbanna, IV.-VII.1993, 1 ex. ZISB, 1 ex. SMNS 3434.
PSEUDOSCORPION FAUNA OF CHINA 1053
Remarks: This material is characterized by a distinct granulation on the palpal
femur, furthermore the club of the palpal patella is distinctly separated externally
from the stick. The palps are sexually dimorphic and somewhat more slender in males
(Fig. 20).
Distribution: Vietnam (locus typicus), Thailand (SCHAWALLER 1994), China
(new record).
15. Bisetocreagris kaznakovi (Redikorzev, 1918)
Material: Not seen.
Distribution: Tibet (locus typicus), Nepal (SCHAWALLER 1987, 1991), Sikkim
(kexsEGG):
16. Bisetocreagris thailandica Schawaller, 1994 (Fig. 19)
Material: Yunnan Prov., Xishuangbanna, IV.-VII.1993, 11 ex. ZISB, 3 ex. SMNS 3432.
Yunnan Prov., Kunming, 4.1V.1992, 2 ex. ZISB. Yunnan Prov., Mengleng Tropical Garden,
9.1V.1992, 5 ex. ZISB, 2 ex. SMNS 3438. Yunnan Prov., Mengla-Mengleng, 10.1V.1992, 5 ex.
ZISB.
Remarks: The Chinese series show no significant differences to the type series
from Thailand. This species is characterized by a relatively small body size, smooth
palps and short palpal fingers (Xishuangbanna specimens see Fig. 19) among other
characters. The specimens from Kunming, however, have an indistinctly granulated
palpal femur.
Distribution: Thailand (locus typicus), China (new records).
17. Bisetocreagris cf. ussuriensis (Redikorzev, 1934)
Material: Ilin Prov., Mt. Changbai, no date, leg. H.F. Wang, 1 ex. ZISB.
Remarks: This single record from the northeastern province is represented by a
tritonymph only, which probably belongs to ussuriensis (Redikorzev, 1934). Without
having adults a sure identification is impossible.
Distribution: Eastern Siberia (SCHAWALLER 1995), ? China (Ilin, new record).
18. Chinacreagris chinensis (Beier, 1943) (Figs 12-13)
Material: Zhejiang Prov., Mt. Tianmu, 1987-1989, 7 ex. ZISB. Hunan Prov., Mt.
Hengshan, 1988-1989, 1 ex. ZISB, 1 ex. SMNS 2894, 1 ex. SMNS 2895. Fujian Prov., Mt.
Wuyi, 31.VII.1983, leg. H.F. Wang, 3 ex. ZISB, 1 ex. SMNS 3445. Guizhou Prov., Huaxi,
X.1986, leg. G.M. de Rougemont, 7 ex. MHNG, 2 ex. SMNS 3460. Sichuan Prov., Mt. Emei,
1800 m, 24.KX.1994, 3 ex. MHNG.
Remarks: The measurements, the palpal proportions and the smooth palpal
femur coincide with the original description. A single specimen (SMNS 2894)
1054 WOLFGANG SCHAWALLER
possesses a few tubercles (not an even granulation as in gigas) on the palpal femur.
The series from Mt. Wuyi is represented by tritonymphs only but could fit to the
adults from the other adjacent localities. Proportions of the slightly sexually
dimorphic palps of the Mt. Hengshan specimens see Figs 12-13.
Distribution: China (locus typicus: Kiangsu).
19. Chinacreagris kwantungensis (Beier, 1967)
Material: Not seen.
Distribution: China (Kwangtung)
20. Chinacreagris nankingensis Curcic, 1983
Material: Not seen.
Remarks: The status of the genus Chinacreagris seems doubtful to me, because
the separation from Bisetocreagris by a somewhat different shape of the flagellum and
a somewhat different setation of the abdominal tergites is typological and far away
from scientific argumentation (see remarks under 11. Bisetocreagris sp. A).
Distribution: China (Kiangsu).
21. Microcreagris gigas Balzan, 1892 (Fig. 11)
Material: Zhejiang Prov., Mt. Tianmu, 1988-1989, 4 ex. ZISB, 2 ex. SMNS 2893.
Hunan Prov., Mt. Hengshan, 1.1988, 1 ex. ZISB. Sichuan Prov., Wolong Nature Reserve, 1000
m, 24.V.1994, 7 ex. MHNG, 4 ex. SMNS 3512. Sichuan Prov., Wolong Nature Reserve, 1700
m, 18.V.1994, 3 ex. MHNG. Sichuan Prov., Mt. Emei, 1500 m, 21.1X.1994, 2 ex. MHNG.
Remarks: The species, typus generis of Microcreagris, is redescribed by
MAHNERT (1979). The specimen from Mt. Hengshan coincides well by the distinct
even granulation, by the measurements and by the distinct epistome, the specimens
from Sichuan and from Mt. Tianmu, however, are partly quite smaller (palpal femur
1.2 mm minimum, Fig. 11) and the epistome is reduced. I hope not to fail in
considering these differences as infraspecific variations.
Distribution: China.
22. Stenohya chinacavernicola n.sp. (Figs 21-28)
Holotype (3): China, Sichuan Prov., Huaying, cave "Dei" (="Xian Nin"), 1065 m,
17.VI11.1993, leg. "Centro Ybleo Ricerche Speleo-Ydrogeoliche", CGG.
Paratype (2): China, Sichuan Prov., Huaying, cave "Chao-Tian", 930 m, 18.VIII.1993,
leg. "Centro Ibleo Ricerche Speleo-Idrogeologiche", CGG.
Description (4): Carapace (Fig. 23) smooth, (1.12/1.09 mm) 1.10x longer than
wide, epistome rounded and indistint, a single small eye spot without lense on each
side; carapace with 26 setae, anterior margin with 4, posterior margin with 7. Setation
PSEUDOSCORPION FAUNA OF CHINA 1055
1 HAN) Dn 27
i
A f
[ERRRAN SLITTA
PENNE
Fics 21-28
Stenohya chinacavernicola n.sp. holotype male, scales 1.0 mm - 21: Palp from dorsal; 22:
Palpal chela from lateral with trichobothriotaxie and dentation in the middle of the fingers; 23:
Carapace; 24: Chelicera; 25: Galea from lateral; 26: Flagellum; 27: Setation around. genital
opercula, 28: Leg IV IV with subterminal seta.
on tergites 9-8-10-10-12-12-13-14-13-12-10, on sternites x-x-18-18-18-19-18-18-16-
10, setation around the genital opercula see Fig. 27. Chelicera (Fig. 24): 7 setae on the
hand, movable finger with 1 seta; galea deeply bifurcated, each branch with 3 sub-
branches (Fig. 25); flagellum with 10 pinnate blades (Fig. 26); serrula with about 35
1056 WOLFGANG SCHAWALLER
blades; both fingers each with about 8 small and rounded teeth. Pedipalp (Figs 21-22):
all segments smooth, without granulation; femur (2.04/0.34 mm) 6.0x, patella (2.06/
0.44 mm) 4.7x, chela without pedicel (3.30/0.68 mm) 4.86x longer than wide; pedicel
of patella as long as club; finger somewhat longer than hand; trichobothriotaxie see Fig.
22, ist at the base of the fixed finger, b and sb separated; both palpal finger with about
120 equal and closely situated teeth, teeth on the fixed finger acute and pointing
backwards. Medial process of the palpal coxa with 5+1 long setae. Leg IV (Fig. 28):
femur and patella 2.18 mm, tibia 1.95 mm, basitarsus 0.66 mm, telotarsus 0.89 mm
long; telotarsus 1.35x longer than basitarsus; subterminal seta bifurcated and with
serrate branches. Body length 4.5 mm.
Description (2): Sexual dimorphism indistinct. Flagellum and galea as in the
male. Trichobothriotaxie and setation as in the male. Palpal proportions similar to those
in the male: femur (2.02/0.36 mm) 5.7x, patella (1.90/0.40 mm) 4.75x, chela without
pedicel (3.00/0.68 mm) 4.4x longer than wide. Body length 4.8 mm.
Discussion: This first cavernicolous species of the genus can easily be separated
from the "free" living congeners by the palpal proportions with a long pedicel of the
patella, by a relatively high number of setae on the carapace (which might be an
adaption to cave life) and mainly by the trichobothriotaxie: sb just between b and st and
not close to b, ist at the base of the fixed finger. This trichobothrial pattern is quite
unusual within the genus and it could be necessary to place this species in an own genus
after having revised the genus validity around "Microcreagris". The number of the
flagellar blades seems variable in the genus: for example in martensi and mahnerti 8, in
hamatus 7-10 and in chinacavernicola n.sp. 10 pinnate blades.
23. Microbisium brevifemoratum (Ellingsen, 1903)
Material: Ilin Prov., Mt. Changbai, no date, leg. H.F. Wang, 1 ex. ZISB. Shaanxi Prov.,
E Xian, Mt. Huashan, 1000 m, 9.-11.V.1994, 9 ex. MHNG, 3 ex. SMNS 3510. Beijing, no date,
leg. H.F. Wang, 1 ex. ZISB. Beijing, 18.IX.1985, leg. H.F. Wang, 2 ex. SMNS 3443.
Remarks: These Chinese specimens show no differences to eastern Siberian
records.
Distribution: Wider distribution in the Palaearctic region from Europe to
eastern Siberia including Sakhalin and Kuriles, China (Ilin, Qinling, Beijing, new
records).
Cheiridiidae (1 species)
24. Cheiridium minor Chamberlin, 1938.
Material: Not seen.
Remarks: The status of this species seems doubtful, probably it is a synonym
of museorum (Leach, 1817).
PSEUDOSCORPION FAUNA OF CHINA 1057
Atemnidae (4species)
25. Anatemnus orites (Thorell, 1889)
Material: Yunnan Prov., Xishuangbanna, IV.-VII.1993, 2 ex. ZISB, 1 ex. SMNS 3435.
Yunnan Prov., Mengyang Nature Reserve, 500 m, 10.-14.[X.1994, 5 ex. MHNG.
Remarks: The species taxonomy within the genus Anatemnus is doubtful
(SCHAWALLER 1994). This Yunnan material coincides with material from Thailand,
which originates from an area close to the locus typicus of orites (Burma).
Distribution: Burma (locus typicus), southeastern Asia, China (new record).
26. Atemnus politus (Simon, 1878)
Material: Not seen.
Distribution: Widely distributed in the Palaearctic region, known also from
Mongolia, Bhutan and China (BEIER 1967 sub turkestanicus).
27. Diplotemnus insolitus Chamberlin, 1933
Material: Not seen.
Remarks: Miratemnus piger sinensis Schenkel, 1953 is transferred by HARVEY
(1990) to Diplotemnus ophthalmicus Redikorzev, 1949, which is synonymized by
DASHDAMIROV & SCHAWALLER (1993) with insolitus Chamberlin, 1933.
Distribution: Wider distribution in the Palaearctic region from Algeria to
Middle Asia, Sudan, Himalayas (locus typicus of insolitus) and China ("Süden des
Ordos", localisation ?).
28. Paratemnoides sinensis (Beier, 1932)
Material: Not seen.
Distribution: China (Kwangtung).
Given enindiave (6:species)
29. Dactylochelifer gansuensis Redikorzev, 1934
Material: Not seen.
Distribution: China (Kansu).
30. Eremochernes secundus Beier, 1937
Material: Not seen.
Remarks: This genus belongs to the Cheliferidae near Rhacochelifer (BEIER
1973) and includes 2 species (gracilipes Redikorzev, secundus Beier) from Mongolia
1058 WOLFGANG SCHAWALLER
and northwestern China respectively. Eremochernes tropicus Beier, 1967 from
Thailand and Sichuan is transferred herein provisionally to the chernetid genus
Allochernes Beier, 1932 (see 36.).
Distribution: China (Liaoning).
31. Lophochernes gracilis Beier, 1943
Material: Not seen.
Distribution: China (Fujian).
32. Lophochernes tibetanus Beier, 1943
Material: Not seen.
Distribution: Tibet.
33. Macrochelifer tibetanus (Redikorzev, 1918)
Material: Not seen.
Distribution: China (Chinghai).
34. Sinochelifer kwantungensis Beier, 1967
Material: Not seen.
Distribution: China (Kwangtung).
Chernetidae (I0 species)
35. Allochernes asiaticus (Redikorzev, 1922)
Material: Not seen.
Distribution: Middle Asia, Nepal, Tibet.
36. Allochernes tropicus (Beier, 1967) n.comb.
Material: Sichuan Prov., Xiangcheng County, 2800 m, 7.VII.1982, leg. H.F. Wang, 1
ex. SMNS 3459.
Remarks: This single male coincides well with the description of tropicus
Beier, 1967 (sub Eremochernes Beier, 1932), concerning for example the proportions
of the palps and in having 3 flagellar setae, the serrula with 19 lamellae (description:
22), the tarsus IV distally with a tactile seta and the palpal chela with several
accessory teeth. However, Eremochernes (see 30.) belongs to the Cheliferidae (BEIER
1973), thus tropicus must be transferred to a chernetid genus. Because of the
unsatisfactory taxonomic situation concerning the genus separation in this family,
PSEUDOSCORPION FAUNA OF CHINA 1059
tropicus Beier, 1967 can be assigned only provisionally to the genus Allochernes
Beier, 1932. The presence of a tarsal tactile seta is quite "unusual" among the
congeners, but Allochernes liwa Harvey, 1988 from Sumatra also possesses such a
seta.
Distribution: Thailand, China (Sichuan).
37. Chernes hahni (Koch, 1873)
Material: Beijing, no date, leg. H.F. Wang, 2 ex. ZISB, 1 ex. SMNS 3441.
Remarks: This series shows no differences in the palpal proportions, the
setation, the granulation and in the form of the galea to European and eastern Siberian
specimens.
Distribution: Europe, Caucasus, northern Iran, Kasakhstan, Siberia, Sakhalin,
China (Beijing, new record).
38. Chernes sinensis Beier, 1932
Material: Not seen.
Remarks: The status of this species seems doubtful. The description point to a
short tactile seta on the tarsus IV, which is unusual for Chernes but characteristic for
Dinocheirus.
Distribution: China (Kwangtung).
39. Lamprochernes savignyi (Simon, 1881)
Material: Guizhou Prov., Huaxi, X.1986, leg. G.M. de Rougemont, 1 ex. MHNG.
Distribution: Cosmopolitan, China (new record).
40. Megachernes himalayensis (Ellingsen, 1914)
Material: Not seen.
Distribution: Himalayas, China (Sichuan, BEIER 1932 sub sinensis).
41. Megachernes cf. himalayensis (Ellingsen, 1914)
Material: Guangxi Prov., Contea Gongcheng, cave "Hei Yan", VIII.1994, leg. R.
Zorzin, 1 ex. MSNV.
Remarks: This single male is very similar in many characters (specially palpal
proportions) to the few specimens which I know from the Himalayas. However, the
body is somewhat smaller, the last coxa is not swollen and the setation on the tergites
and sternites is somewhat different, so probably we face a different (? new) species.
Without having both sexes and without a revision of the Asian congeners I avoid in
1060 WOLFGANG SCHAWALLER
describing this form/species. In this connection it should be checked in the type
material if sinensis Beier, 1932 is really a synonym of himalayensis (Ellingsen, 1914).
Distribution: see 40.
42. Megachernes cf. titanius Beier, 1951
Material: Yunnan Prov., Mengyang Nature Reserve, 500 m, 10.-14.IX.1994, 3 ex.
MHNG, 1 ex. SMNS 3506.
Remarks: This series consists of 1 male and 3 females, which are somewhat
smaller (body length about 4 mm) than noted in the description of fitanius (6 4.5 mm,
2 5 mm), but distinctly bigger than himalayensis. Both sexes show no distinct
dimorphism concerning the setation on the carapace and the palps but a distinct
dimorphism concerning the proportions of the palpal patella and fingers. The palpal
patella in males is round and thick (as figured in the original description) and in
females distinctly more slender; the fingers are somewhat longer in females than in
males.
Distribution: Vietnam, China (new record).
43. Megachernes cf. vietnamensis Beier, 1967
Material: Hubei Prov., Xing Shan, cave "Da Dang", 1992, leg. J. Lips, 1 ex. MHNG.
Hubei Prov., Yishang, cave "Cygne", 1992, leg. J. Lips, 1 ex. MHNG. Sichuan Prov., Huaying,
cave "Chao-Tian", 930 m, 18.VIII.1993, leg. "Centro Ibleo Ricerche Speleo-Idrogeologiche", 1
exe EGG:
Remarks: Megachernes vietnamensis is significantly different from the conge-
ners by its slender palps (in particular the femur) without distinct sexual dimorphism.
The above listed specimens also possess such slender palps, but the palpal hand is
somewhat more convex both on the lateral and on the medial side. Without further
material it seems impossible to decide, whether this is a infraspecific variation or a
character for separating different biospecies.
Distribution: Vietnam, China (new record).
44. Verrucachernes oca Chamberlin, 1947
Material: Yunnan Prov., Xishuangbanna, IV.-VII.1993, 2 ex. ZISB, 1 ex. SMNS 3436.
Yunnan Prov., Mengyang Nature Reserve, 500 m, 10.-14.1X.1994, 10 ex. MHNG. Sichuan
Prov., Wolong Nature Reserve, 1700 m, 18.V.1994, 5 ex. MHNG, 2 ex. SMNS 3503.
Remarks: The species is redescribed by HARVEY (1988). Probably, Pselapho-
chernes indicus Beier, 1974 is a synonym of Verrucachernes oca, at least the material
which I recorded from Nepal (SCHAWALLER 1991) and Thailand (SCHAWALLER 1994)
belongs to Verrucachernes oca and shows no differences to the new Chinese records.
Distribution: Wide distribution in southeastern Asia, China (new record), New
Guinea, Solomon Islands.
PSEUDOSCORPION FAUNA OF CHINA 1061
Withiidae (3 species)
45. Hyperwithius tonkinensis Beier, 1951
Material: Sichuan Prov., Mt. Emei, 1700 m, 23.IX.1994, 1 ex. MHNG, 1 ex. SMNS
5307.
Remarks: This genus contains 3 "species": annamensis (Redikorzev, 1938),
dawydoffi Beier, 1951 and tonkinensis Beier, 1951, which are mainly separated by the
proportions of the palps and the setation on tergites and sternites. This might be only a
typological separation, furthermore variability of the characters is unknown. The
above listed 2 females fit quite well with the description of tonkinensis, based on |
female and 1 male. In comparison with Metawithius spiniventer, which can be
expected also in that region, the congeners of Hyperwithius are smaller, the form of
the carapace is different and the trichobothrium on tarsus IV inserts somewhat distally
and not just in the middle of the tarsus.
Distribution: Vietnam, China (new record).
46. Withius pekinensis (Balzan, 1892) (Figs 29-32)
Material: China, Peking, 1 © (holotype) MSF.
Description (by V. Mahnert): Carapace coarsely granulate (0.69/0.49 mm),
1.4x longer than wide, with 2 distict transverse furrows, 2 eyes present; setae of
carapace and tergites clavate, mostly broken; tergites and sternites divided, tergal
chaetotaxy not noted; setation of half-sternites 4/5-4-7-6/7-7-8-7-6, with 2 and 3
suprastigmal setae and 1 anterior lateral seta, a few sensory (?) setae present: VI 1,
VII 2-3, VII 2, IX 1 (?); anterior genital operculum with 12 setae. Spermatheca (Fig.
32) with paired short tubes. Chelicera: 5 setae on the hand (db and ib dentate),
movable finger with | seta, galea broken (on both fingers), serrula with 16 blades,
flagellum with 4 setae, fixed finger with 3 teeth, movable finger with cone-like
subapical lobe. Pedipalp (Fig. 29): all segments granulate; trochanter with pointed
dorsal hump, femur (0.55/0.18 mm) 3.0x, patella (0.55/0.21 mm) 2.7x, chelal hand
with pedicel (0.54/0.28 mm) 1.9x longer than wide and 1.35x longer than finger,
length of finger 0.40 mm, chela with pedicel (length 0.88 mm) 3.1x longer than wide;
trichobothriotaxie see Figs 30-31, ist dorsal, it internal. Leg I: femur (0.13/0.12 mm)
1.0 x, patella (0.27/0.11mm) 2.3x longer than wide and 2.1x longer than femur, tibia
(0.27/0.07 mm) 3.7x, tarsus (0.24/0.05 mm) 4.7x longer than wide; leg IV: femur and
patella (0.51/0.17 mm) 3.0x, tibia (0.41/0.10 mm) 4.1x, tarsus (0.30/0.06 mm) 4.7x
longer than wide; all setae broken, arolia broken, claws smooth.
Remarks: The trichobothrial pattern and the pedipalpal measurements are quite
similar to those of Withius piger (Simon, 1878), but the spermatheca seems to be
different. Males should be available before the specific identity of this species can be
established.
Distribution: China (Hopeh).
1062 WOLFGANG SCHAWALLER
Fics 29-32
Withius pekinensis, holotype female, scales 0.3 mm (29, 30), 0.1 mm (32). - 29: Palp from
dorsal; 30: Palpal chela with trichobothriotaxie; 31: Palpal fingers; 32: Spermatheca.
47. Withius piger (Simon, 1878)
Material: Not seen.
Distribution: Cosmopolitan, China (Kwangtung).
PSEUDOSCORPION FAUNA OF CHINA 1063
ACKNOWLEDGMENTS
Many thanks are due to Prof. Dr. Song Daxiang (Beijing), who trusted me with
the valuable material for study, who allowed me to keep duplicates and who helped in
transscribing Chinese labels into English. Dr. Giulio Gardini (Genova) transferred to
me a few cavernicolous species from the Verona museum and from his own
collection. Dr. Volker Mahnert (Geneva) loaned me the material from the Geneva
museum, allowed me to use his redescription of Withius pekinensis and helped with
the identification of Stenohya chinacavernicola n.sp. and Allochernes tropicus.
REFERENCES
BEIER, M. 1932. Pseudoscopionidea II. Subord. Cheliferinea. Das Tierreich 58: 1-294.
BEIER, M. 1951. Die Pseudoscorpione Indochinas. Mém. Mus. natn. Hist. nat. (ser. A) 1: 47-123.
BEIER, M. 1967. Pseudoscorpione vom kontinentalen Siidost-Asien. Pacific Insects 9: 341-369.
BEIER, M. 1973. Pseudoskorpione aus der Mongolei. Ann. naturhist. Mus. Wien 77: 167-172.
CHAMBERLIN, J.C. 1930. A synoptic classification of the false scorpions or chela-spinners, with
a report on a cosmopolitan collection of the same. - Part II. The Diplosphyronida
(Arachnida, Chelonethida). Annals Mag. nat. Hist. (ser. 10) 5: 1-49, 585-620.
Curcic, B.P.M. 1983. A revision of some Asien species of Microcreagris Balzan, 1892
(Neobisiidae, Pseudoscorpiones). Bull. Br. arachnol. Soc. 6: 23-36.
DASHDAMIROV, S. & W. SCHAWALLER, 1993. Pseudoscorpions from Middle Asia, Part 3
(Arachnida: Pseudoscorpiones). Stuttgarter Beitr. Naturk. (A) 497: 1-16.
HARVEY, M.S. 1988. Pseudoscorpions from the Krakatau Islands and adjacent regions, Indo-
nesia (Chelicerata: Pseudoscorpionida). Mem. Mus. Victoria 49: 309-353.
HARVEY, M.S. 1990. Catalogue of the Pseudoscorpionida. 1-726. Manchester, Manchester
University Press.
Hong, Y. & T.H. Kim, 1993. A check list and key to the order Pseudoscorpiones of Korea with
description of one new species from the genus Tyrannochthonius. Korean J. Ent. 23: 1-4.
Kim, T.H. & Y. HONG, 1994. Two new species of Chthoniidae (Arachnida: Pseudoscorpiones)
from Korea. Korean J. syst. Zool. 10: 47-53.
MAHNERT, V. 1979. The identity of Microcreagris gigas Balzan (Pseudoscorpiones, Neo-
bisiidae). Bull. Br. arachnol. Soc. 4: 339-341.
REDIKORZEV, V. 1938. Les pseudoscorpions de l'Indochine francaise recueillis par M.C.
Dawydoff. Mém. Mus. natn. Hist. nat. (NS) 10: 69-116.
SATO, H. 1979. Pseudoscorpions from Mt. Takao, Tokyo. Mem. Educ. Inst. Priv. Schools Japan
64: 79-105.
SCHAWALLER, W. 1987. Neue Pseudoskorpion-Funde aus dem Nepal-Himalaya, II (Arachnida:
Pseudoscorpiones). Senckenbergiana biol. 68: 199-221.
SCHAWALLER, W. 1991. Neue Pseudoskorpion-Funde aus dem Nepal-Himalaya, II (Arachnida:
Pseudoscorpiones). Revue Suisse Zool. 98: 769-789.
SCHAWALLER, W. 1994. Pseudoskorpione aus Thailand (Arachnida: Pseudoscorpiones). Revue
Suisse Zool. 101: 725-759.
SCHAWALLER, W. 1995. Review of the pseudoscorpion fauna of the Far East of Russia
(Arachnida: Pseudoscorpionida). Arthropoda Selecta 3: 123-126.
REVUE SUISSE DE ZOOLOGIE
Tome 102 — Fascicule 4
Zoologia 95, Zürich, 15-17 March 1995 (Annual Conference of the Swiss
ZLOOLO SI Cal SOCIE LY) MANS MEURT ara a as LOS UE Ronen En A
Watt, Ward B. Allozymes in evolutionary genetics: beyond the twin pit-
fallstofsgNeutralisma and =Selectionismn 25) men eo ee es.
POMIANKOWSKI, Andrew & Yoh IwAsA. What causes diversity in male sexual
characters sin Seth ae MEME Raia CASINA RN RE eC AI
HOFER, Heribert & Marion L. EAST. Virilized sexual genitalia as adaptations
Ofsiemalespottedshyacnasyte. MERE O TETE RE
LANG, Claude & Olivier REYMOND. Recovery from eutrophication delayed in
the profundal of Lake Neuchâtel: evidence from the oligochaete com-
MINUTE Sire SSL I ASP LATTA A RNA ET SEN OSE
MAHUNKA, Sandor. Oribatids from Brunei I (Acari: Oribatida). ...........
KURBATOV, Serguei A. Sur les Euthiini et Cephenniini (Coleoptera, Scyd-
maenidae) de |’Extréme-Est de la Russie et du Japon. ...............
ASSING, Volker. The types of some species of Lathrobium Gravenhorst and
of Xantholinus sejugatus G. Benick (Coleoptera: Staphylinidae)......
PERREAU, Michel. Espèces nouvelles ou mal connues de Cholevidae (Cole-
OPICIA) SERE LES RS ar IAA OA IAA DEMI ISO
KNOFLACH, Barbara. Two remarkable afromontane Theridiidae: Probosci-
dula milleri n. sp. and Robertus calidus n. sp. (Arachnida, Araneae). . .
Macko, Josef K. & Claude VAUCHER. Redescription and lectotype designa-
tion of Aploparaxis pseudofilum (Clerc, 1902) (Cestoda: Hymeno-
lépididae ) er ac eA U EE RE RAR RU
BURCKHARDT, Daniel & Stanislaw Adam SLIPINSKI. A review of the Passan-
dridae of the world (Coleoptera, Cucujoidea). IV. Genus Ancistria. . . .
SCHAWALLER, Wolfgang. Review of the Pseudoscorpion Fauna of China
(ArachnıdasBseudescorpionida)e ar a2 2a
Pages
853-867
869-882
883-894
895-906
907-912
913-942
943-959
961-968
969-978
979-988
989-994
995-1044
1045-1064
REVUE SUISSE DE ZOOLOGIE
Volume 102 — Number 4
Zoologia 95, Zürich, 15-17 March 1995 (Annual Conference of the Swiss
DOO SI CALS OCIELY)) LUN AS obo. ash eva ASI io TIA CR PERTE
WATT, Ward B. Allozymes in evolutionary genetics: beyond the twin pit-
fallsomeNeutralismeand Selectonisne er oe 2
POMIANKOWSKI, Andrew & Yoh Iwasa. What causes diversity in male
sexualscharaelters er arrosti E
HOFER, Heribert & Marion L. EAST. Virilized sexual genitalia as adaptations
oßremalesspottedihyaenas. 2.1.2 0. SES
LANG, Claude & Olivier REYMOND. Recovery from eutrophication delayed
in the profundal of Lake Neuchatel: evidence from the oligochaete
COMMUNE ST ek as cer M nel Ve ty ae ee
MAHUNKA, Sandor. Oribatids from Brunei I (Acari: Oribatida). ..........
KURBATOV, Serguei A. On the Euthiini and Cephenniini (Coleoptera, Scyd-
maenidae) from Russian Far East and Japan. ....................
ASSING, Volker. The types of some species of Lathrobium Gravenhorst and
of Xantholinus sejugatus G. Benick (Coleoptera: Staphylinidae)......
PERREAU, Michel. New or poorly known species of Cholevidae (Coleoptera).
KNOFLACH, Barbara. Two remarkable afromontane Theridiidae: Probosci-
dula milleri n. sp. and Robertus calidus n. sp. (Arachnida, Araneae). . .
Macko, Josef K. & Claude VAUCHER. Redescription and lectotype designa-
tion of Aploparaxis pseudofilum (Clerc, 1902) (Cestoda: Hymeno-
lepri) RE TE II LEO GRAS E
BURCKHARDT, Daniel & Stanislaw Adam SLIPINSKI. A review of the Passan-
dridae of the world (Coleoptera, Cucujoidea). IV. Genus Ancistria. . . .
SCHAWALLER, Wolfgang. Review of the Pseudoscorpion Fauna of China
(ArachnidazBseudoseorpiontda) 2a re RESO
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