cn ®, © tu e © NN LL. Q LL. n — n Lal-rd #iÿ > CL 5 A SWISS JOURNAL OF ZOOL REVUE SUISSE DE ZOOLOGIE TOME 118 — FASCICULE 4 Publication subventionnée par: ACADÉMIE SUISSE DES SCIENCES NATURELLES (SCNAT) VILLE DE GENÈVE SOCIÉTÉ SUISSE DE ZOOLOGIE Comité de rédaction DANIELLE DECROUEZ Directrice du Muséum d'histoire naturelle de Genève ALICE CIBOIS, PETER SCHUCHERT Chargés de recherche au Muséum d’histoire naturelle de Genève Comité de lecture A. Cibois (oiseaux), G. Cuccodoro (coléoptères), S. Fisch-Muller (poissons), B. Merz (insectes, excl. coléoptères), J. Mariaux (invertébrés excl. arthropodes), M. Ruedi (mammifères), A. Schmitz (amphibiens, reptiles), P. Schwendinger (arthropodes excl. insectes). ; Le comité soumet chaque manuscrit pour évaluation à des experts d'institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: taxonomie, systématique, faunistique, phylogénie, évolution, morphologie et anatomie comparée. Administration MUSÉUM D'HISTOIRE NATURELLE 1211 GENÈVE 6 Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm | PRIX DE L’ABONNEMENT: SUISSE Fr. 225.— UNION POSTALE Fr. 250.— (en francs suisses) Les demandes d’abonnement doivent être adressées à la rédaction de la Revue suisse de Zoologie, Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse ANNALES de la SOCIÉTÉ SUISSE DE ZOOLOGIE et du MUSÉUM D'HISTOIRE NATURELLE de la Ville de Genève tome 118 fascicule 4 204 Ë à ; FL GENEVE DÉCEMBRE 2011 ISSN 0035 - 418 X REVUE SUISSE DE ZOOLOGIE SWISS JOURNAL OF ZOOLOGY REVUE SUISSE DE ZOOLOGIE TOME 118 — FASCICULE 4 Publication subventionnée par: ACADÉMIE SUISSE DES SCIENCES NATURELLES (SCNAT) VILLE DE GENÈVE SOCIÉTÉ SUISSE DE ZOOLOGIE Comité de rédaction DANIELLE DECROUEZ Directrice du Muséum d'histoire naturelle de Genève ALICE CIBOIS, PETER SCHUCHERT Chargés de recherche au Muséum d'histoire naturelle de Genève Comité de lecture A. Cibois (oiseaux), G. Cuccodoro (coléoptères), S. Fisch-Muller (poissons), B. Merz (insectes, excl. coléoptères), J. Mariaux (invertébrés excl. arthropodes), M. Ruedi (mammifères), A. Schmitz (amphibiens, reptiles), P. Schwendinger (arthropodes excl. insectes). Le comité soumet chaque manuscrit pour évaluation à des experts d'institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: taxono- mie, systématique, faunistique, phylogénie, évolution, morphologie et anatomie comparée. Administration MUSÉUM D'HISTOIRE NATURELLE 1211 GENÈVE6 Internet: http://www.ville-ce.ch/musinfo/mhng/page/rsz.htm PRIX DE L’ ABONNEMENT: SUISSE Fr. 225.— UNION POSTALE Fr. 250.— (en francs suisses) Les demandes d'abonnement doivent être adressées à la rédaction de la Revue suisse de Zoologie, Muséum d'histoire naturelle. C.P. 6434, CH-1211 Genève 6. Suisse REVUE SUISSE DE ZOOLOGIE 118 (4): 599-637; décembre 2011 On two Heptathela species from southern Vietnam, with a discussion of copulatory organs and systematics of the Liphistiidae (Araneae: Mesothelae) Peter J. SCHWENDINGER! & Hirotsugu ONO? Muséum d'histoire naturelle de la Ville de Genève, c. p. 6404, CH-1211 Genève 6, Switzerland. E-mail: peter.schwendinger@ville-ge.ch 2 Department of Zoology, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba-shi, Ibaraki-ken, 305-005, Japan. E-mail: ono(@kahaku.go.jp On two Heptathela species from southern Vietnam, with à discussion of copulatory organs and systematics of the Liphistiidae (Araneae: Mesothelae). - Songthela australis Ono, 2002 is re-defined on the basis of new material from the type locality and from a second locality nearby. The male of this species 1s described for the first time. A closely related species, Heptathela nui sp. n., is described from males and females collected at two other localities in the same province. Variation in the copulatory organs of both species 1s 1llustrated and notes are given on their biology, particularly on their association with an ectoparasitic mite species of the genus Ljunghia Oudemans, 1932. “Twig-lining” is reported for one population of Æ. nui sp. n. Songthela australis and H. nui sp. n. are compared with Nanthela ton- kinensis (Bristowe in Bristowe & Millot, 1933) and Æ. tomokunii Ono, 1997a from northern Vietnam, with Æ. kimurai (Kishida, 1920), H. kikuyai Ono, 1998, Ryuthela owadai Ono, 1997b and R. nishihirai (Haupt, 1979) from Japan, and with Liphistius ornatus Ono & Schwendinger, 1990 and Z. thaleri Schwendinger, 2009 from Thailand. Additional taxonomic charac- ters are 1llustrated for most of these species. Judging from apomorphies in the male copulatory organs, S. australis and H. nui Sp. n. are sister species, but their female copulatory organs are very different. This challenges the generic concepts of Songthela Ono, 2000 and Abcathela Ono, 2000 (the latter earlier placed in the synonymy of Heptathela). The genus Nanthela Haupt, 2003 has no clear apomorphies, and transitions of diagnostic characters between Nanthela, Songthela and Heptathela were found. Thus Nanthela and Songthela are here placed in the synonymy of Heptathela. Morphology and conflicting terminology of copulatory organs in Liphis - tiidae, and the current confusion in Heptathelinae systematics are discussed. The copulatory organs of males and females of Liphistius Schiôdte, 1849 and Ryuthela Haupt, 1983 are here considered as more derived than those of Heptathela. Some observations on moulting of hphistiid spiders are given. Keywords: Taxonomy - new species - Liphistius - Nanthela - Ryuthela - Songthela - Ljunghia - Acari. Manuscript accepted 14.08.2011 600 P. J. SCHWENDINGER & H. ONO INTRODUCTION The Heptathelinae (considered as a family by Haupt, 2003: 69) is a group of 41 species (including the new one described in here), which are very distinct in their genital morphology from the 49 currently known species of Liphistiinae (all in the genus Liphistius) and which are also geographically separated. Heptathelinae occur in eastern Asia, Liphistiinae in southeastern Asia, both ranges roughly divided (but not everywhere; Schwendinger, in preparation) by the river Mekong. Originally only comprising the genus Æeptathela Kishida, 1923, other genera were added to the Heptathelinae as the number of species increased: Ryuthela Haupt, 1983, Abcathela Ono, 2000, Songthela Ono, 2000, Vinathela Ono, 2000, Nanthela Haupt, 2003 and Sinothela Haupt, 2003. Unfortunately many of the recently added genera were based on characters of a single sex. The discovery of heptatheline spiders at four localities in Lam Dong Province, including the type locality of Songthela australis, by one of us (PJS) in 2003 yielded the previously unknown male of that species and a new, closely related one. With this also came a surprise: the male copulatory organs of both species are very similar, but the female copulatory organs so strongly different that, according to the current system, they would need to be placed in different genera. Thus an examination of additional heptatheline species had to be carried out to re-evaluate the existing generic concepts in this group, and this further revealed serious confusion in the terminology used to describe copulatory organs of liphistiid spiders in the literature. MATERIAL AND METHODS External morphology was studied and drawn with a Zeiss SVII stereomicro- scope, the vulvae (tissue removed with forceps) with a Nikon Optiphot compound microscope (both with a drawing tube). SEM micrographs were taken with a Zeiss DSM-940A scanning electron microscope; prosoma photos were taken at several focal planes with a digital camera on a Leica MZ APO stereomicroscope and assembled with the AutoMontage® system. Hairs were partly or completely omitted in the drawings of the male palp. Body measurements were taken with a stereomicroscope and are given in mm. The total body length includes the chelicerae. The carapace length was measured in a slightly forward-inclined position, with the anterior and the posterior margins at the same focal plane. Lengths of leg articles and palpal articles were mea- sured on the dorsal side, from midpoint of anterior margin to midpoint of posterior margin, and are given in the following order: total (femur + patella + tibia + metatarsus + tarsus). | Terminology of the copulatory organs in Heptathelinae largely follows Song & Haupt (1984), but what they called “receptacles” (other authors called them bursae) are here called “receptacular clusters” (except for the Æ. tomokunii females examined). The term “genital plate” is inappropriate for Heptathelinae because that refers to the ventral wall of the genital atrium plus the entire bursa copulatrix with its dorsal and ventral walls, all of which are unsclerotised. “Slerotised” refers to areas in the cuticle that are hard, stiff and strongly pigmented; “unsclerotised” refers to cuticular structures that are thin and unpigmented or thick and moderately pigmented (as is the bursa copulatrix) but that are always flexible. We use “cymbial projection” as a shorter form of Haupt’s (2003: 69) “distal projection on the ventral side (of the cymbium)”, which HEPTATHELA FROM VIETNAM 601 can also be described as the ‘“elongate proventral-distal lobe of the cymbium”. “Carapace” is used instead of “dorsal plate of prosoma”. The informal appellation “allotype” refers to the paratype on which the description of the female of the new species is based. Positioning (dorsal, ventral, etc.) on the palpal organ refers to an unexpanded, outstretched palp. In the following text generic names, which are currently in synonymy or which are here placed in synonymy, are given between inverted commas. Abbreviations not explained in the figure legends are: AME, ALE, PME, PLE = anterior (posterior) median (lateral) eyes; IEBR = Institute of Ecology and Biological Resources of the Vietnamese Academy of Science and Technology, Hanoï, Vietnam; MHNG = Muséum d'histoire naturelle de la Ville de Genève, Switzerland; MOQ - median ocular quadrangle; MNHN = Muséum national d'Histoire naturelle, Paris, France; NSMT = National Museum of Nature and Science (formerly National Science Museum), Tokyo, Japan. TAXONOMY Liphistiidae Thorell, 1869 Heptathelinae Kishida, 1923 Heptathela Kishida, 1923 Abcathela Ono, 2000; type species Heptathela abca Ono, 1999: placed in the synonymy of Heptathela by Haupt, 2003: 71, 79. Nanthela Haupt, 2003; type species Liphistius tonkinensis Bristowe in Bristowe & Millot, 1933; syn. n. Sinothela Haupt, 2003; type species Heptathela sinensis Bishop & Crosby, 1932; placed in the synonymy of Songthela by Platnick, 2011 (in an earlier version of that online catalogue): Syn. n. Songthela Ono, 2000; type species Heptathela hangzhouensis Chen, Zhang & Zhu, 1981; placed in the synonymy of Sinothela by Haupt, 2003: 71; syn. n. Vinathela Ono, 2000; type species Heptathela cucphuongensis Ono, 1999: placed in the syno- nymy of Heptathela by Haupt, 2003: 71, 79. TYPE SPECIES: Liphistius kimurai Kishida, 1920. EMENDED DIAGNOSIS: Distinguished from Ryuthela Haupt, 1983 by males pos- sessing a conductor on sclerite III of the palpal organ and lacking an enlarged denticle (= contrategular spine) on its sclerite Il; females with 2, 3 or 4 receptacles or recepta- cular clusters on the anterior margin of the bursa copulatrix (if 2, then these more wi- dely separated from each other than in Ryuthela), or with 2 anteriorly and 2 dorsally on the bursa, or with 2 anteroventrally and 2 anterodorsally. Distinguished from Liphistius Schiôdte, 1849 by the presence of 7-8 spinnerets (posterior medians more or less dis- tinctly fused) and by the lack of clavate trichobothria on tarsi of legs and palps; males without retrolateral tibial apophysis on palp, palpal organ with large tegulum carrying two prominent apophyses, contrategulum and subtegulum without apophyses, para- cymbium without cumulus; females with leathery but unsclerotised walls of the bursa copulatrix devoid of thickened rim and vesicles, gland pores usually restricted to receptacles or receptacular clusters situated anteriorly, dorsally or only slightly antero- ventrally on the bursa. 602 P. J. SCHWENDINGER & H. ONO THE TWO SPECIES FROM SOUTHERN VIETNAM Heptathela australis (Ono, 2002) Figs 1-20, 55-58, 67 Songthela australis Ono, 2002a: 120-122, figs 1-8 (description of female). Heptathela australis (Ono, 2002).- Platnick, 2011 (transferred without explanation in an earlier version of that online catalogue; this generic placement is maintained here). MATERIAL EXAMINED: NSMT-Ar 9617 (1 male), IEBR, without registration numbers (1 male and 1 female), MHNG, without registration numbers (all other specimens), sample SV- 03/18; 9 males (matured early June 2004, 2.VIL.2004, 30.VIL.2004, 30.VI.2006, 10.V.2007, 24.V.2007, May 2007, 17.1V.2008, 27.1V.2008), 10 females and 1 juvenile; near Dambri Waterfall, about 18 km N of Bao Loc, 11°38’42”N, 107°44’37°E, 850 m; 1.-2.1X.2003; leg. P. J. Schwendinger. - MHNG, without registration number, sample SV-03/17; 1 male; Lam Dong Province, off the road from Da Hoa (— Da Huoai) to Bao Loc, about 13 km SW of Bao Loc, 11°27°20°N, 107°43°04”E, 690 m; 31.VIIL.2003; leg. P. J. Schwendinger. EMENDED DIAGNOSIS: Males of Æ. australis differ from those of 4. tonkinensis by a narrower cymbium with a longer and narrower elongate proventral-distal pro- jection; paracymbium much longer and basally much narrower, almost forming a right angle with the cymbium; an unpigmented and unsclerotised zone present distoventrally on cymbium; marginal tegular apophysis much longer and pointed, terminal tegular apophysis with its dorsal edge much less projecting and less dentate; contrategulum with two distal edges running parallel to each other: an outer sharp edge and an inner dentate edge; the sharp edge with a pronounced, beak-like extension prolaterally, the dentate edge not connected with the isolated denticles in the ventro-proximal part of the contrategulum; conductor shorter, situated close to the embolus, its apex without denticles; embolus longer. Females of Æ. australis are similar to those of H. hang- zhouensis, differing by larger posterior “bursae” (= receptacular clusters) (see Ono, 2002a: 120, referring to Song & Haupt, 1984: fig. 3c-d). Another distinction 1s found in the shape of the bursa copulatrix in dorsal and ventral view: longer and trapezoiïdal or triangular in Æ. australis, shorter and rectangular in 4. hangzhouensis. DESCRIPTION OF MALE (matured 10.V.2007): Colouration in alcohol (live spe- cimens distinctly darker): Ground colour of carapace light brown, with indistinct dark brown W-shaped pattern behind dark ocular mound and with dark brown band along anterior and lateral margins. Opisthosoma cream, mottled with light grey; tergites uni- formly light greyish brown; sternites light orange-brown; spinnerets cream. Chelicerae light brown distally, cream proximally. Legs and palps dorsally uniformly light reddish brown except for a dark red-brown distal zone of the cymbium (not including cymbial projection); no dark annulation present. Ventral side of body and limbs generally lighter than dorsal side. Total length 15.2. Carapace 6.4 long, 5.2 wide, set with a few short, peg-like hairs (with blunt tips) on margin (mostly anteriorly and posteriorly), behind ocular mound and on coxal elevations, only few longer pointed hairs running over ocular mound in a longitudinal row. Eye group 0.92 long, 1.05 wide. Eye sizes and interdis- tances: AME 0.04, ALE 0.59, PME 0.30, PLE 0.53; AME-AME 0.09, AME-ALE 1.14, PME-PME 0.04, PME-PLE 0.12, ALE-PLE 0.08. MOQ 0.43 long, front width 0.19, back width 0.46. Labium 0.4 long, 1.2 wide. Sternum 2.5 long, 2.2 wide (1.2 on ven- tral surface). Anterior margin of sternum distinctly elevated, thus a deep suture present between sternum and labium. Maxillae 2.2 long, 1.4 wide. Promargin of cheliceral HEPTATHELA FROM VIETNAM 603 groove With 10 small teeth on each chelicera. Paired tarsal claws with 4-5 teeth; un- paired claws without denticles. Measurements of limbs: palp 11.4 (3.4 + 1.9 + 3.9 + tn T38+r49 +25) ler I91(50+24+37+5.3 +27): 7 23300t.0.2 + 3.0); leg 1V26.5 (65 +2,74 5.0 + 8:6 + 3,7). “Tibial spurs” (sensu Platnick & Goloboff, 1985) absent on all legs. Opisthosoma 5.7 long, 3.6 wide; posterior margin of genital sternite (— sternite of second opisthosomal segment) widely rounded, with slightly protruding median part (Fig. 1). Palp (Figs 2-15) with a quite long and narrow cymbial projection (Fig. 2); distoventral zone of cymbium (below subtegulum) unpigmented and unsclerotised, formed by extension of unpigmented and unsclerotised inner (retrolateral distal) side of cymbium onto its ventral side (Fig. 3); paracymbium relatively long and narrow, with only indistinct proximal tubercle; cymbium and paracymbium almost at right angles (Fig. 3). Tegulum with two large apophyses: 1) marginal apophysis long, pointed, distad-directed, with a sharp edge running across it from prolateral to retro- lateral side (Figs 2-4); 2) terminal apophysis retrolaterad- and slightly proximad- directed, with an abruptly narrowed and slightly distad-bent apex (Fig. 3), dorsal side of apophysis only moderately extended and carrying a low, weakly dentate edge, dorsal and ventral sides of apophysis almost parallel to each other (Figs 3, 5, 10-14); edges of both apophyses not in contact with each other. Contrategulum with a few isolated denticles ventro-proximally and with two parallel distal edges, the outer one sharp, the inner one finely dentate, both uniformely pigmented (Fig. 3); sharp edge prolaterally abruptly truncate, with a beak-like extension pointing toward marginal apophysis of tegulum (Fig. 2). Conductor arising ventro-proximally on embolus (— bulbal sclerite IT), its proximal half fairly wide and fused with base of embolus, its distal half free, blade-like, with continuously narrowing sides and pointed apex (Figs 3-9). Embolus largely sclerotised, prolateral half strengthened by numerous longitudinal ribs (Figs 2-5), terminal fringe around opening of sperm duct (= spermophore) hyaline and finely serrate (Figs 3-4, 6, 8-9). DESCRIPTION OF FEMALE: See Ono (2002a: 120-122, figs 1-8). Anterior margin of sternum elevated as in male, thus also with a deep suture between sternum and labium. “Tibial spurs” present on legs I-IIT. Vulva as in Figs 16-19, 55-58. VARIATION: Range of measurements in males (n=10) and in newly collected females with clearly developed vulva (n=10; in parentheses): body length 12.7-17.6 (15.3-24.1), carapace length 5.4-7.1 (6.0-9.4), carapace width 4.2-6.2 (4.8-8.1). Variation in the overall shape of the vulvae of four new females is not strongly pro- nounced (Figs 16-18, 55). The largest female (only it) has a dark brown spot on the ventral wall of the bursa copulatrix, and several small sclerotised vesicles (as the ones on the receptacular clusters), penetrated by gland pores, sitting on the dorsal wall of the bursa copulatrix, in between the four receptacular clusters (Fig. 16). The latter charac- ter can be an atavism (see Discussion: evolution of female copulatory organs). In one female the posterior margin of the genital sternite is indented (probably from an injury sustained in an earlier instar; Fig. 20). For variation in the shape of the palpal organ, see Figs 6-9 (conductor), Figs 3, 10 (marginal tegular apophysis) and Figs 5, 10-14 (terminal tegular apophysis). One male has a proximally narrow paracymbium (Fig. 15). The number of isolated denticles ventro-proximally on the contrategulum varies from 2-7. 604 P. J. SCHWENDINGER & H. ONO 7 14-415 5-14 a FiGs 1-15 Heptathela australis (Ono, 2002), male (mature 10.V.2007) from the type locality (1-6), five other males from the type locality (7, 11; 9; 10; 12; 13), male from Dao Hoa - Bao Loc (8, 14- 15). (1) Posterior margin of genital sternite, ventral view. (2) Distal part of left palp, prolateral view. (3) Same, ventral view. (4) Same, retrolateral view. (5) Same, distal view. (6, 8-9) Embolus and conductor, proventral view. (7) Conductor, proventral view. (10) Marginal and terminal apo- HEPTATHELA FROM VIETNAM 605 FiGs 16-20 Heptathela australis (Ono, 2002). (16) Vulva of largest female, dorsal view; dorsal and lateral walls of genital atrium not shown. (17-18) Bursae copulatrices of two other females, dorsal view. (19) Schema of vulva, lateral view (the oviduct is actually anterodorsad- and not posterodorsad- directed). (20) Posterior margin of deformed genital sternite, ventral view. BC = bursa copula- trix; GA= genital atrium; Od = oviduct; RC = receptacular cluster. Scale lines 1.0 mm. RELATIONSHIPS: Despite pronounced differences in female copulatory organs, H. australis appears most closely related to N. nui sp. n. Their palps share distinct synapomorphies, and both species occur in the same mountain range, about 80 km apart. Their closest known relatives are Æ. tonkinensis and H. tomokunii from northern Vietnam, but more closely related species most likely await discovery in the mountains between Da Lat and Hanoi. The male of the geographically closer A. cucphuongensis (from south of Hanoi) is unknown and therefore the relationships of this species are not yet clear. physis of tegulum, ventral view. (11-14) Terminal apophysis of tegulum, ventral view. (15) Outline of paracymbium, ventral view. Co = conductor; CP = cymbial projection; CT = contra- tegulum; Cy = cymbium; DE = dentate distal edge of contrategulum; DT = dorsal extension of terminal apophysis of tegulum; E = embolus; MA = marginal apophysis of tegulum; PC = para- cymbium; PE = prolateral extension of sharp distal edge of contrategulum; SE = sharp distal edge of contrategulum; ST = subtegulum; T = tegulum; TA = terminal apophysis of tegulum:; Z = unpigmented distoventral zone of cymbium. Scale lines 1.0 mm. 606 P. J. SCHWENDINGER & H. ONO DISTRIBUTION: Known only from two localities near Bao Loc village in southern Vietnam. BIOLOGY: The mature male was discovered when scraping leaf litter from the floor of an evergreen broadleaf forest. He probably had left his burrow in search of females and was taking temporary shelter in the leaf litter during the day. AI other spiders were extracted from their burrows in sun-exposed earth banks (with little vegetation) on one side of a dirt road in an old secondary forest. None of the burrows inspected at the beginning of September contained mature males, and by then the mating period was probably at its end. AIT burrows were closed with a single trapdoor that had its hinge on the upper side of the entrance; the lower rim of the burrow entrance was slightly protruding. Trapdoors of females were up to 3.3 cm wide and 2.5 cm long; those of penultimate males 2.0-2.3 and 1.5-1.8 cm, respectively. No spiders were seen when their doors were carefully opened with forceps for inspection. They were resting in the depth of their burrows and could not be provoked to come to the entrance by inserting a flexible grass stem and disturbing them (which often works for extracting spiders of the genus Liphistius). AÏl these spiders had to be dug up. Large and medium-sized spiders (including adult males) react to disturbance by repeatedly raising their body above the ground while the leg tarsi remained on the ground (a behaviour called “tiptoeing”) and by spreading their chelicerae, as large Liphistius also do. Four egg cases (2.5-3.5 cm in diameter and 1.4-1.9 cm high) were built between the end of November and late December 2003. Three of them held clusters of 129-164 beige or light yellow eggs wrapped in a thin skin made of a coagulated vulval secretion and some fine silken threads (as present in all liphistiid egg cases examined, but less distinct in Liphistius), resting on a mesh of fine threads above the bottom of the inner chamber of the egg case. Ninety-nine third (or later?) instar juveniles were taken from the fourth egg case in mid-April 2004. Five of them were males that reached maturity in June 2006, May 2007 and April 2008, which is presumably earlier in the year than in nature (since the adult male was found at the end of August). Development from eggs to adult males in captivity thus took between one and a half years and three years and four months. Females which laid eggs between November and December moulted in the following March, May and June. Three of the large females, which were collected in the field and subsequently laid eggs in Geneva, each carried more than 50 ectoparasitic mites of different instars on their prosoma. These mites were usually sitting on the carapace, but when disturbed took shelter on the ventral side, between the coxae and the sternum, and between the prosoma and the opisthosoma. They pierced the carapace at numerous places and left clearly visible dark round scars that make the spiders look as if they had measles (Fig. 67). The carapace, a quite strongly sclerotised plate, is an unlikely place for sucking body fluids. Surprisingly, no scars or any other signs of damage are visible on the soft membranes between the sclerites of the prosoma or on the opisthosoma. HEPTATHELA FROM VIETNAM 607 Heptathela nui Sp. n. Figs 21-33, 68 HOLOTYPE: MHNG, without registration number, sample SV-03/21; male (matured 10.VI.2004); Vietnam, Lam Dong Province, Mt Penhatt near Quang Trung Reservoir, about 8 km SW of Da Lat, 11°52°37°N, 108°25’58"E, 1500 m; 6.1X.2003; leg. P. J. Schwendinger. PARATYPES: MHNG, without registration number, sample SV-03/21; 2 males (matured 3.V.2004, 10.VII.2004) and 3 females (“allotype” constructed egg case in XI1.2003 and moulted 5.1V.2004); collected together with the holotype. - NSMT-Ar 9618, 9619 (1 male, 1 female), IEBR, without registration numbers (1 male, 1 female), MHNG, without registration numbers (all other specimens), sample SV-03/22; 3 males (matured early June 2004, 15.VI.2004, 21.1V.2005) and 4 females;, Cam Ly Waterfall, western outskirts of Da Lat city, 11°56°37”"N, 108°25’14”E, 1450 m; 8.-9.1X.2003; leg. P. J. Schwendinger. ETYMOLOGY: “Nui”, a noun in apposition, is the Vietnamese word for “mountain”. DIAGNOSIS: Males of the new species can be distinguished from those of the closely related 7. australis by: anterior margin of sternum low; paracymbium stronger, with distinct proximal tubercle; conductor narrower, with more strongly bent apex; tegulum with larger marginal apophysis, terminal apophysis more continuously narrowing towards apex, its dorsal and ventral sides at an acute angle to each other; prolateral side of contrategulum with light-coloured central portion; sharp (outer) distal edge of contrategulum convex, broadly rounded, without beak-like extension; denticles on inner edge of contrategulum running down to ventro-proximal area in a continuous row; embolus shorter, its terminal fringe more coarsely serrate. Females of Æ. nui sp. n. differ from those of Æ. australis by: bursa copulatrix shorter, with more strongly inclined lateral margins; receptacular clusters smaller, with more or less distinct stalks, all four sitting on the anterior margin of the bursa copulatrix. DESCRIPTION: MALE (holotype). Colouration in alcohol (live specimens dis- tinctly darker): Carapace and legs light reddish brown; ocular mound dark; bulbal sclerites and distal part of cymbium red-brown, cymbial projection light brown. No annulation on legs or dark pattern on carapace and opisthosomal tergites. Ventral side of body and limbs generally lighter than dorsal side. Opisthosoma cream, mottled with light grey; tergites uniformly light greyish brown. Total length 14.4. Carapace 6.3 long, 5.0 wide, set with a few short, peg-like hairs (with blunt tips) on margin (mostly anteriorly and posteriorly), behind ocular mound and on coxal elevations; only few long pointed hairs running over ocular mound in a longitudinal row. Eye group 0.87 long, 0.99 wide anteriorly, 0.91 pos - teriorly. Eye sizes and interdistances: AME 0.04, ALE 0.60, PME 0.32, PLE 0.50; AME-AME 0.14, AME-ALE 0.07, PME-PME 0.05, PME-PLE 0.08, ALE-PLE 0.06. MOQ 0.43 long, front width 0.21, back width 0.52. Labium 0.5 long, 1.2 wide. Sternum 2.7 long, 2.2 wide (1.3 on ventral surface). Anterior margin of sternum low, thus only a shallow furrow present between sternum and labium. Maxillae 2.2 long, 1.4 wide. Promargin of cheliceral groove with 10 teeth on right chelicera, 12 on left. Paired tarsal claws with 3-5 teeth; unpaired claws without denticles. Measurements of lHimbs: PA MG ED EE + 2.3); les 19:76:66 +2,5.+ 40:50 7 2:6); les 19:33 ER ES 26622D; lee I 20.6(5.0+2.5+38762#31);'leg TV 25.9(6.6 + 2.7 + 4,9 + 8.2 + 3.5). “Tibial spurs” absent on all legs. Opisthosoma 5.6 long, 3.7 wide; posterior margin of genital sternite widely rounded, with an indistinct invagi- nation in the middle (Fig. 21). 608 P. J. SCHWENDINGER & H. ONO Palp (Figs 22-26) with paracymbium carrying a proximal tubercle (Fig. 23); cymbium and paracymbium almost at right angles to each other; distoventral zone of cymbium (below subtegulum) unpigmented and unsclerotised (Fig. 23). Tegulum with a very long, pointed, distad-directed marginal apophysis with a sharp edge (Figs 22-25), and with a retrolaterad- and slightly proximad-directed terminal apophysis continuously narrowing to a rounded, distad-bent apex (Figs 23, 25); dorsal side of terminal apophysis less extended than in F4. australis, carrying a weakly dentate edge (Fig. 25) not being in contact with edge on marginal apophysis. Contrategulum with a cream central portion on prolateral side (Fig. 22); two parallel distal edges present: the outer one sharp, the inner one finely dentate (Fig. 23); sharp edge convex, widely rounded, without beak-like extension (Fig. 22); row of denticles on inner edge running down to near ventro-proximal margin of contrategulum and there forming two rows of denticles together with a few irregularly arranged additional denticles situated more proximally (Fig. 23). Conductor situated ventro-proximally on embolus; proximal portion of conductor fairly wide and fused with embolic base; distal portion of conductor free, narrow, blade-like, with slightly hook-like apex (Figs 23-26). Embolus largely sclerotised, prolateral half strengthened by numerous longitudinal ribs (Figs 22, 25); terminal fringe around opening of sperm duct (= spermophore) hyaline, coarsely serrate, With fairly deep indentations (Figs 22-24, 26). FEMALE (‘“allotype”). Colouration similar to that of male but shightly lighter; palpal tarsi brown. Total length 21.6. Carapace 8.3 long, 6.3 wide. Eye group 0.97 long, 1.03 wide anteriorly, 1.00 posteriorly. Eve sizes and interdistances: AME 0.06, ALE 0.60, PME 0.35, PLE 0.50; AME-AME 0.19, AME-ALE 0.14, PME-PME 0.04, PME-PLE 0.06, ALE-PLE 0.06. MOQ 0.49 long, front width 0.30, back width 0.52. Labium 1.1 long, 2.0 wide. Sternum 3.6 long, 2.5 wide (1.9 on ventral surface), its anterior margin distinctly elevated, thus separated from labium by a deep suture (not so in males). Maxillae 2.8 long, 1.9 wide. Promargin of cheliceral groove with 11 teeth on each chelicera. Paired tarsal claws of legs with 2-4 teeth, unpaired claws without denticles; each palpal claw with 2 denticles. Measurements of limbs: palp 13.3 (4.5 + 2.5 + 3.0 +33): les115249+26%730730 107); le2114/6 442 6002 70 I lès IT 15.5 43 F27+27%%38+20) lee N219 (C1 0e 020022227720 “Tibial spurs” present on legs I-IIL. Opisthosoma 7.7 long, 6.4 wide. Posterior margin of genital sternite broadly rounded, with a slight median invagination. Vulva (Fig. 31). Bursa copulatrix short, plano-convex in dorsal and ventral view, its anterior margin Widely rounded and carrying four anteriad-directed recepta- cular clusters on indistinct short stalks, the median pair slightly larger than the lateral one. Genital atrium densely set with fine hairs on dorsal and ventral walls. VARIATION: Range of measurements in males (n=6) and in females with clearly developed vulvae (n=7; in parentheses) examined: Body length 13.2-16.5 (16.4-22.5), carapace length 5.2-6.5 (6.1-8.2), carapace width 4.2-5.3 (4.9-6.5). For variation in the shape of the vulva in three females, see Figs 31-33. In one female the receptacular clusters are raised on relatively long stalks (Fig. 33), in two other females the stalks are stout and rather indistinct (Figs 31-32). Variation in the shape of the palpal organ 1s less pronounced than in A. australis: the marginal tegular apophysis carries a subterminal HEPTATHELA FROM VIETNAM 609 21-24, 30 RÉ 25-29 Er PTS FiGs 21-30 Heptathela nui sp. n., male holotype (21-26) and three male paratypes from Cam Ly Waterfall (27-28; 29; 30). (21) Posterior margin of genital sternite, ventral view. (22) Distal part of left palp, prolateral view. (23) Same, ventral view. (24) Same, retrolateral view. (25) Same, distal view. (26) Embolus and conductor, proventral view. (27) Conductor, proventral view. (28) Marginal and terminal apophysis of tegulum, ventral view. (29) Marginal apophysis of tegulum, ventral view. (30) Outline of paracymbium, ventral view. PT = proximal tubercle of para- cymbium,; other abbreviations as in Figs 1-15. Scale lines 1.0 mm. 610 P. J. SCHWENDINGER & H. ONO FIGs 31-33 Heptathela nui sp. n., three female paratypes. (31) Vulva of “allotype”, dorsal view; dorsal and lateral walls of genital atrium not shown. (32-33) Bursae copulatrices of two other females, dor- sal view. Abbreviations as in Figs 16-20. Scale lines 1.0 mm. denticle in two males (Figs 25, 28), in others it 1s bare (Fig. 29); the proximal tubercle of the paracymbium is very distinct in two males (Figs 23, 30), less so in others; in some males the scattered denticles below the continuous row of denticles on the ventro-proximal side of the contrategulum are indistinct. In one female the posterior median spinnerets are fused only along their ental sides but the apices are still free, whereas in all other conspecific spiders examined these spinnerets are completely fused. REMARKS!: The paired leg claws of the “allotype” are inflated, especially on the posterior legs. This is not so in any of the other specimens of both species examined and thus it was probably caused by preservation. Similar rare cases of inflated claws were also observed in alcohol-preserved specimens of mygalomorph spiders. RELATIONSHIPS: Heptathela nui Sp. n. possesses a vulva similar to that of H. abca from northern Vietnam, and to F4. bristowei Gertsch, 1967, H. sinensis and H. schensiensis (Schenkel, 1953) from central China. However, strong similarity and several likely synapomorphies in the male palps of H. australis and H. nui sp. n. show that these species are the most closely related. See also Discussion - Systematics. DISTRIBUTION: Known only from two localities near Da Lat city in southern Vietnam. BIOLOGY: The spiders examined were collected from a steam bank in an ever- green hill forest, from the recess of an earth bank at the side of a path in a pine forest (both at Mt Penhatt), and from an earth bank in a park with pine trees (at Cam Ly Waterfall). Burrows were as those of Æ. australis, but at the Cam Ly site (not so on Mt Penhatt) they all had several grass stems attached to their entrances to enlarge the sensory area. “Twig-lining” is known from individual populations of several species of HEPTATHELA FROM VIETNAM 611 mygalomorph trapdoor spiders in Australia, North America and Japan (see Haupt, 1995 for a summary), and 1t was also reported for Ryuthela populations (Haupt, 1983: 286) and for some . abca at the type locality (Ono, 1999: 41, figs 13-14). Trapdoors of females were up to 2.6 cm wide and 2.0 cm long; those of penultimate males 1.6-2.1 and 1.1-1.6 cm, respectively. Large and medium-sized spiders react to disturbance by “tiptoeing”, spreading their chelicerae and raising their palps. One egg case (2.5 cm in diameter and 1.7 cm high) was built on 19.XI1.2004. It held 83 light yellow eggs suspended above the bottom of the inner chamber. The female with eggs moulted in early April of the following year; other females moulted in January, March, April, September, October and November, usually twice per year. Males (collected in early September 2003) became adult between May and July 2004 and in April 2005. The largest female (gravid; died before laying its eggs) had about 50 ecto- parasitic mites of the genus Ljunghia on its prosoma. These caused the same kind of scars on the carapace of the spider (Fig. 68) as described earlier in this paper for H. australis. SPECIES EXAMINED FOR COMPARISON Heptathela tonkinensis (Bristowe, 1933) Figs 34-38 Liphistius birmanicus Thorell, 1897.- Simon, 1909: 70-71 (misidentification, description of male). Liphistius tonkinensis Bristowe in Bristowe & Millot, 1933: 1022, text-fig. 4 (naming of species; illustration of palp of male holotype).- Bristowe, 1976: 4 (listing). Heptathela tonkinensis (Bristowe, 1933).- Haupt, 1983: 284-285, figs 8a-b, d, 13e (transfer; description and illustration of palp of male holotype); Platnick & Sedgwick, 1984: 3 (transfer). Vinatela tonkinensis (Bristowe, 1933).- Ono, 2000: 150 (transfer). Nanthela tonkinensis (Bristowe, 1933).- Haupt, 2003: 69, 71, figs S1a-b (transfer; illustration of palp of male holotype). Here transferred back to Heptathela. MATERIAL EXAMINED: MNHN 29170, AR-4104; male holotype; Vietnam, Tonkin, forêt de Khà là; no collecting date; leg. Blaise. EMENDED DIAGNOSIS: The previously published descriptions of the holotype do not mention some of the following characters: cymbial projection quite broad and dis- tinctly inclined from axis of cymbium (Fig. 34); distoventral zone of cymbium (below subtegulum) pigmented and sclerotised as normal; paracymbium short and exceptio- nally wide at base, at an acute angle to axis of cymbium (Fig. 35); contrategulum distally with a broad band of denticles (arranged in several irregular rows) running down to near ventro-proximal margin of contrategulum, no sharp distal edge present (Fig. 36); marginal apophysis of tegulum fairly short, compressed and arched, with a sharp distal edge running from prolateral to retrolateral side (Figs 34, 37-38); terminal apophysis of tegulum beak-like, dorsally with a large and broadly arched extension with a coarsely dentate edge (Figs 34, 36-38) not in contact with edge on marginal apophysis; conductor long (about as long as embolus), its distal part fairly distant from embolus (Figs 34, 36). “Tibial spurs” absent on all legs. Posterior margin of genital sternite widely and uniformly rounded, without protruding median part. Anterior margin of sternum not markedly elevated, suture to labium therefore shallow (as in male of H. nui sp. n.). 612 P. J. SCHWENDINGER & H. ONO 34 FiGs 34-38 Heptathela tonkinensis (Bristowe, 1933), male holotype. (34) Distal part of left palp, prolateral view. (35) Paracymbium of left palp, ventral view. (36) Left palpal organ, distal view. (37) Marginal and terminal apophysis of tegulum of left palp, ventral view. (38) Same, right palp. Abbreviations as in Figs 1-15. Scale lines 1.0 mm. REMARKS: The retrolateral view of the right palp of the holotype given by Bristowe (Bristowe & Millot, 1933: text-fig. 4) is detailed and correct, but it gives the false impression of a quite long and pointed marginal tegular apophysis, as present in H. australis and H. nui Sp. n. À comparison of these three species showed that all of them have a fairly sharp distal edge on their marginal tegular apophysis, and in pro- and retrolateral view this apophysis always looks pointed (see also Fig. 34). Ventral and dorsal views show that in 1. tonkinensis this apophysis is actually scale-like, fairly short and moderately arched (Figs 37-38; Haupt, 1983: fig 8b; Haupt, 2003: fig. 51B), whereas in #1. australis and H. nui sp. n. it is long, triangular and truly pointed (Figs 3, 5, 102325, 28-29); | RELATIONSHIPS: Among the hepthatheline species of which males are known, H. tonkinensis currently appears most closely related to 4. tomokunii; both are more distantly related to Æ. australis and H. nui sp. n. Heptathela cucphuongensis (known only from females), which also occurs in northern Vietnam (south of Hanoi), 1s geographically closer and possibly also more closely related to Æ. tonkinensis and H. tomokunii than to H. australis and H. nui sp. n. from the south of the country. DISTRIBUTION: Heptathela tonkinensis is known only from the holotype collected in a forest near Khà là (this spelling is given on the laeser-printed label in the tube with the specimen; in the original description the name of the locality is spelled “Kha-lé”), in the valley of the Song (= river) Luc Nam, northeast of Hanoï (Simon, 1909: 69, 71). This locality probably corresponds to the village of Kha Le, about 15 km east of the township of Luc Nam in Bac Giang Province, northern Vietnam. HEPTATHELA FROM VIETNAM 613 Heptathela tomokunii Ono, 1997 Figs 39-43 Heptathela tomokunii Ono, 1997a: 24-26, figs 1-8 (description of male and female). MATERIAL EXAMINED: NSMT-Ar 3396-3398; male holotype (matured 8.X.1995), female “allotype”, 1 female paratype; Vietnam, Vinh Phu Province, Tam Dao, 900 m:; 15./21.1X.1995; leg. H. Ono. REMARKS: Diagnosis and description were given by Ono (1997a). The following can be added: cymbial projection only moderately developed, quite wide (Fig. 39); paracymbium short and wide at base, at an acute angle to axis of cymbium (Ono, 1997a: fig. 2), similar to that of Æ. tonkinensis; subtegulum exceptionally long (Fig. 40); contrategulum with denticles on distal edge modified into a series of parallel ribs running down to 1-2 isolated ventral denticles near proximal margin (Figs 39-40); tegulum with quite low, narrowly arched marginal apophysis and with quite short terminal apophysis (Figs 39, 41) carrying a pronounced, strongly bulging dorsal extension with a coasely dentate edge (Fig. 39; Ono, 1997a: fig. 4) not in contact with edge on marginal apophysis; genital atrium with some hairs on ventral side near posterior and lateral margins (Fig. 42); bursa copulatrix short, anterior margin widely arched; all receptacles simple and smooth [Fig. 43; also confirmed for the “allotype”, though the corresponding illustration (Ono, 1997a: fig. 6) indicates indistinctly cauli- flower-shaped receptacles], not developed as caulifower-shaped receptacular clusters as in other Heptathelinae examined; larger lateral pair of receptacles sessile, slightly displaced onto dorsal wall of bursa copulatrix; smaller median pair of receptacles stalked [on common socket in paratype (Fig. 43), on individual stalks in “allotype” (Ono, 1997a: fig. 6)] and slightly displaced onto ventral wall (Fig. 43). “Tibial spurs” present on legs I-IIT of female, on last exuvia of male and also on mature male (with a few “‘spurs” reduced). RELATIONSHIPS: Due to similarities in the palpal organ of males (moderately developed cymbial projection; short, wide paracymbium at an acute angle to the axis of the cymbium; long subtegulum; low, arched marginal tegular apophysis; pronoun- ced dorsal extension of terminal tegular apophysis), H. tomokunii appears to be more closely related to Æ. tonkinensis (female unknown) than to Æ. australis and H. nui sp. n. More or less pronounced displacements of the lateral pair of receptacles onto the dorsal wall of the bursa copulatrix in Æ. tomokunii, H. australis and some Ryuthela species are considered homoplasic (see Discussion). DISTRIBUTION: This species is only known from the type locality, the Tam Dao hill station, about 85 km northwest of Hanoi, in northern Vietnam. Heptathela kikuyai Ono, 1998 Figs 44-45 Heptathela kikuyai Ono, 1998: 16-19, figs 11-16 (description of male and female). MATERIAL EXAMINED: NSMT-Ar 8718; 1 male; Japan, Kyushu Island, Oita Prefecture, Hita-shi; 15.X.2006; leg. T. Irie. —- NSMT-Ar 3545; 2 females; Kyushu Island, Miyazaki Prefecture, Nishi-ushuki-gun, Takachiho-cho; 8.-14.1X.1996; leg. H. Ono. DISTRIBUTION: Known from several localities in the northwestern part of Kyushu Island, Japan. 614 P. J. SCHWENDINGER & H. ONO RE ne FIGs 39-43 Heptathela tomokunii Ono, 1997, male holotype (39-41) and female paratype (42-43). (39) Distal part of left palp, prolateral view. (40) Left palpal organ, ventral view. (41) Marginal and terminal apophysis of tegulum of left palp, ventral view. (42) Vulva, dorsal view: dorsal and lateral walls of genital atrium not shown. (43) Bursa copulatrix, dorsal view. R= receptacle; other abbreviations as in Figs 1-20. Scale lines 1.0 mm. Heptathela kimurai (Kishida, 1920) Figs 46-47 Liphistius kimurai Kishida, 1920: 362, fig. 1 (description of male and female); type species of Heptathela. For a complete synonymy, see Platnick (2011). MATERIAL EXAMINED: NSMT-Ar 3549; 1 male; Japan, Kyushu Island, Kagoshima-shi:; 7.X1.1970; leg. K. Tanaka. — NSMT-Ar 8717; 1 female; Kyushu Island, Kagoshima-shi, Shiroyama; 25.X1.1997; leg. H. Ono. DISTRIBUTION: Known from several localities in the southern part of Kyushu Island, Japan. HEPTATHELA FROM VIETNAM 615 F1iGs 44-50 Heptathela kikuyai Ono, 1998, male (44-45); Heptathela kimurai (Kishida, 1920), male (46-47); Ryuthela owadai Ono, 1997, male holotype (48-50). (44, 46, 49) Left cymbium, prolateral view. (45, 47, 50) Left palpal organ (45 and 47 slightly expanded), ventral view. (48) Same, retrolateral view. CS = contrategular spine; other abbreviations as in Figs 1-15. Scale lines 1.0 mm. 616 P. J. SCHWENDINGER & H. ONO Ryuthela owadai Ono, 1997 Figs 48-50 Ryuthela owadai Ono, 1997b: 155-157, figs 15-18 (description of male).- Ono, 2001: 151-153, figs 1-3 (description of female). MATERIAL EXAMINED: NSMT-Ar 3459; male holotype; Japan, Okinawa Prefecture, Tokashiki Island, Aharen, about 100 m; 11.X.1990; leg. M. Owada. —- NSMT-Ar 8717; 1 female; Okinawa, Tokashiki Island, Tokashiku, 30 m; 17.1.2001; leg. H. Ono. DISTRIBUTION: Known only from the Tokashikijima Island, Ryukyu Islands, Japan. Ryuthela nishihirai (Haupt, 1979) Heptathela nishihirai Haupt, 1979: 356-362, 365, figs 6-13 (description of male and female); type species of Ryuthela. For a complete synonymy, see Platnick (2011). MATERIAL EXAMINED: NSMT-Ar 422-423; |] male (matured X.1977) and 1 female syntypes; Japan, Shuri, Okinawa, Ryukyu Islands; 15.11.1976; leg. M. Nishihira & J. Haupt. — MHNG (coll. S. Heimer), without registration number; 1 female; Japan, Ryukyu Islands, without further data. DISTRIBUTION: Known from several localities on Okinawa Island, Japan. Liphistiinae Thorell, 1869 Liphistius ornatus Ono & Schwendinger, 1990 Figs 51-54 Liphistius ornatus Ono & Schwendinger, 1990: 166-170, figs 1-8 (description of male and female). MATERIAL EXAMINED: MHNG, without registration number; male holotype, 1 male paratype, 4 female paratypes; Thailand, Chanthaburi Province, Khao Soi Dao Wildlife Sanctuary, 300-400 m; 9.V.1987; leg. P. Schwendinger. DISTRIBUTION: Known only from the type locality in the southern part of eastern Thailand. Liphistius thaleri Schwendinger, 2009 Figs 59-64 Liphistius thaleri Schwendinger, 2009: 1255-1260, figs 1-12 (description of male and female). MATERIAL EXAMINED: MHNG, without registration number; male holotype, 5 male paratypes, 8 female paratypes; Thailand, Trang Province, Libong Island, 30 m; 20.VII.2005; leg. P. Schwendinger. DISTRIBUTION: Known only from Libong Island in southern Thailand. DISCUSSION TAXONOMIC CHARACTERS AND TERMINOLOGY: Different and partly contra - dictory terminology (for a summary see Table 1) for genital structures of liphistiid spiders has been used in taxonomic publications by various authors. Conductor: Haupt (1979: 358, 360-361, figs 8-10 and subsequent papers of the same author) used the term “conductor” for a distad-directed spine-like structure on the palpal organ of Ryuthela males. This spine arises on the ventral side of the contra - tegulum and is actually a strongly elongate denticle, the last (proximal-most) one in a series (continuous or interrupted) of denticles that run down from the distal edge of the contrategulum in all heptatheline palps examined (denticles indistinctly developed in HEPTATHELA FROM VIETNAM 617 SE(DET) / Ve RS | 5 OS En ? AATTAEES. : te FIGs 51-54 Liphistius ornatus Ono & Schwendinger, 1990, male holotype (51-52) and female paratype (53-54). (51) Distal part of left palp, ventral view. (52) Same, retrolateral view. (53) Vulva, dorsal view; dorsal and lateral walls of bursa capulatrix and genital atrium not shown. (54) Same, ventral view. CDO = central dorsal opening; Co = conductor; CT = contrategulum; Cy = cymbium; DEC = distal edge of “contrategulum”; DET = dorsodistal edge of “tegulum”; DPE — dentate proximal edge of ‘“contrategulum”; DT — dorsal extension of terminal apohysis of tegulum; E = embolus; GA = genital atrium; MA = marginal apophysis of tegulum; P = medium- sized pore (leading to ampulliform vesicle); PC = paracymbium; PeP = paraembolic plate; PP] — poreplate; RC = receptacular cluster; SE = sharp distal edge of contrategulum; ST = sub- tegulum; T = tegulum; TiA = tibial apophsis; V = ampulliform vesicle. Scale lines 1.0 mm. Modified from Ono & Schwendinger, 1990: figs 1-4, with permission from the Bulletin of the National Science Museum (Tokyo). H. kimurai, Fig. 47; modified to ribs in Æ. tomokunii, Figs 39-40). In À. owadai the three proximal denticles are elongate to various degrees (Figs 48, 50; Ono, 1997b: figs 16-18). Thus the “conductor” of Ryuthela is not fused with the contrategulum as suggested by Haupt (1983: 286; 2003: 71), it is an original part of the contrategulum and as such also of the bulbal sclerite II sensu Kraus (1978: 237, figs 2-4). In other heptatheline genera the conductor is a structure of various shapes that arises from the ventral base of the embolus (or embolus complex) and thus belongs to bulbal sclerite II. Consequently the “conductors” of both groups are not homologous and therefore should not both be called the same. In his descriptions of various Ryuthela species Ono correctly notes: “‘contrategulum with a basal spine; conductor not developed” (e.g., Ono, 1997b). The elongate basal denticle of the contrategulum, in the following called contrategular spine, is unique within the Liphistiidae and a clear synapomorphy for species in Ryuthela. Raven (1985: 15, 16) also did not recognize the different origins of the Ryuthela and Heptathela ‘“‘conductors” but just their different shapes, and gave this as one of the reasons to synonymise Ryuthela with Heptathela (Ryuthela was later removed from synonymy by Haupt, 2003: 71). 618 P. J. SCHWENDINGER & H. ONO The conductor of Heptathela 1s on the same bulbal sclerite (IT) and essentially in the same position as the “‘paraembolic plate” (also called ‘‘scale-like plate of the ventral embolus edge”, “proximal edge of embolus” and “Basalkante des Embolus”) present in many species of Liphistius (especially in those of the #rang-group; Figs 51-52). These sclerites differ only in shape and in distance to the embolus proper: in Liphistius the paraembolic plate 1s wide, short or long, fairly remote from the embolus and slightly inclined from it, whereas in non-Ryuthela heptathelines the conductor is wide or narrow, long, usually lies much closer to the embolus and runs more or less parallel to its surface. The relative position of the conductor in 4. tonkinensis (type species of “Nanthela”; Figs 34, 36) is intermediate between that of Liphistius and Heptathela. Differences in shape [paraembolic plate scale-like (Fig. 51) or only deve- loped as a low wide edge in Liphistius (Schwendinger, 1998: fig. 1 A-D, I); conductor leaf-shaped in Heptathela from Japan (Haupt, 2003: fig. 52F-K, M-P, fig. 62G-L), proximally broad and usually carrying three more or less pronounced, acutely pointed tips in “Sinothela” (Haupt, 2003: fig. 52L, fig. 62D-F), blade-like, spiniform and smooth, or spiniform and apically dentate in “Nanthela” (Haupt, 2003: fig. 62C)] do not exclude that these structures are homologous. This was stated by Raven (1985: 15) and vigorously refuted by Haupt (2003: 92); we here support Raven’s view. The terms “‘conductor” and ‘“contrategulum” were confused in Ono’s (1998) taxonomic treatment of the males of five Heptathela species from Kyushu Island (Japan). What is correctly referred to as “conductor” in the descriptive text (except on page 21), 1s incorrectly called “contrategulum” in the legends to the corresponding figures. Tegulum: Tegulum and contrategulum are the two more or less distinctly outlined parts of the ring-shaped bulbal sclerite IT of the palpal organ in Liphistiidae. In the Heptathelinae the tegulum extends over the dorsal and retrodorsal side of the bulb and carries two apophyses: 1) Dorsally lies the distad-directed marginal apophysis over which runs a sharp distal edge from the prolateral to the retrolateral side. This apo- physis 1s long and pointed in Æ. australis and H. nui sp. n. (Figs 2-5, 10, 22-25, 28-29), short and arched in the other heptatheline species examined (Figs 34, 37-38, 39, 45, 47-48). 2) Retrolaterally lies the retrolaterad-directed terminal apophysis over which runs a more or less coarsely dentate (least developed in À. owadai) dorsal edge perpendicular to the distal edge on the marginal apophysis. The terminal tegular apo- physis is distinctly developed in “Nanthela” and Heptathela (Figs 3, 5, 10-14, 23, 25, 28, 34, 36-39, 41, 45, 47), indistinctly so in Ryuthela (Fig. 48). In Æ. australis and H. nui Sp. n. the dorsal side of the terminal tegular apophysis is only moderately extended and slightly arched, with the dorsal dentate edge only little elevated (Figs 5, 25), in the other Heptathelinae examined it is strongly protruding into a widely arched extension (Figs 34, 36, 39, 47-48; not visible in Fig. 45). In #1. kikuyai and H. kimurai there is a coarsely dentate third edge on the tegulum (= the “first edge” of Song & Haupt, 1984: 444) which starts at the point where the edges of the marginal and terminal tegular apo- physes meet, and from there it runs down the prodorsal side to the base of the tegulum. This edge is not produced into an apophysis as on the other two tegular edges. This condition appears to be characteristic for H. kimurai (the type species of Heptathela) and its closest relatives. In the other Heptathelinae the edges of the marginal and terminal apophyses do not meet prodorsally and no third edge is present. HEPTATHELA FROM VIETNAM 619 In Liphistius the tegulum 1s less obvious, because it seemingly does not possess the same apophyses as the tegulum of the Heptathelinae. The so-called ‘“tegular process” of some Liphistius Species arises from the opposite (= ventral to proventral) side of bulbal sclerite II and thus belongs to the contrategulum. The contrategular process 1s probably apomorphic for Liphistius, as is also the subtegular apophysis (both present only in some species); both apophyses are absent in all Heptathelinae. What was previously called “tegulum” in Liphistius therefore does not correspond to the te- gulum in the Heptathelinae. There 1s, however, a sclerotised area on the retrodorsal side of bulbal sclerite IT in Liphistius that is connected to the larger part (— the contra- tegulum) of bulbal sclerite II by a narrow sclerotised area and that carries a sharp distal edge in most species and a dentate marginal or submarginal proximal edge in all species (F1g. 52, Schwendinger, 1996: figs 1, 12, 21, 31, 39, 46, 55, 65, 74, cf. Figs 4, 24). These two edges, though being more two-dimensional, resemble the marginal and terminal tegular apophyses of Heptathelinae and they lie at the same position on the palpal organ. Haupt (1983: figs 1c, 3-4) called the sclerite of the Liphistius palp which carries these two edges the “contrategulum”, and that appellation was followed in pa- pers on Liphistius by Ono and by Schwendinger. We assume that Haupt regarded this sclerite as the “‘contrategulum” because of a misinterpretation of his own illustration (see Haupt, 1983: fig. 1c) that shows a distinctly expanded male palp of L. malayanus Abraham, 1923 in which the so-called “contrategulum (K)” was rotated to the ventral side where the contrategulum 1s situated in the Heptathelinae (see Haupt, 1983: fig. 1b). In his illustrations of non-expanded Liphistius palps (see Haupt, 1983: figs 3b-c, 4a-b), however, the so-called “contrategulum (K)” lies on the other side, retro- dorsally, where the tegulum of the Heptathelinae 1s situated. Correspondingly the latter figures show the so-called “tegulum (T})” proventrally, where the Heptathelinae have the contrategulum. Haupt has obviously confused these terms in both taxa. The tegulum of Liphistius (hitherto incorrectly called “contrategulum”) is much shorter and narrower than that of the Heptathelinae, and it is confined to the retrodorsal side of bulbal sclerite II. The dentate proximal or subproximal edge of the tegulum of Liphistius is presumably homologous with the dentale edge on the dorsal side (pro- truded into a dorsal extension in some species) of the terminal tegular apophysis of the Heptathelinae, and the more or less distinctly developed distal edge of the Liphistius tegulum is presumably homologous with the marginal apophysis of the Heptathelinae (Fig. 52 cf. Figs 4, 24). A fairly simple tegulum (more reduced than in Æeptathela but less than in Liphistius) is also present in Ryuthela owadai (Fig. 48). We consider the small tegula of Liphistius and Ryuthela as homoplasic reductions, which implies that the palpal organs of Liphistius and (to a lesser degree) Ryuthela are more derived than those of Heptathela. Contrategulum: Haupt (1983: 277) introduced the term “contrategulum” for the prolateral to ventral part of bulbal sclerite II that opposes the tegulum. In the Heptathelinae tegulum and contrategulum are more or less distinctly separated by a wide membranous zone (haematodocha) retroventrally, thus forming a broken ring. Prodorsally, however, both parts are widely fused and it is difficult to tell where the contrategulum ends and the tegulum begins. An indicator is the sharp distal edge of the contrategulum portion which more or less abruptly ends prolaterally in Æ. australis 620 P. J. SCHWENDINGER & H. ONO (Fig. 2), A. nui sp. n. (Fig. 22), H. kikuyai and H. kimurai, Wereas in H. tonkinensis (Figs 34, 36), H. tomokunii (Fig. 39) and À. owadai (Fig. 48) it fades out between the marginal tegular apophysis and the embolus. More proximally at that place contrate- gulum and tegulum are usually fused with each other without a transition, though in Hepthatela males from Japan examined a small invagination in the proximal margin of bulbal sclerite IT presumably marks the border between both parts. In Ryuthela this pat- tern is modified and the distinction between contrategulum and tegulum is more visible on the dorsal side of the palpal organ: the contrategulum is much wider than in other heptathelines (as wide as in Liphistius) and extends distally past the tegulum to the retrolateral side in an almost complete circle; a deep, almost horizontal furrow divides both sclerites dorsally (Fig. 48), and only on the ventral side are tegulum and contra- tegulum completely fused (Fig. 50). In À. nishihirai tegulum and contrategulum appear to be completely separated (see Haupt, 1979: 360, fig. 9). In Liphistius the contra - tegulum is much more extensive than the tegulum, almost completely embracing the middle portion of the palpal organ (as also in Ryuthela, but there the tegulum is much wider than in Liphistius and overlaps the contrategulum). Retrolaterally the large contrategulum and the much smaller tegulum are connected by sclerotised bridges (the more dorsal one always narrower than the more ventral one) and form a closed ring (Fig. 52). What was previously called “tegular process”, ‘“distal edge of tegulum” and “dorsal edge of tegulum” in various publications on Liphistius by Haupt and by Schwendinger, are in fact parts of the contrategulum. In Heptathelinae the contrategulum usually carries on 1ts distal edge (which is divided into a sharp outer edge and a dentate inner edge in A. australis and H. nui Sp. n.) a row of denticles which are more or less distinctly connected to a few (in some species isolated) denticles ventro-proximally. These denticles were subject to evolu - tionary modifications of taxonomic significance. In several species they have multiplied and become more than one row deep: in Æ. nui Sp. n. two rows proximally (Fig. 23), in À. owadai three rows proximally (Fig. 50), in Æ. kikuyai several rows proximally (Fig. 45) and in A. tonkinensis several irregular rows on most of its distal edge (Fig. 36); in A. tomokunii (Figs 39-40) and seemingly also in F4. hongkong Song & Wu, 1997 (Haupt, 2003: fig. S1D) they have transformed into a series of parallel edges or ribs. Such ribs are also present in some Liphistius species where they are probably homoplasic (see Schwendinger, 1995: figs 22-23, 26-27; Schwendinger, 1996: figs 12, 15-17; Schwendinger, 1998; fig. 5b, e-f). In Ryuthela spp. the basalmost of these denticles (in À. owadai to a lesser degree also two adjacent ones; Figs 48, 50) has become strongly elongate and spine-like. This. modified contrategular denticle of Ryuthela is clearly not homologous to the conductor (or paraembolic plate) of other Liphistiidae (see discussion of conductor), but it presents a clear synapomorphy for species in Ryuthela. Cymbial projection: Haupt (2003: 69, 94, character 23 in table 11 and fig. 61) considers the elongate proventral cymbial lobe as “autapomorphic” (Haupt, 2003: 94; it should read “synapomorphic”) and distinctive for males of species in the genus “Nanthela”. However, this modification is also present in two Heptathela species from Japan examined: F4. kikuyai (Fig. 44) and H. kimurai (Fig. 46; the type species), and it is also discernible in Song & Haupt’s (1984: fig. 1c) illustration of the male palp of HEPTATHELA FROM VIETNAM 621 H. schensiensis (previously in “Songthela”) and in Haupt’s (1983: fig. 10a-d) illus- trations of the male palp of Ryuthela nishihirai and R. ishigakiensis Haupt, 1983. In R. owadai the cymbial projection is only weakly developed (Fig. 49). In some Liphistius species (e.g., L. bristowei Platnick & Sedgwick, 1984 and Z. batuensis Abraham, 1923; in MHNG examined), on the other hand, it is the prodorsal cymbial lobe (instead of the proventral one) that is moderately elongate. Thus it appears that one of the cymbial lobes (the proventral one in Heptathelinae, the prodorsal one in Liphistius) has become elongated to various degrees independently in different phylo- genetic lineages. The taxonomic value of this character at the generic level has thus been overestimated. Vulva: HEPTATHELINAE. The vulva of heptatheline spiders is composed of two main parts (see Fig. 19): 1) The posterior part is a long and wide entrance area (= the genital atrium) furnished with numerous hairs on the dorsal and ventral walls in H. australis (Fig. 16), H. nui sp. n. (Fig. 31) and Æ. tomokunii (Fig. 42), but only on the dorsal wall in A. kikuyai, H. kimurai, R. owadai and R. nishihirai. The genital atrium 1s overlapped by a more or less tongue-shaped projection of the posteromedian margin of the genital sternite. 2) The anterior part of the vulva is a blind-ending pouch (= the bursa copulatrix) in the form of a flattened bell (see Ono, 2002a: fig. 3), plano- convex or trapezoidal in dorsal and ventral view, with walls consisting of a light yellow, tough, leathery but not sclerotised cuticle (Figs 16-19, 31-33, 42-43, 55). As already stated by Forster (1980: 280), there is no sclerotised structure in the vulva of heptatheline spiders that can be called a genital plate (an incorrect term used by several authors; see Table 1]. Usually no pores are present in the walls of the bursa copulatrix (Figs 55-56), but the largest H. australis female has tiny pores on the small vesicles that lie scattered between the receptacular clusters (Fig. 16). This could be an atavism (see Evolution of female copulatory organs). From the dorsal side of the vulva, where both main parts meet at an obtuse angle, a short and wide, dorsoventrally depressed membranous collar (— the uterus externus) leads dorsad to the mesodermal part of the oviduct (= the uterus internus) and further to the ovary (Fig. 19). In the illustrations by Haupt (1983: fig. 2b) the entrance to the uterus externus 1s erroneously shown at the anterior end of the bursa copulatrix, which 1s in fact a cul-de-sac. The anterior margin of the bursa copulatrix carries between one and four sessile or stalked, mostly cauli- flower-shaped protuberances (= the receptacular clusters) composed of numerous more or less strongly sclerotised vesicles (— the receptacles). The composite structure of the receptacular clusters in heptathelines (for Ryuthela nishihirai) was first recognized by Haupt (1979: 365). The walls of these vesicles are perforated by tiny micropores through which surrounding glands empty. The proximal, cuticular portions of the gland ducts that are sticking out of the micropores like fine hairs (Figs 56-58; SEM-micro- graphs taken from exuviae) clearly show that these are indeed gland pores. In the two females of H. tomokunii examined these anterior protuberances on the bursa copulatrix are simple in shape and have a fairly smooth surface (though also perforated by micro- pores; Figs 42-43) and may be called receptacles. This indicates a trend towards simplification, which is also seen in the receptacular clusters of some Liphistius species that have become smooth and digitiform (Sedgwick & Schwendinger, 1990; fig. 6; Schwendinger, 1996: figs 18-20, 52-54). In A. australis and other species previously 622 P. J. SCHWENDINGER & H. ONO i LU FIGS 55-58 Heptathela australis (Ono, 2002), SEM-micrographs of genitalia from the exuvia of one female, dorsal views. (55) Entire bursa copulatrix with receptacular clusters. (56) Border between dorsal wall of bursa copulatrix (without pores) and lower left receptacular cluster (with gland pores). (57) Lower left receptacular cluster. (58) Detail of the same showing gland pores and cuticular parts of gland ducts. BC = bursa copulatrix; GD = cuticular (proximal) part of gland duct; GP = gland pore (micropore); RC = receptacular cluster. Scale lines: 14 um (58), 31 um (56), 56 um (57), 236 um (55). placed in “Songthela”’ by Ono (2000), two of these receptacular clusters (the lateral ones) have become displaced onto the dorsal wall of the bursa copulatrix and are directed dorsad instead of anteriad (Figs 16-19, 55). Illustrations in the original description of H. goulouensis Yin, 2001 show a very different type of vulva (without a bursa copulatrix but instead with a pair of bifid “curved, tube-like stalks”, each carrying a median and a lateral receptacle or recep- tacular cluster; Yin, 2001: figs 5-6, 8-9) which is similar to the vulvae of certain mygalomorph spiders. If correct, this would be most unusual for liphistiid spiders. À HEPTATHELA FROM VIETNAM 623 re-examination of these specimens may show that the true boundaries of the bursa copulatrix were not recognized and that the “stalks” are actually zones of different pig- mentation, or a mass of sperm inside the bursa copulatrix. Similar ‘“‘stalks” and a short bursa copulatrix were 1llustrated for the holotype of Æ. sapana (Ono, 2010: fig. 16). The so called “didymous phenomenon” displayed by the female paratype of H. gou- louensis (Yin, 2001: figs 8-9) is most likely based on a spider that was preserved prior to moulting, which has the new cuticle of the vulva already formed underneath the old cuticle. The number of receptacles or receptacular clusters and their position on the bursa copulatrix 1s variable and this has been used to recognize groups of species which were given generic rank. Ryuthela is characterized by only two anterior receptacular clusters situated close to each other (Haupt, 2003: 71, fig. 53), but a rudimentary pair of additional (lateral) receptacular clusters 1s still present in the female “allotype” of R. owadai (see Ono, 2001: figs 2-3). This is not so in the corresponding female para- type examined. Non-Ryuthela Heptathelinae usually have four more or less distinctly developed receptacles or receptacular clusters, but distinction from Ryuthela is not clear-cut. Lateral receptacular clusters have become more or less fused with median ones in several Heptathela species (see Ono, 1998: figs 9-10, 15-16, 17-20, 31-32; Haupt, 2003: fig. 54), resulting in only two anteromedian receptacular clusters which are generally more widely separated from each other than in Ryuthela. The holotypes Of H. hunanensis Song & Haupt, 1984 and of 4. cucphuongensis have the median pair fused and the lateral pair still isolated, resulting in a vulva with three anteromedian receptacular clusters (Song & Haupt, 1984: fig. 3e; Ono, 1999: fig. 8). Ono (2000) used this intermediate character state to establish the genus “Vinathela” (misspelled “Viathela” and placed in the synonymy of Heptathela by Haupt, 2003: 71, 79). Fusion of receptacular clusters has gone even further: the illustrated vulva of R. iheyana Ono, 2002 has a single, wide and short anteromedian receptacular cluster (Ono, 2002b: figs 2-3); the holotype and one female paratype of À. sasakii Ono, 1997 have a single, long and narrow anteromedian receptacular custer (Ono, 1997b: figs 5-8). We consider all these forms as intermediate stages of reduction and fusion of the four anterior recepta- cular clusters that are plesiomorphically present in heptatheline spiders. Such modi - fications probably have evolved independently in different phylogenetic lineages and should be treated with caution when establishing relationships between species. Two receptacular clusters displaced to the dorsal side of the bursa copulatrix (Figs 16-19, 55) is another kind of modification in heptatheline vulvae. This character state was used by Ono (2000) to establish the genus “Songthela”, but as H. australis (described in “Songthela”) has it and the obviously closely related 7. nui Sp. n. not, it can no longer be used as distinctive on the generic level. AIT the more so because se- veral transitional steps exist: the lateral receptacular clusters of Æ. tomokunii (see Figs 42-43; Ono, 1997a: fig. 6), H. sinensis, H. schensiensis (see Song & Haupt, 1984: fig. 3a-b, f) [the latter two placed in “Abcathela” by Ono, 2000, in “Sinothela” by Haupt, 2003 and then in “Songthela” by Platnick, in an earlier version of his 2011 online catalogue] and of H. yunnanensis Song & Haupt, 1984 (see Song & Haupt, 1984: fig. 3f) are also slightly displaced onto the dorsal wall of the bursa copulatrix, but not as far back as in A. australis (Figs 16-18, 55) and as in . hangzhouensis (see Song & 624 P. J. SCHWENDINGER & H. ONO Haupt, 1984: fig. 3c-d) (both species previously in “Songthela”). The median pair of receptacles (reduced receptacular clusters) of 4. tomokunii (Figs 42-43; in addition to the lateral pair which was slightly displaced onto the dorsal wall) and of some Ryuthela species (see e.g., Ono, 1997b: figs 5-14, 19-22) is even sitting slightly below the anterior margin of the bursa copulatrix, on its ventral wall (thus on the same side as the receptacular cluster of Liphistius). LIPHISTIIDAE. The heptatheline vulva 1s largely different from the vulva of Liphistius which carries no anterior receptacular clusters. In that genus the dorsal wall of the bursa copulatrix is a thin transparent membrane that never carries any modifi- cations apart from a pigmented spot opposing the macropore in the ventral wall in a few large females. (In Liphistius the entrance to the uterus externus 1s also situated at the border between atrium and bursa copulatrix, not at the anterior end of the bursa copulatrix as shown in Haupt, 1983: fig. 2a). The ventral wall of the bursa copulatrix in Liphistius is strongly sclerotised and includes a true genital plate (= the poreplate) perforated dorsally by medium-sized pores that lead inside small ampulliform vesicles on the ventral side (Figs 53-54, 59-60). The appellation “gland pores” used in a paper by Schwendinger (1996) for these vesicles is incorrect; the gland pores (= micropores) are actually on the ouside of the vesicles (ventrally on the poreplate), as clearly visible in SEM-micrographs by Kraus (1978: fig. 8; see also Fig. 62). Kraus (1978: 242-243, figs 6-8) called these vesicles “circular structures [or circular areas] ... interpreted as a pore system for glands ... [which] must not be confused with the receptacula”. Kraus (1978: fig. 8) first showed SEM-micrographs of micropores in Liphistius and recognized the function of the ampulliform vesicles, but the corresponding lettering 1s misleading [in fig. 8 the ventral surface of a vesicle is labelled as “gland pore”, in fig. 6 the opening (= medium-sized pore) of a vesicle on the dorsal side of the pore- plate]. Sperm that enters into the ampulliform vesicles from the dorsal side of the poreplate is thus supplied with (or flushed out by) a secretion of the receptacular glands from the other side. Thus the vesicles on the ventral side of the Liphistius poreplate are sperm receptacles. On the dorsal side of the Liphistius poreplate a fairly large central opening (or macropore; CDO in Figs 53, 59) leads to an unpaired, strongly sclerotised receptacle or (in most species) receptacular cluster (Figs 54, 60-61; Haupt, 2003: 68 called this an “unpaired ventral sac”) on the ventral side of the poreplate. The central receptacular cluster 1s perforated by gland pores, and it varies in shape from simple and digitiform (in that case called a receptacle, see Sedgwick & Schwendinger, 1990; fg. 6; Schwendinger, 1996: figs 18-20, 52-54) to complex and cauliflower- or popcorn- shaped (Kraus, 1978: figs 5-8; Platnick & Sedgwick, 1984: figs 69, 80, 87). In the heptatheline vulva no receptacle or receptacular cluster is found at this position (although shown there in Haupt, 1983: fig. 2b), but a slight displacement of the median pair onto the ventral side of the bursal copulatrix can be seen in A. tomokunii (Figs FIGs 59-64 Liphistius thaleri Schwendinger, 2009, SEM-micrographs of vulvae from the exuviae of three female paratypes. (59) Ventral wall of opened bursa copulatrix and genital atrium, dorsal view. (60) Same, ventral view. (61) Receptacular cluster. (62) Ampulliform vesicle. (63) Anterolateral HEPTATHELA FROM VIETNAM 625 _=+ © D 4 ‘ protuberance on rim of poreplate. (64) Posterolateral protuberance on rim of poreplate. ALP — anterolateral protuberance on rim of poreplate; AMP = anteromedian protuberance on rim of poreplate; CDO = central dorsal opening (macropore) to receptacular cluster on other side; GA = genital atrium; GD = cuticular (proximal) part of gland duct; GP = gland pore (micropore); P — medium-sized pore (leading to ampulliform vesicle); PLP — posterolateral protuberance on rim of poreplate; PPI = poreplate; RC = receptacular cluster; V = ampulliform vesicle. Scale lines: 14 um (64), 16 um (62), 32 um (63), 35 um (61), 195 um (59), 205 um (60). 626 P. J. SCHWENDINGER & H. ONO 42-43) and in several Ryuthela species (Ono, 1997b: figs 9-14, 19-22). In À. sasakii this pair has become fused, it possesses a common unpaired, slightly ventrally situa- ted, fairly large opening (macropore; see Ono, 1997b: figs 5, 7), and thus resembles the receptacular cluster in Liphistius. Several Liphistius species possess 1-3 pairs of ventrad-directed protuberances on the thickened anterior and lateral margins of the poreplate (Fig. 60). These appear to be synapomorphic for members of superspecies D of the #rang-group (co-occurring with a synapomorphic subtegular apophysis in males of these species, see Schwendinger, 2009; subtegular apophyses in the bristowei-group are homoplasic), but also occur in distantly related species in other species groups (e.g., Platnick & Sedgwick, 1984: figs 13, 21, 78; Schwendinger, 1990: figs 2-4, 37-39, 43-45, 47-49, 53-56). These protuberances are hollow, in contact with the bursa copulatrix through medium-sized pores on the dorsal side of the poreplate, and their walls are perforated with micropores through which gland ducts enter (Figs 60, 63-64). Therefore the ven- tral marginal protuberances are functional sperm receptacles. Function and position of these protuberances suggest that they are homologous with the receptacular clusters of the heptatheline vulva, but considering that the poreplate in Liphistius itself is an apo- mophic structure, that is quite unlikely. We regard these protuberances as outgrowths on the thickened margin of the poreplate, which have developed several times inde- pendently in Liphistius. They are useful for species distinction, and their number and position on the poreplate allow recognizing relationships between species. Micropores are not only found in the elevated parts of the ventral side of the poreplate, but also in the flat parts between them (Figs 60, 62). Most of the pores there are grouped and sitting in shallow depressions, with membranous gland duct bases sticking out of them, and these groups of sessile micropores are in contact with medium-sized pores on the dorsal side of the poreplate. Thus essentially the whole of the poreplate serves as a sperm receptacle. Evolution of female copulatory organs: The fact that two very different types of vulvae exist within the Liphistiidae raises the questions: how did they evolve and which one is more derived than the other? Five hypotheses were put forward in an attempt to explain the evolution of female copulatory organs in Liphistiidae and in Araneae as a whole: 1) Platnick & Gertsch (1976: 6), and subsequently Platnick (1977: 13), pos- tulated that in ancestral spiders the palps were pressed simultaneously into the female genital tract during copulation, and that the tips of embolus and conductor of each palpal organ left imprints in the wall of the vulva which developed into receptacles. A vulva with four receptacles, as found in most Heptathelinae and in some basal Mygalomorphae and basal Araneomorphae, 1s thus plesiomorphic for the Araneae. 2) An alternative hypothesis, put forward by Forster (1980: 277, fig. 12) and further elaborated by Forster, Platnick & Gray (1987: 94-98), presents an archetypical spider vulva with only a large and simple bursa copulatrix supported by a secretory gland system that empties through pores in the anterior wall of the bursa. According to this hypothesis, numerous receptacles (vesicles) developed later by invagination (better called exsacculation) of the bursal wall where the gland pores are situated. In this view the Liphistius vulva is relatively primitive because it still has numerous indi- HEPTATHELA FROM VIETNAM 627 TABLE 1. Different terminology used for homologous genital characters in three groups of liphistiid spiders by various authors in 14 selected publications. Forster (19801); Haupt (19792, 19833, 2003+); Kraus (1978); Ono (1997b6); Ono (19987); Ono & Nishikawa (19898); Schwendinger (1990°, 199610, 200911), Song & Haupt (198412), Yin (200113), present paperl4. Abbreviations (in parentheses) correspond to those given in Figs 1-64. Terms between inverted commas are considered as inappropriate or incorrect. Heptathela (see Figs 1-47, 55-58, 67-68) Ryuthela (see Figs 48-50) Liphistius (see Figs 51-54, 59-64) conductor (Co)2: 3 4 7, 12, 14 “contrategulum” (CT)7 ventro-proximal denticles on contrateguluml# contrategulum (CT)? # 7.8, 12, 14 tegulum (T)?: # 7, 8, 12, 14 terminal apophysis of teguluml?; third tegular edgel?; dentate edge of dorsal extension of terminal apophysis of tegulum (DT)l4 marginal Re of tegulum (MA)!2: 14; second tegular edgel genital plate: 12. 15; Genital- platte; dorsal and ventral walls of bursa copulatrixl4 receptaculal: 3 4 12, 13; receptacles (R)14; receptacula organl>; receptacular clusters (RC)1#; spermathecae?: “conductor”2: à: 4; spine on contrategulumt; contrategular spine (CS)l4 Medianapophyse?; contrategulum (CT)6: 14 tegulum (T)2: 3: 4 6, 14 dentate edge of dorsal extension of terminal apophysis of tegulum (DT)!4 marginal apophysis of tegulum (MA) Genitalplatte?; Atrialplatte}: dorsal and ventral walls of bursa copulatrix 14 receptacula?: 3: 4; receptacular clusters (RC)14; spermathecae® para-embolic plate (PP)10. 11, 14: “posterior” edge of embolus”: scale-like proximal embolus edge”; broad lamella at base of embolus*: ‘; Chitinplatte auf Basalkante des Embolus}: conductorl+ “tegulum” (T3: 4 9. 10, 11; contrategulum (CT)!4 “contrategulum” (CT): 4 10, 11: tegulum (T)!4 dentate proximal edge of “contrategulum” (DPE)!0; “ventral” edge of “contra- tegulum”1l; dentate edge of dorsal extension of terminal apophysis of tegulum (DT)!4 distal edge of “contra- tegulum” (DEC)10; “dorsal” edge of “contrategulum”1l: distal edge of teguluml#; marginal apophysis of tegulum (MA)14 Genitalplatte?: Atrialplatte: receptaculum#; chitinous plate’; inner plate”; internal plate”; poreplate (PP1)!: 14 receptaculum®?. 10, 14: receptacular system; central receptacular cluster (RC)°: 10. 11, 14: bseudo-receptaculum!; unpaired ventral sac4 vidual receptacles [plus an autapomorphic unpaired “pseudo-receptaculum” (= the central receptacular cluster)] embedded in the poreplate, whereas the heptatheline vulva has developed further away from the ancestral form through a bilateral aggrega - tion of receptacles into receptacular clusters and through a reduction of the poreplate (Forster, Platnick & Gray, 1987: 98). 628 P. J. SCHWENDINGER & H. ONO 3) A similar interpretation was given by Kraus (1978: 235, 249), who considers the Liphistius-type vulva (with an unpaired central receptaclular cluster) as the most primitive within the Araneae and postulated that vulvae with paired receptacles (and also those with unpaired ones) derived from it. 4) Haupt took elements of the previous hypotheses to explain the evolution of female copulatory organs in spiders. He postulated an ancestral vulva with a large bursa copulatrix and irregularly arranged gland pores (Haupt, 1983: 288), as did Forster (1980), but referred to Kraus (1978) who proposed a different hypothesis (see above). Localised receptacles were assume to have formed later in co-evolution with the split tips of the embolus (Haupt, 1983: 288). In his cladogram of mesothelid syste- matics and in the corresponding character evaluation Haupt (1990: 136, fig. 1) follows Kraus (1978) and regards the vulva of Liphistius as plesiomorphic and that of Heptathelinae as apomorphic. Later, however, he noted that “it cannot be decided with certainty whether unpaired or paired receptacula in mesothelid spiders have to be considered as apomorphic for the group” (Haupt, 2003: 93). 5) Raven (1985: 15-16) regarded the vulva of Ryuthela (with a single pair of anteromedian receptacles) as plesiomorphic and assumed that the vulvae of Heptathela and Liphistius derived from it. In view of the evolutionary tendencies for fusion and posteriad-displacement (to the dorsal and ventral walls) of four anterior receptacles or receptacular clusters in different lineages of heptatheline spiders, we believe that the same has also happened during the evolution of Liphistius species. We thus favour a combination of hypotheses l and 2 and consider the Liphistius vulva as derived from a proto-Heptathela vulva. That presumably resembled the vulvae of Æ. nui sp. n. (Figs 31-33) and of species from China and Vietnam previously placed in the genus “Abcathela” by Ono (2000), but had a much larger bursa copulatrix carrying four receptacular clusters on its anterior margin and possibly also scattered micro-receptacles on the walls of the bursa copu- latrix more posteriorly. Such isolated micro-receptacles (vesicles penetrated by gland pores) are still present in the anterior part of the dorsal surface of the bursa copulatrix of the largest #. australis female examined (Fig. 16). This possibly is an atavism. The pronounced differences in the vulvae of Liphistius and Heptathela indicate that the separation of both genera goes back much further in time than the separation of Heptathela and Ryuthela. The vulva of the latter 1s essentially a Heptathela vulva with a strongly reduced bursa copulatrix, with the receptacular clusters of each side fused to one another resulting in a single pair. This interpretation is in line with the hypothesis of Schwendinger (2009: 1265- 1266) that the Mesothelae originated in Euramerica before its integration into Pangaea, and from Euramerica spread eastward onto terranes that successively accreted to eastern Laurasia. In such a scenario, and knowing the very limited powers of dispersal of extant mesothelid spiders, one would rather expect to find the more basal extant species in southern and eastern China than further south, in Myanmar, Thailand, peninsular Malaysia and Sumatra. The latter lands mostly lie on the Sibumasu terrane which accreted to Laurasia later than the Chinese terranes and thus was colonized by Laurasian species later. This zoogeographic hypothesis and its implications for liphis- tud phylogeny will be falsified as soon as an early (Palaeozoic or Mesozoic) meso- thelid fossil is discovered on land of Gondwanan origin. HEPTATHELA FROM VIETNAM 629 “Tibial spurs”: The function of these paired structures distally on leg tibiae I-IIT (Platnick & Goloboff, 1985: figs 1-2; Haupt, 2003: fig. 16B-D), which must not be confused with coupling spurs ventrally or prolaterally on tibia I or tibia IT in males of many mygalomorph spiders, is not yet understood. Platnick & Goloboff (1985) consider them as part of proprioreceptive sense organs in which the “tibial spurs” press against smooth oval patches at the bases of metatarsi I-IIT (Platnick & Goloboff, 1985: figs 3-4; Haupt, 2003: fig. 16B, D). Haupt (2003: 22, 95, fig. 16), however, assumes that the “tibial spurs”, situated close to lyriform organs, are more likely used to monitor hemolymph pressure and strains in the cuticle, because they are allegedly not in contact with the metatarsal patches (“the bristle has no contact to the circular area of thinner cuticle”). This statement 1s incorrect. On alcohol-preserved specimens with stretched legs, or on a dry, stub-mounted and sputter-coated leg as shown by Haupt (2003: 16D), these structures are usually not in contact with each other, but they are so on live spiders sitting in ambush with slightly bent legs at the burrow entrance. Whatever the function of this sensory system is, its phylogenetic significance as a clear apomorphy of the Liphistiidae 1s undisputed. We found it to be present in all juvenile (also exuvia of penultimate males) and female liphistiids ever examined by both of us. “Tibial spurs” (but not always the light smooth paired patches on metatarsi I-IIT) are usually absent in mature males. In the male holotype of Æ. tomokunii and in the male paratype of L. ornatus (Ono & Schwendinger, 1990: 169-170) some “tibial spurs” are present but probably no longer functional. SYSTEMATICS: Discoveries of new species or of the missing sex of described species occasionally challenge the concepts of established supraspecific taxa. This 1s also the case in the two Heptathela species from southern Vietnam. The female of H. australis corresponds perfectly to Ono’s (2000: 150, fig. 4D) concept of “Songthela”, where this species was originally placed. Its newly discovered male, however, possesses the two characters considered by Haupt (2003: 69) as diagnostic for “Nanthela” (a cymbial projection present, conductor slender and much of its base fused to the embolus), and it also has a character considered in the same paper, on the same page, as diagnostic for Heptathela [paracymbium almost as long as cymbium (though relatively shorter in relation to the cymbium of Heptathela from Japan because that usually has no pronounced cymbial projection)]. The same type of male palp is present in the closely related Æ. nui sp. n., the female of which, however, possesses a vulva as in species previously placed in “Abcathela” (see Ono, 2000: 149-150, fig. 4C). This suggests that three of these four nominal genera (“Abcathela”, Heptathela, “Nanthela”, “Songthela”) do not reflect phylogenetic relationships. We rule out that Æ. australis and H. nui sp. n. are only distantly related. Geographical proximity and strong overall resemblance of the male palps show that these species are the most closely related. They share four likely synapomorphies in their male palps: 1) a very pronounced cymbial projection (more than in other heptathelines); 2) a long and pointed marginal tegular apophysis; 3) two parallel distal edges (one sharp, the other dentate) on the contrategulum; 4) an unpigmented and unsclerotised distoventral zone on the cymbium. Consequently these two species cannot be kept in different genera, and at the moment it appears most suitable to place them in the genus Feptathela. The generic concept of “Songthela” is firmly rejected because the species included clearly do not represent a monophyletic group. Its single diagnostic character 630 P. J. SCHWENDINGER & H. ONO (the dorsally displaced lateral receptacular clusters on the bursa copulatrix) is also present in species outside the genus, but it is not shared by the closely related species pair “Songthela” australis and Heptathela nui sp. n. The original concept of this genus does anyway no longer correspond to the currently included species apart from the type species, S. hangzhouensis (see Platnick, 2011). “Songthela” australis and “Song- thela ‘'cipingensis Wang, 1989 were transferred to Heptathela in an earlier version of Platnick’s (2011) online catalogue, without giving an explanation. Three other species [H. heyangensis (Zhu & Wang, 1984), H. schensiensis and H. sinensis, all with a different type of vulva than in the type species] were included in “Songthela” by the synonymisation of “Abcathela” with “Sinothela” and then by the synonymisation of “Sinothela” With “Songthela”. The generic concept of “Nanthela” is also rejected because it is not based on any clear apomorphies. One of the two diagnostic characters, the cymbial projection is also present in species outside “Nanthela”, e.g., in H. kikuyai (Fig. 44), H. kimurai (Fig. 46) and “Sinothela” schensiensis (see Song & Haupt, 1984: fig. 1c-d). The other diagnostic character of “Nanthela”, the slender conductor, is possibly plesiomorphic (Haupt, 1990: 137, fig. 1; Haupt, 2003: fig. 61, table 11). Instead of retaining this poorly defined genus and transferring species from the well-established genus Heptathela to it (“Songthela” australis, H. nui sp. n., “Nanthela” tonkinensis, H. to - mokunii and presumably also other species form Vietnam form a distinct clade), we here place “Nanthela” in the synonymy of Heptathela. We want to point out that the systematics of the Heptathelinae (except for the fairly well-defined and monophyletic group of species currently placed in the genus Ryuthela), as proposed by Ono (2000), Haupt (2003), Platnick (2011) and summarized in Table 2, are in dire need of a comprehensive revision based on genitalic characters of both sexes, and ideally also on molecular data. Until this is done (and the missing sexes are found), transferring or (in most cases) returning all non-Ryuthela Heptathelinae to Heptathela solves at least the problem of having closely related species in different genera. This solution is not ideal. Ryuthela appears to be an offshoot from within Heptathela, leaving the latter paraphyletic. At the moment no other monophyletic groups of species, which are sufficiently distinct to warrant generic rank, can be recognized within Heptathela. However, a more comprehensive study including the missing sexes (and new species yet to be discovered) may show that distinct clades do exist in Heptathela as defined here. If so, the 34 nominal taxa here included in Heptathela could be split into several genera again. In that case plenty of generic names Ww1ll be available. SUPPLEMENTARY OBSERVATIONS: Venom glands and moulting position: While Haupt’s (2003) monograph is a very useful and comprehensive compilation of the knowledge of the Mesothelae of that time, his characterisation of the group includes two misinterpretations: 1) Haupt stated that mesothelid spiders do not possess venom glands (“Mesothelae, however, lack such venom glands. ... This can only mean that venom glands are an apomorphic character of Opisthothelae”; Haupt, 2003: 6), despite the detailed description and illustration of such a gland in Liphistius desultor Schiôdte, 1849 by Millot (Bristowe & Millot, 1933: 1046, pl. 3, figs 1-2). 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In liphistiid spiders the venom probably does not play an important role during prey capture. Upon se1zing prey, the spiders almost immediately start crushing it with their strong and un- usually mobile chelicera, which can be spread at a right angle to each other. Liphistius bites to one of us (PJS) had no recognizable toxic effects. 2) He also stated that “the moulting specimen turns round and lies on its back for the moulting process .…..” (Haupt, 2003: 49-50, 95), in the same way as theraphosid spiders do, and he concludes “As this turning for moulting does not occur in other Megoperculata ... it seems to be an autapomorphic habit characteristic of Araneae” (Haupt, 2003: 50). This 1s not the case. Haupt was presumably generalizing from a single observation captured in a photo (Haupt, 2003: fig. 35D) which shows the final moult of a L. trang Platnick & Sedgwick, 1984 male lying on its back. Looking at the exuvia in this photo, however, it becomes clear that the spider actually moulted in a venter-down position. When pulling the anterior limbs out of the old cuticle, the posterior legs obviously remained stuck in it. As the spider struggled to get free, both legs IV and one leg II (still soft and flexible) bent backwards and the spider fell on its back. The damage to the discarded exuvia seen in another photo (Haupt, 2003: fig. 35E), with the fourth pair of legs missing, clearly show that this was not a normal moult and that the spider’s fourth pair of legs probably got permanently stuck in the exuvia. An examination of this specimen in Haupt’s collection should confirm our interpretation. We have had the chance to observe dozens of moults of liphistid spiders in captivity, and we have collected hundreds of exuviae in the field and in captivity. In all cases the exuviae were in a venter-down position, with the leg claws and palpal claws gripping the substrate (Figs 65-66). In the interior of their burrows (in nature lHiphistiids always moult inside them) there is not enough space for the spiders to arch up and bend backward during moulting. They thus normally moult in the same position as uropygids and amblypygids. Ectoparasitic mites: The same kind of mites, as were collected from the two Heptathela species in southern Vietnam (Figs 67-68 show bite marks), were also found on Liphistius at several localities in Laos, Thailand and peninsular Malaysia. At one locality in eastern Thailand the majority of liphistiid spiders collected were carrying mites. They stay on their hosts permanently and pierce the hard carapaces and cheli- cerae but not the soft membranes. These mites were identified as belonging to the laelapid genus Ljunghia and are currently being studied by I. Juvara-Bals & B. Halliday. Ljunghia includes obligate ectoparasites which naturally occur on primitive spiders in SE-Asia and Australia (Domrow, 1975), and which were also found on a Central American theraphosid spider in captivity (Moraza ef al., 2009). So far only one Ljunghia species associated with liphistiid spiders is known, Ljunghia bristowi (Finnegan, 1933) living on Liphistius malayanus in peninsular Malaysia (Finnegan, 1933). Further species, including the one living on A. australis and H. nui sp. n., Will be described soon. It appears worth pointing out that all Heptathela specimens from southern Vietnam that carried such mites were reproducing females that either laid eggs (3 X. australis) or were gravid (1 Æ. nui sp. n.). No mites were seen on juveniles or mature HEPTATHELA FROM VIETNAM FiGs 65-66 Liphistius sumatranus Thorell, 1890, moulting of an immature male in captivity. (65) Spider partly emerged from the old cuticle. (66) Spider fully emerged. Note that the animal does not lie on its back. 634 P. J. SCHWENDINGER & H. ONO FIiGs 67-68 Dorsal view of prosomata of two females showing bite marks caused by ectoparasitic Ljunghia mites. (67) Heptathela australis (Ono, 2002); chelicerae spread due to preservation. (68) H. nui Sp. n., paratype; chelicerae in normal position. HEPTATHELA FROM VIETNAM 635 males. This may just be a coincidence, because in Liphistius such mites were also found on females that did not reproduce in captivity, and on immature and mature males (unpublished observation). ACKNOWLEDGEMENTS The MHNG supported the first author’s collecting trip to southern Vietnam. The Japan Society for the Promotion of Science (JSPS No. 21540487) supported the second autor’s study. Christine Rollard (MNHN) kindly sent us the holotype of Heptathela tonkinensis for examination. Ilinca Juvara-Bals (Cologny, Switzerland) identified the mites; Bruce Halliday (CSIRO, Canberra, Australia) confirmed her identification and provided additional information. Rainer Foelix (Aarau, Switzerland) pointed out the cuticular bases of the receptacular glands and commented on the ultrastructure of spi- der genitalia. Norman Platnick (American Museum of Natural History, New York, USA) kindly reviewed the manuscript and pointed out hypotheses and publications that we have overlooked. Yoshimi Watanabe (NSMT) produced most of the ink drawings, Christina Lehmann-Graber (MHNG) completed them and added other ones, André Piuz (MHNG) took the SEM micrographs, Florence Marteau and Lionel Monod (both MHNG) arranged all illustrations into plates. John Hollier (MHNG) checked the English text and suggested improvements. The Bulletin of the National Science Museum (Tokyo), renamed the Bulletin of the National Museum of Nature and Science, allowed us to reproduce figures 51-54 from Ono & Schwendinger (1990). REFERENCES ABRAHAM, H. C. 1923. A new spider of the genus Liphistius from the Malay Peninsula and some observations on its habits. Proceedings of the Zoological Society of London 1923: 769-774. BISHOP, S. C. & CROSBY, C. R. 1932. A new species of the spider family Liphistiidae from China. Peking Natural History Bulletin 6(3): 5-7. BRISTOWE, W. S. 1976. A contribution to the knowledge of liphistiid spiders. Journal of Zoology 178: 1-6. BRISTOWE, W. S. & MILLOT, J. 1933. The liphistiid spiders [by W. S. Bristowe], with an appendix on their internal anatomy [by J. Millot]. Proceedings of the Zoological Society of London 1932(4): 1015-1057, plates 1-6. CHEN, Z., ZHANG, Z. & ZHU, C. 1981. À new species of genus Heptathela. Journal of the Hangzhou University 8(3): 305-308. DOMROW, R. 1975. 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Liphistius thaleri, a new mesothelid spider species from southern Thailand (Araneae, Liphistiidae). Contributions to Natural History 12: 1253-1268. SEDGWICK, W. C. & SCHWENDINGER, P. J. 1990. On a new cave-dwelling Liphistius from Thailand (Araneae: Liphistiidae). Bulletin of the British arachnological Society 8(4): 109-112. SIMON, E. 1909. Etude sur les arachnides du Tonkin (1°'€ partie). Bulletin scientifique de la France et de la Belgique 42: 69-147. SONG, D. & HAUPT, J. 1984. Comparative morphology and phylogeny of liphistiomorph spiders (Araneae: Mesothelae). 2. Revision of new Chinese heptathelid species. Verhandlungen des Naturwissenschafilichen Vereins in Hamburg (new series) 27: 443-451. SONG, D. & WU, K. 1997. On a new species of the genus Heptathela (Araneae: Liphistiidae) from Hong Kong, China. Chinese Journal of Zoology 32(3): 1-3. THORELL, T. 1869. On European spiders. Part I. Review of the European genera of spiders, pre- ceded by some observations on zoological nomenclature. Nova acta regiae Societatis scientiarum Upsaliensis (series 3) 7: 1-108. THORELL, T. 1897. Viaggio di Leonardo Fea in Birmania e regione vicine. LXXIII. Secondo saggio sui ragni birmani. Annali del Museo Civico di Storia Naturale di Genova (series 2) 17: 161-267. YIN, C.-M. 2001. A new species of the genus Heptathela and its variant type from China. Acta zootaxonomica sinica 26(3): 297-300. tr ecrire : i » td ré hr 00 ol PAPA trade maté sait UT qomsedté cal M tt : LA ct RS | = sr anA MMS SPORE I > à Cine S'anteg NS it crC LIRE ie À dt BLX hs tPieweec 6 à vf: en dés, M “10e NDS — p D in tæ # at "t sé 14 she von À © A = 1 FA, Her ARIAUE ir2T ba \s re : TT - ire «+ mt dont de DOME LEUR, 2. re” L id En? REVUE SUISSE DE ZOOLOGIE 118 (4): 639-649; décembre 2011 A new genus and species of Spilomelinae (Lepidoptera, Pyralidae) from the Galapagos Islands, Ecuador Bernard LANDRY! , Lazaro ROQUE-ALBELO? & James E. HAY DEN? l Muséum d'histoire naturelle, C.P. 6434, 1211 Geneva 6, Switzerland. E-mail: bernard.landry(@ville-ge.ch 2 Ecologia Environment, 1025 Wellington St., West Perth, W.A. Australia and Curtin Institute for Biodiversity and Climate, Curtin University, PO Box U1987, Perth, WA 6845, Australia. E-mail: lazaro.roque-albelo(@ecologia.com.au 3 Florida State Collection of Arthropods, FDACS, Division of Plant Industry, P.O. Box 147100, Gainesville, Florida, U.S.A., 32614. E-mail: james.hayden(@freshfromflorida.com A new genus and species of Spilomelinae (Lepidoptera, Pyralidae) from the Galapagos Islands, Ecuador. - Cheverella galapagensis Landry, gen. n. and sp. n. is described as an endemic of the Galapagos Islands, Ecuador. Based on a combination of two apomorphies (reduced uncus and presence of setose pads on anterodorsal extensions of the male valva medially) this taxon is possibly related to Choristostigma Warren, now placed in the Hydriris group of Munroe (1995), but Cheverella lacks the setose projection at base of the valve’s costa present in Choristostigma. The female has a clearly circumscribed corpus bursae with a short, spine-like signum. Cheverella also shares a few characters with members of the Siga group. The larva is a borer in stems of the endemic Zournefortia pubescens Hook. f. (Boraginaceae). INTRODUCTION During his first expedition to the Galapagos in 1989 at the onset of his inven- tory of the Lepidoptera of the archipelago, B. Landry (BL) came across the nicely patterned pyraloid treated here, and began to investigate its generic affinities. The double praecinctorium pointed to a Spilomelinae, and along the years since 1989 the species was shown to several colleagues (James Hayden, Eugene Munroe, Matthias Nuss, Michael Shaffer, Alma Solis), who concluded that it was undescribed and that it did not belong to any described genus, while various proposals were made with regard to phylogenetic affinity to other genera. In 1983, Eugene Munroe had, in fact, examined a specimen preserved in the Museum of Comparative Zoology, Cambridge, Massachusetts, and added a label saying ‘New Pyraustine”, a group which at the time included the Spilomelinae. Here this new taxon is finally described. Its recognition will allow for an evaluation of its conservation status and to plan future studies on its ecology. Manuscript accepted 28.07.2011 640 B. LANDRY ET AL. With more than 3700 species described, the Spilomelinae is one of the two largest subfamilies of the Pyraloidea, a group including either one (Pyralidae) or two (Crambidae and Pyralidae) families, depending on authors. It is included in the mono- phyletic group of pyraloid subfamilies here referred to as Crambiformes (the Crambidae, or Crambinina), the other group of subfamilies being known as Pyralidae s. str., or Pyralinina, or Pyraliformes. Munroe (1995) treated the subfamily as a tribe of Pyraustinae and recorded 1437 species in the Neotropical Region. The Spilomelinae are believed to be polyphyletic (Minet, 1982; Solis & Maes, 2003), but this has never been tested. The moths are characterized by the absence of chaetosemata, a bilobed praecinctorium, a projecting fornix tympani (tympanic frame), a pointed spinula, the absence of a gnathos, and the absence of a rhomboidal signum on the bursa copulatrix of the female genitalia (Minet, 1982; Nuss ef al., 2011). Other diagnostic characters are the loss of the subcostal hamus (frenulum hook) on the male forewing and the male valva with a clasper in some other conformation than in Pyraustinae (Munroe, 1976: 8). When Cheverella is compared with the matrix and characters used by Solis & Maes (2003) in their phylogenetic analysis of the crambiforme subfamilies, the genus scores as a representative of the Spilomelinae for all 17 characters used, although the shape of the median ridge of the tegumen 1s here complete from base to apex, not u-shaped at base. In the Galapagos Islands, there are 42 species of Spilomelinae currently known, of which nine are undescribed and under investigation (BL, unpublished). MATERIAL AND METHODS Specimens were collected mostly by BL at light during five expeditions to all of the major islands of the Galapagos in 1989, 1992, 2002, 2004, and 2005. The 1989 material was deposited at the Canadian National Collection of Insects, Ottawa, Canada (CNC) while most of the rest is deposited in the Muséum d’histoire naturelle, Geneva (MHNG). Many valuable specimens, including the single reared specimen came from co-author L. Roque-Albelo while he worked at the Charles Darwin Research Station, on Santa Cruz Island, Galapagos (CDRS). Additional material came from the California Academy of Sciences, San Francisco, California, USA (CAS), the Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA (MCZ), and the Natural History Museum, London, U.K. (BMNH). The terminology of the tympanal organs follows Landry (1995). That of geni- talia follows Solis & Maes (2003). The manner of giving the label data of the holotype and paratypes is presented in Landry (2006) as are the methods used for specimen collecting. DESCRIPTIONS Cheverella Landry gen. n. Figs 1-8 Type species: Cheverella galapagensis sp. n. Gender: feminine. DIAGNOSIS: Cheverella can be separated from the other genera of Spilomelinae by two apomorphies of the male genitalia, i.e. the reduced uncus and the presence of NEW GENUS AND SPECIES OF SPILOMELINAE FROM THE GALAPAGOS ISLANDS 64] FIGS 1-2 Cheverella galapagensis Landry, sp. n. (1) Holotype (CNC). (2) Darker coloured female paratype (MHNG). setose pads on anterodorsal extensions of the valva mediobasally, and the clearly circumscribed corpus bursae with a short, spine-like signum in the female. Choristo - stigma Warren, a mainly North American genus with nine species, also has a reduced unçcus and setose pads on a transtilla or extensions of the valva as in Cheverella (C. ele - gantalis Warren and C. plumbosignalis (Fernald) examined), but its valva is more narrow and it has a short, setose projection at base of costa. The female genitalia of Choristostigma (C. plumbosignalis (Fernald) examined) differ more strongly in that the wide ductus bursae has a large colliculum, the papillae anales are elongate rather than triangular, and the signum 1s very large, oval, and with multiple spines that are 4-5 X longer than their basal width and irregularly distributed. DESCRIPTION: Head (Fig. 3) with antennae filiform, slightly thinner in female, reaching almost 2/3 of length of forewing, with dense, short ciliation ventrally, with single short seta arising from scale-coated dorsal edge near middle of first 12 flagel- lomere approximately, with last flagellomere terminating into distinct sensillum stylo- conicum; with ocelli; without chaetosemata; maxillary palpus very short; labial palpus curved upward at about half right angle, reaching slightly above eye. Forewing rather narrow, 2.4 X longer than largest width. Frenulum simple in male, with 2 acanthae in female. Retinaculum a short bunch of scales below cubital stem; male without frenulum hook. Wing venation (n=1) (Fig. 4): Forewing Sc, RI, and R2 free, latter from before upper angle of cell; R3 and R4 stalked for most of length, stemming from upper angle of cell; RS, M1-3, and CuAÏI-2 veins free; MI and M2 well separated at base; M3 from lower angle of cell; 1A+2A clear; 3A faintly indicated. Hindwing with Sc+RI connected with Rs from 1/2 to 3/4: M1-3, and CuA1-2 veins free; M1 and M2 well separated at base; M2, M3 and CuAÏI stemming from lower angle of cell; anal veins clearly distinct. Abdomen: Male intersegmental membrane VIII-IX without associated sclerites or hair-like scales. Sternum VIII broadly sclerotized at base, with short median extension and long, thin lateral extensions reaching apex; tergum VIII with broad sclerotized band along whole segment medially. Female segment VII well sclerotized, narrower and longer than preceding segment, with tergum a large quadrangular plate. Tympanal organs (n—6) (Fig. 5): Tergo-sternal sclerite with broadly rounded, deeply concave ventral margin. 642 B. LANDRY ET AL. Tympanum plane almost at right angle from sternal plane. Tympanic frame slightly projecting ventrad of margin of segment. Tympanic crest short, situated slightly anterad of middle. Tympanic drum short, slightly longer than wide, extending anterad to base of bridge. Transverse ridge slightly concave medially, without tympanic pockets, or unapparent, blending with surface of sternum. Tympanic bridge about 1/3 length of drums. Praecinctorium only slightly bilobed. Male genitalia (n=3) (Figs 6, 7). Tegumen with wide median ridge, narrowly ex- tended laterally at apex, widened at base and narrowly connected with lateral ridges; area between median and lateral ridges more thinly sclerotized, slightly bulged and with scale sockets, as opposed to bare ridges. Uncus reduced, mostly thinly sclerotised, rounded, apically setose, occasionally with very short median depression dorsally; ven- tral margin more thickly sclerotized, occasionally with very tiny point medially. Short arms of gnathos (sensu Solis & Metz, 2011) fused with narrow apicolateral ridges of tegumen, narrowly triangular, not connecting medially. Dorsal articulation of valva with vinculum of adjacent type (see Solis & Maes, 2003). Costa of valva with medially directed projection posterad of dorsal articulation of valva with vinculum; projections not connected medially and supporting rounded setose pads ventrally. Valva short, nar- rowing to half basal width near middle, apically rounded, with digit-like, medio- dorsally recurved projection (sella) medially between pair of ridges, with shallow rounded cavity ventrad of sella, with long, abundant setation on basal part of sella dorsally. Juxta a thin, elongate plate with lobed ventral and apical margins, with short wing-like extensions laterally before middle. Vinculum shorter than tegumen + uncus, narrow, with anterior end curved upward, apically blunt in lateral view. Phallus short, stout, without pronounced coecum penis; vesica with bunch of about 20 short, slender cornuti. Female genitalia (n=3) (Fig. 8). Papillae anales simple, rounded, setose, uncon- nected dorsally, with straight sclerotized band at base. Apophyses posteriores straight, reaching middle of segment VIII. Latter with well sclerotized plates laterally, sparsely setose, expanding apicoventrally toward midline, but medially not connected, dorsally approximate on distal half and fused on proximal half. Apophyses anteriores slightly curved and longer than posterior ones, not quite reaching middle of segment VII. Lamella postvaginalis triangular, located at base and between apical ventral extensions of sclerotized plates of segment VIII. Ostium bursae at bottom of cup-like, thinly sclerotized antrum. Ductus bursae with girth about 1/3 width of middle of antrum, more or less ridged on distal half, proximal half gradually expanding, without colli- culum. Ductus seminalis arising subdistally, at-slightly less than 1/3 of length of ductus bursae from ostium. Corpus bursae circular, with one small, spine-like signum ven- trally near distal end. ETYMOLOGY: The name is derived from a frequent interjection heard in Ecuador, chévere, which means great, nice, or cool. The unusual maculation of the moth prompted this interjection, or a synonym, to me and others who examined it. This type of maculation is found in another species of Galapagos Spilomelinae, but not in any other members of this subfamily as far as we know. BIOLOGY: The caterpillar of Cheverella galapagensis is a borer in stems of Tournefortia pubescens Hook. f. (Boraginaceae). One moth was reared by Lazaro NEW GENUS AND SPECIES OF SPILOMELINAE FROM THE GALAPAGOS ISLANDS 643 FIG. 3 Cheverella galapagensis Landry, sp. n. Head of paratype female from Santa Cruz Island (CNC). Roque-Albelo in 1999, from a plant growing on the Barranco, just behind the Charles Darwin Research Station on Santa Cruz Island. This endemic species of ZTournefortia is known to contain pyrrolizidine alkaloids (Roque-Albelo ef al., 2009). These are known to protect Ufetheisa connerorum Roque-Albelo & Landry (Lepidoptera, Arctiidae) (formerly mostly known as U. galapagensis (Wallengren)) moths from being consumed by Eustala (Araneidae) spiders (Garrett et al., 2008). Whether or not Cheverella also stores pyrrolizidine alkaloïids remains to be discovered. REMARKS: The medially directed projections arising from the costal edge of the valvae posterad from the dorsal articulations of the valvae with the vinculum are not called a transtilla as the definition of this structure is restricted to ‘the [dorsal] sclerotisation of the diaphragma’ (Solis & Maes, 2003). SYSTEMATIC POSITION: The current classification of Neotropical Spilomelinae (Munroe, 1995) recognizes 14 groups of genera and S1 unplaced genera. Un - fortunately, Munroe did not provide diagnostic characters to support his groups, and there is no classification available for any other Spilomelinae fauna. Therefore, we examined representatives of Munroe's generic groups to find apparently diagnostic character combinations, and we comment on their validity and applicability to the newly described genus. Cheverella was found to have affinities with the Hydriris and 644 B. LANDRVY ET AL. the Siga groups of Munroe (1995). With the Hydriris group, Cheverella shares a reduced uncus without robust bifid spines and the presence of setose pads in the vici- nity of the transtilla in Choristostigma Warren. However, in the other genera of the group, the setose pads are in different positions (on the costa in Geshna Dyar, on the tegumen in Hydriris Mevyrick) or absent in Nehydriris Munroe. Most of the members of this group share tufts of setae at or around the base of the costa of the valva, but these setae are lacking in Cheverella and Nehydriris, and the female genitalia vary among the three genera for which they are known. The Diagnosis above explains some of the differences between the female genitalia of Cheverella and Choristostigma. Those of Geshna show a poorly differentiated, elongate corpus bursae without signum, while those of Hydriris have a very short ductus bursae and an elongate corpus bursae with an appendix bursae and with two large signa showing about 12 long, thin and curved projections. The labial palpi are variable. They are upturned in Hydriris and Geshna but porrect with downturned apical meron in Choristostigma, so the upwardly directed palpi of Cheverella fit in this range. Hence, there 1s no clear indication that the Hydriris group is monophyletic as presently constituted. The Siga group of Munroe (1995) comprises large, robust-bodied Neotropical moths. Our concept of the group is here informed by our addition of Loxomorpha Amsel and Maracayia Amsel, because they share the same structural characters and known larval habits, despite their much smaller size. So defined, the Siga group varies greatly in maculation, but the forewing postmedial line 1s usually roundly concave on the anal fold. The labial palpi are short or obliquely ascending with a short apical meron. The male genitalia are robust with a moderately inflated sacculus and well- sclerotized costa, and the saccus is absent or weakly developed. The sella is aciculate in most member taxa (as in the Hydriris group), but it is quadrate in some (Cirrho- cephalina Munroe) or with a basal process (Laniifera Hampson). The uncus 1s bifid with a short stalk or none at all (with the two uncus arms arising from the tegumen separately), and the apices of the uncus are armed with robust, bifid chaetae. In the female genitalia, where known, the ostium bursae is strongly sclerotized and shaped like a funnel or pitcher plant, and the corpus bursae lacks a signum. Known larvae (Laniifera, Beebea Schaus, Loxomorpha, Maracayia) are all borers in Cactaceae. Among these characters, Cheverella shares the straight and obliquely ascending labial palpi, robust genitalia with inflated sacculus, and the distally quadrate sella of some members. The breadth of the ostium bursae is similar, but it is only weakly sclerotized in Cheverella. The absence of an uncus, bifid or otherwise, would be explained as derived from the reduced bilobate condition seen in Zeuzerobotys Munroe. Although Cheverella is much smaller than most, it is comparable in size to Loxomorpha species. The black and white wing coloration is shared with Zeuzerobotys. The following characters are shared by the two groups and Cheverella, so although they do not favor placement in either group, they exclude Cheverella from many other spilomeline groups. The valve costa is straight or slightly concave, and the apex of the valve is rounded but attenuate (narrower than the valve width across the costa). The valve sella originates variably from the costal half of the valve or near middle of the valve. The hindwing maculation is nearly absent, and the praecinctorium is weakly (not strongly) bilobed. NEW GENUS AND SPECIES OF SPILOMELINAE FROM THE GALAPAGOS ISLANDS 645 KS (4) Cheverella galapagensis Landry, sp. n. Wing venation of paratype male (MHNG). (5) Cheverella galapagensis Landry, sp. n. Tympanal organs of paratype male, without praecinc- torium (MHNG). FIGS 4-5 Other characters were examined (tympanal organs, venation) but were found to be either invariant or so variable as to be uninformative of relationships at the level of generic group. The diagnostic characters in these groups (and others) should be evalu- ated in a phylogenetic context to determine their influence and to help uncover the relationships of Cheverella. The biogeographic relationships of Cheverella are ambiguous but suggest testable hypotheses. The Hydriris group includes one Neotropical member (Nehydriris of southern Brazil) and the cosmopolitan Hydriris; the other members range to the southern Nearctic. As far as known, the Siga group is exclusively Neotropical to southern Nearctic, with members distributed in many subregions. Among these, Beebea guglielmi Schaus is endemic to the Galapagos Islands, so despite the great difference in body size and maculation, the possibility of a close relationship should be considered. If Cheverella belongs to the Siga group, two hypotheses suggest them- selves. A close relationship with Beebea would support rapid evolutionary divergence in body size and wing maculation, which might be expected on the Galapagos. Conversely, a closer relationship with a mainland taxon would suggest more than one dispersal event to the islands or a more complex biogeographic scenario. If Cheverella is more related to some other spilomeline group, similarly intriguing hypotheses may present themselves. Cladistic and biogeographic analyses of Spilomelinae are greatly needed for choosing among these alternative hypotheses. Cheverella galapagensis Landry, sp. n. Figs 1-8 MATERIAL EXAMINED: Holotype male: 1- ‘ECUADOR | GALAPAGDOS | Santa Crüz | Los Gemelos | 31.1.1989, Mfercury]V{[apour]L[ight] | B. Landry’ [printed in black ink on white card stock, with ‘ECUADOR’ sideways on left]; 2- ‘HOLOTYPE | Cheverella | galapagensis | B. Landry’ [hand-written in black ink on red card stock]. Deposited in the CNC. 646 B. LANDRY ET AL. PARATYPES: 26 d, 49 ©, 1 of unknown sex from the Galapagos Islands, Ecuador: — Fernandina: 1 4, SW side, crater rim, G[lobal]P[ositioning]S[system]: 1341 m, S 00° 21.910’, W 091° 34.034’, 12.11.2005, ufltra-|v{iolet|l[ight] (B. Landry, P. Schmitz). — /sabela, Alcedo: 1 ®, Lado Este, 700 m elev{ation]., 6.iv.1999, uvl-f[?].I[ight] (L. Roque); 1 &, 1 9, NE slope, about 400 m up (S) Los Guayabillos camp, GPS: 892 m elev., S 00° 25.208’, W 91° 04.765, 1.iv.2004, uvl (B. Landry, P. Schmitz); 1 &,1 8 (DNA voucher, Lepidoptera, M. Nuss 2007, no. 263), lado NE, 400 m [elev.], camp pega-pega, 15.iv.2002, uvl (B. Landry, L. Roque); 2 4,1 9%, 570 m elev., 11.x.1998, uvli (L. Roque). — /sabela, Sierra Negra: 3 & , 11 km N Pluer]to Villamil, 13.11.1989, Mfercury]V{apour]L{ight] (B. Landry); 1 ©, same locality, Gf[lobalP[osi- tioning]S[system]: S 00° 87.613’ [sic], W 91° 0.903’, 9.iv.2004, uvl (P. Schmitz); 1 , San[to] Tomas, 1200 Ff[eelt. Alt[itude]., 22.viu.1906 (F. X. Williams); 1 à, 1 $, Alemania, xi.1974 (T.J. de Vries); 2 ®, Corazon Verde, xi-xii.1974 (T.J. de Vries). — Zsabela, Volcan Darwin: 1 4, 300 m elev., 15.v.1992, MVL (B. Landry); 1 4, 630 m elev., 16.v.1992, MVL (B. Landry); 1 & (dis- sected, slide MHNG ENTO 6070), same data except 17.v.1992; 2 d (one dissected, slide MH- NG ENTO 3041), 300 m elev., 20.v.1992, MVL (B. Landry). — San Cristobal: 1 ® (dissected, slide CNC PYR 366), pampa zone, 18.11.1989, MVL (B. Landry); 1 ® , La Toma, ca. 5.6 km East El Progreso, GPS: 299 m'elev.,.S:00%55:356% W°089° 31089 23=n.2005 uv (B lande) Santa Cruz: 1 %, Cfharles|D{[arwin]R[esearch]S[tation], arid zone, 19.1.1989, MVL (B. Landry); 1 ©, 4 km N Puerto Ayora, 20.i.1989, MVL (B. Landry); 4 ®, Media Luna, pampa zone, 21.1.1989, MVL (B. Landry), 2 4, 1 ©, same data as holotype; 1 d, Cfharles] Darwin Research]. Sta[tion]., alt. + 5 m, 2..1970, at u.v. “blacklight” (R. Silberglied); 1 &, Tortuga Resferve]., W S[an]ta Rosa, 6.1.1989, MVL (B. Landry); 1 of sex unknown, Media Luna, pampa zone, 8.11.1989, MVL (B. Landry); 1 ®, Horneman Farm, 220 m, 16.11.1964 (D. Q. Cavagnaro),; 1 $, same data except 25.11.1964; 1 ©, same data except 30.11.1964; 3 ©, same data except 5.iv.1964; 2 ©, same data except 3.v.1964; 1 ©, same data except 7.v.1964; 1 ©, 2 km W Bella Vista, 27.11.1989, MVL (B. Landry), 1 4, Academy Bay, Darwin Research Sta[tion]., 27.11.1964 (D. Q. Cavagnaro), 1 4,3 9, [no precise locality], 111.1969, B.M. 1970-172, Ref. No. L.80 (no collector); 1 ®, Ef[stacion|C{ientifica|C[harles]D{[arwin]., 6.11.1992, uvl (B. Landry); 1 4 (dis- sected, slide MHNG ENTO 6069), transition zone, recently cut road, GPS: S 00° 42.528’, W 90° 18.849’, 12.11.2004, uvl (B. Landry, P. Schmitz); 1 ©, low agriculture zone, GPS: S 00° 42.132, W 90° 19.156’, 13.11.2004, uvl (B. Landry, P. Schmitz); 1 ®, Finca S[teve]. Devine, 17.11.1989, MVL (B. Landry), 1 à, grassiand, 750 m, 6.iv.1964 (D. Q. Cavagnaro); 1 d, vic[inity]. “Mirador”, W of Media Luna, alt. + 620 m, 26.v.1970 (Silberglied); 4 ® (one dissected, slide MNHG ENTO 6068), Los Gemelos, 27.v.1992, MVL (B. Landry); 2 ®, Station Darwin, lumière, x.1964 (J. & N. Leleup); 2 © , Hacienda Schiess., x1.1974, B.M. 1975-7, Ref. No. L (no collector); 1 4, CDRS, Barranco, barrenador de tallos [de] Tournefortia pubecens [sic], emergio 17.xi.1999 (L. Roque); 1 %, 80 [referring to note in notebook of ?T. J. de Vries, deposited at CDRS|. — Santiago: 1 ®, N side, GPS: 527 m elev., S 00° 13.690”, W 90° 44.135’, 5.111.2005, uvl (P. Schmitz); 1 , Aguacate [camp], 520 m elev., 6.iv.1992, MVL (B. Landry); 3 ©, Central [camp], 700 m elev., 9.iv.1992, MVL (B. Landry); 1 4 (wings on slide MHNG ENTO 3043), 1 9, Aguacate, 520 m elev., 12.iv.1992, MVL (B. Landry). Deposited in the BMNH, CAS, CDRS, CNC, MCZ, and MHNG. ETYMOLOGY: From the name of the archipelago where this species 1s pre- sumably endemic. DIAGNOSIS: Currently unique within the genus Cheverella, this species can be separated from presumably related taxa as mentioned in the generic diagnosis above. In the Galapagos, this species is unique by virtue of its whitish grey colour with deep dark brown markings on the forewing, head, and thorax. An undescribed species of endemic Agathodes Guenée (Pyralidae, Spilomelinae) has a similar colour and pattern, but its wings are narrower and longer, and its hindwing is completely suffused with brown, with a light purple shine. DESCRIPTION: MALE (n=27) (Fig. 1). Head white with greyish brown as spot between antennae, and ventrally behind eyes. Antenna white, with brown on basal NEW GENUS AND SPECIES OF SPILOMELINAE FROM THE GALAPAGOS ISLANDS 647 FIGS 6-8 Cheverella galapagensis Landry, sp. n. Male genitalia of paratype (MHNG). (6) Genitalia without phallus. (7) Phallus. (8) Cheverella galapagensis Landry, sp. n. Female genitalia of paratype (CNC). flagellomeres. Haustellum and maxillary palpus white. Labial palpus mostly white, brown laterally on palpomeres II and III. Thorax white with brown spots as shown, deeper brown (almost black) at collar, greyish brown on metascutellum. Foreleg with greyish brown on dorsal side, white ventrally; midleg white with brown at tips of femur and tibia; hindleg entirely white. Forewing white with deep dark brown spots and lines as shown, sometimes (1 specimen) with more extensive dark brown suffusion, especially between subapical line and outer margin. Forewing length 6.4-9.6 mm (holotype 8.7 mm). Hindwing white with more or less strongly marked subapical and apical lines, except in anal sector. Abdomen mostly white, with some greyish brown dorsally on all or most segments. Male genitalia: See generic description. FEMALE (n=49) (Figs 2, 3, 8). Forewing length 7.7-10.3 mm. Female geni- talia: See generic description. REMARKS: As mentioned above, L. R.-A. reared the only known larva from the host plant Tournefortia pubescens Hook. f. (Boraginaceae). This taxon is endemic to the Galapagos and found on the islands of Fernandina, Floreana, Isabela, Pinzon, San 648 B. LANDRY ET AL. Cristobal, Santa Cruz, Santiago, and Wolf (McMullen, 1999). Thus, the moth may also occur on the islands of Floreana, Pinzon, and Wolf, where adults have not yet been collected. The latitude and longitude data of the specimen collected on Sierra Negra, Isabela, 11 km north of Puerto Villamil, on 9 April 2004 by P. Schmitz were taken in decimal degrees shown above in degrees, minutes, and seconds. In degrees with decimal minutes these data correspond to S 0° 52.568, W 91° 05.418. ACKNOWLEDGEMENTS We are thankful to the authorities of Parque Nacional Galapagos and those of the CDRS for allowing and facilitating the field work of B. Landry and L. Roque, and for permits to export specimens. BL is greatly indebted to Stewart B. Peck, his Ph.D. advisor at Carleton University, Ottawa, for taking him on to explore the Galapagos in 1989 and 1992, and for his inspiring companionship in the field. This fieldwork was supported by an operating grant to S. B. Peck from the Natural Sciences and Engineering Research Council of Canada for field research on arthropod evolution. Other much appreciated field companions during one or more of BL’Ss five expeditions to the Galapagos were Novarino Castillo, Charlotte Causton, Joyce Cook, Moraima Inca, José Loaiza, Ricardo Palma, L. Roque-Albelo, Patrick Schmitz, Bradley J. Sinclair, and Eduardo Vilema. BL is also grateful to the Charles Darwin Foundation and the Galapagos Conservation Trust (UK) for providing financial support for his in- vestigations at the BMNH in 2000 and in the Galapagos in 2002. In the BMNH we would like to acknowledge the great support received from curators Michael Shaffer (deceased) and Kevin Tuck. JEH was supported by a Rea Postdoctoral Fellowship (CMNAÆ, Pittsburgh) during his study of Spilomelinae. JEH thanks Alma Solis for pro- viding a manuscript of a revision of the Polygrammodes group by E.G. Munroe (deceased), which partly informed the discussion of the Siga group. Finally, we thank Philippe Wagneur (MHNG) for his photos of the moths shown here, Florence Marteau (MHNG) for producing the plates, Louis Marcotte (Gatineau, Canada) for helping with GPS data and jargon, pyraloid workers E. Munroe, M. Nuss, M. Shaffer, and Alma Solis for their expertise on this species’ taxonomic and systematic status, and the reviewers for their comments. | REFERENCES GARRETT, S. E., W. E. CONNER & L. ROQUE-ALBELO 2008. Alkaloiïdal protection of Ufetheisa galapagensis (Lepidoptera: Arctiidae) against an invertebrate and a vertebrate predator in the Galapagos Islands. Galapagos Research 65: 2-6. LANDRY, B. 1995. A phylogenetic analysis of the major lineages of the Crambinae and of the genera Of Crambini of North America (Lepidoptera: Pyralidae). Memoirs on Entomo- logy, International, Gainesville. Vol. 1. 245 pages. LANDRY, B. 2006. The Gracillariidae (Lepidoptera, Gracillarioidea) of the Galapagos Islands, Ecuador, with notes on some of their relatives. Revue suisse de Zoologie 113: 437-485. MCMULLEN, C. K. 1999. Flowering plants of the Galäpagos. Cornell University Press, Ithaca and London. xiv + 370 pp. MINET, J. 1982. Les Pyraloidea et leurs principales divisions systématiques. Bulletin de la Société entomologique de France 86: 262-280. NEW GENUS AND SPECIES OF SPILOMELINAE FROM THE GALAPAGOS ISLANDS 649 MUNROE, E. G. 1976. Pyraustinae, Pyraustini (pp. 1-78). /n: Dominick, R. B. ef al. (eds). The Moths of America North of Mexico Fasc. 13.2A, Pyraloidea, Pyralidae (part). £. W Classey Limited and the Wedge Entomological Research Foundation. 78 pp. + plates 1-4 & A-H. MUNROE, E. G. 1995. Pyraustinae (pp.53-79). /n: Heppner, J.B. (ed.), Atlas of Neotropical Lepidoptera, Volume 3. Association for Tropical Lepidoptera and Scientific Publishers, Gainesville. NUSS, M. B. LANDRY, F. VEGLIANTE, A. TRÂNKNER, R. MALLY, J. HAYDEN, A. SEGERER, H. LI, R. SCHOUTEN, M. A. SOLIS, T. TROFIMOVA, J. DE PRINS & W. SPEIDEL 2011. Global Information System on Pyraloidea. Museum für Tierkunde, Dresden. www.pyraloidea.org. ROQUE-ALBELO, L., S. E. GARRETT, W. E. CONNER 2009. Darwin’s moth: Ufetheisa in the Galäpagos Islands (pp.207-222). /n: Conner, W.E. (ed.) Tiger Moths and Woolly Bears: Behavior, Ecology, and Evolution of the Arctiidae, Chapter 14. Oxford University Press, New York. SOLIS, M. A. & K. V. N. MAES 2003. Preliminary phylogenetic analysis of the subfamilies of Crambidae (Pyraloidea Lepidoptera). Belgian Journal of Entomology 4: 53-95. SOLIS, M. A. & M. A. METZ 2011. Male genital homology based on musculature originating from the tegumen in snout moths (Lepidoptera: Pyraloidea). Arthropod Structure & Development 40: 435-448. Cr) 2 CL nat boex - nes D el Les ul DIR MTL serré $a A UP ni f D'EAU PE ‘ê sci l POUPEE | dr à «a ig 7e dE su & ARS | Fr : v - : + # ed 41 ete ALICE AUERRR y CETTE LA 1" VUE LE L Ty hé nn’ NE RER A" 071: | MOE i PR Re Li, LE és ‘'aanire (wi ft IVITARES | Éte | on ere - 4 | y j nr ça Mage mA ÉT ARE Er 4 EF} anve rt MT: ne 14h ua; yQù Crabe ‘rte ù Ro R : Ag © au 06 RARES £ mer eo | - LAN et +16 = 2 ST PE CU . -: LL LU. D. 1 » UN | « dv 2 L = LE 2 & LA = LA — t : = ss (NN _ É > ÿ ® eu - (4 DS UN, | se : 2 © nn 2 s _. es PU | d re : ” = | a del . _ à : e + - L ÉCRIS “E e\ L tj U o PT. C4 : ER a an. LE : & < 7 | ee 27) « _ TT > bre +4 pe 2 ‘ . L4,.6.4 Ÿ Le € QE 2. 4 0,9 mer LE és CE 6 , vu 4 s > Er re Fan ‘Re « + » : té LD : ui 4 (4 é ES : EE" L b SE 1." ra 17 Es » 1 … = ? ù : » CRE ai . | | do F 2 on : . et Le 9 PE : . 4 e : LE | Fe vs D. * . : a Fr QU A 4 | ù L de _ . Ce ne 0x Me 4 YO : 2 PRE) REVUE SUISSE DE ZOOLOGIE 118 (4): 651-657; décembre 2011 A new species of Anthrenus Geoffroy, 1762 (Coleoptera: Dermestidae) from Oman, with a key to related species Marcin KADEJI & Jifi HAVA2 l Department of Biodiversity and Evolutionary Taxonomy, Zoological Institute, University of Wroctaw, ul. Przybyszewskiego 63/77, 51-148 Wroctaw, Poland. E-mail:entomol(@biol.uni.wroc.pl. 2 Private Entomological Laboratory and Collection, Unëtice u Prahy 37, 252 62 Praha-zâpad, Czech Republic. E-mail: jh.dermestidae(@volny.cz; http://www.dermestidae.wz.cz. A new species of Anthrenus Geoffroy, 1762 (Coleoptera: Dermestidae) from Oman, with a key to related species. - Anthrenus (Anthrenus) ardoi sp. n. from Oman is described. The habitus, antennae, and male genitalia are illustrated and compared to related species. The new species 1s characterized by eye with median side broadly and deeply emarginated at about anterior 1/3, 11-segmented antenna and broadly oval scales of the dorsum. Elytra covered with oval white patch on each side of the suture at its base, with three white, transverse, irregularly interrupted patches. The new species 1s most similar to Anthrenus (s. Str.) namibicus Häâva, 2000 and A. (s. str.) flavipes flavipes LeConte, 1854. Identification features to externally similar species of the genus are given. The most distinctive taxonomic characteris- tics concern the tibiae, the male genitalia and the 9h abdominal sternite are also given. Keywords: Coleoptera - Dermestidae - Anthrenus - new species - taxonomy - morphology - comparative study - key - Oman. INTRODUCTION Genus Anthrenus 1s one of the 57 genera within the family Dermestidae (Häva 2007, 2010), and is characterized by the presence of scales covering almost a whole body. The scales often create colorful and very useful in identification patterns on the dorsum, and on the visible abdominal ventrites I-V. Currently genus includes 220 species, grouped in 10 subgenera (Häva 2003, 2008). Most of the species have been described from Palearctic and Afrotropical regions. Only five species have been recorded from Oman (Häva 2007, 2010). Two of them are members of the subgenus Anthrenops Reitter, 1881 (4. cervenkai Häva & Herrmann, 2006, À. coloratus Reïtter, 1881), one of the subgenus Anthrenodes Chobaut, 1898 (4. malkini Mroczkowski, 1980), another one of Anthrenus s. Str. (A. flavipes flavipes LeConte, 1854) and the last one of Nathrenus Casey, 1900 (A. jakli Häâva, 2001). In the material from Oman studied recently we have found specimens representing a new species of the nominotypical subgenus. Its description is given below. Manuscript accepted 28.07.2011 652 M. KADEJ & J. HÂVA MATERIAL AND METHODS The following abbreviations to measurements were made: total length (TL) - linear distance from anterior margin of pronotum to apex of elytra. pronotal length (PL) - maximum length measured from anterior margin to posterior margin. pronotal width (PW) - maximum linear transverse distance. elytral length (EL) - linear distance from shoulder to apex of elytron. elytral width (EW) - maximum linear transverse distance. The following abbreviations are used in the text: IZDBET Institute of Zoology, Department of Biodiversity and Evolutionary Taxonomy, Wroctaw, Poland. JHAC Private Entomological Laboratory & Collection, Jii Hâva, Prague-west, Czech Republic. MZLU Lund University, Lund, Sweden. NHMW Naturhistorisches Museum, Wien, Austria. MHNG Muséum d'Histoire Naturelle, Geneva, Switzerland. OXUM Oxford University Museum of Natural History, England. Locality labels are cited in the original version. Separate labels are indicated by a slash ( / ). Author’s remarks are in square brackets [ ]. Specimens have been labeled with red printed label of the following text: “HOLOTYPE [or PARATYPE, respectively] Anthrenus (Anthrenus) ardoi n. sp.”. Morphological structures were cleared in boiling 10% KOH solution, rinsed in distilled water, mounted in glycerin, and then examined, measured, and 1llustrated un- der a Nikon Eclipse E 600® phase contrast microscope. External structures were examined under a Nikon SMZ-800F stereoscopic microscope. The dissected parts were placed with glycerin in a plastic microvial and attached to the pin of the specimen. Photos were taken with a Canon 500D% and a Nikon Coolpix 4500®. Terminology: The terminology used in this paper follows Beal (1998) and Lawrence & Slipiñski (2010), distribution and classification follows the world cata- logue of Häva (2010). SYSTEMATIC PART Megatominae Leach, 1815 Anthrenini Casey, 1900 Anthrenus Geoffroy, 1762 Anthrenus (Anthrenus) ardoi Sp. n. Figs 1-9 MATERIAL STUDIED: Holotype (4 ): Oman, Ghuzyan rd. AI Khabural-liberi 10.1V.1985 leg. Paul Ardô (MZLU). Allotype ($ ): Oman, Yiti Wadi 7.1V.1985 Paul Ardô (MZLU). Paratypes (16 exx. not sexed): (2 exx.) the same datas as holotype (1 MZLU, 1 JHAC); (8 exx.) the same data as allotype (5 MZLU, 1 MHNG, 1 IZDBTE, 1 JHAC); (5 exx.) Oman, AI Sinain, 10.1V.1985, leg. Paul Ardô (4 MZLU, 1 JHAC),; (1 ex.) Oman, Nizwa, 12.1V.1985 leg. Paul Ardô (MZLU). ETYMOLOGY: The epithet is a patronym honoring Dr. Paul Ardô (NHMW), who collected the type series material. NEW SPECIES OF ANTHRENUS FROM OMAN 653 FIGS 1-4 Anthrenus (Anthrenus) ardoi. (1) Habitus, dorsal aspect (holotype, male). (2) Habitus, dorsal aspect (allotype, female). (3) Visible abdominal ventrites I-V (holotype, male). (4) Visible abdominal ventrites I-V (allotype, female). DIAGNOSIS: The new species can be differentiated from some of the species occurring in Oman by the: 1) The total number of antennomeres: in À. ardoi antennae with eleven antenno- meres; in À. cervenkai Häva & Herrmann, 2006 and 4. coloratus Reitter, 1881 with nine antennomeres; in À. malkini Mroczkowski, 1980 with ten antenno- meres. 2) The morphology of eye: in 4. ardoi eye with median side broadly and deeply emarginated at about anterior 1/3; in À. cervenkai, À. coloratus and À. jakli eye oval. Due to variation in the dorsal setal pattern, the new species dorsal appearance resembles some forms of Anthrenus (s. str.) namibicus Häâva, 2000 and A. (s. str.) flavipes flavipes LeConte, 1854 and can be easily confused with them. The distinction between these two species can be done based on the following characteristics: 1) In À. ardoi and 4. flavipes flavipes tibial spines on first pair of legs absent; in À. namibicus present. 2) In À. ardoi claws of third pair of legs with one small denticle, in À. flavipes flavipes denticle not present; in À. namibicus three denticle present. 3) In À. ardoi apex of paramers slightly curved in to middle, paramers with apex and median lobe slender and narrower, median lobe with distinct extension in 1/2 of its length (Fig. 6); in À. flavipes flavipes and À. namibicus, although male genitalia are similar in general shape to 4. ardoi, their parameres are broader; while in 4. namibicus Häva, 2000 parameres are distinctly curved toward the middle, setae present only on apex of the parameres, median lobe with distinct extension in 3/5 of its length; in À. flavipes flavipes parameres are curved to ward the middle, setae present on apex and in the inner margin of parameres, median lobe without distinct extension. 654 M. KADEJ & J. HAVA 4) In À. ardoi 9 abdominal sternite without distinct waist, with the same width throughout 1ts length, the apex with cordate, deep indentation, sclerotization on the sides and only in the mid section of the upper part - extending to the top and underneath; in 4. flavipes flavipes 9h abdominal sternite spatulate, with the apex narrow and flat, three times wider posteriorly than the apex width, distinct waist in a mid section present, sclerotization on the sides and the mid section only - extending underneath; in 4. namibicus 9h abdominal sternite without distinct waist, with the same width throughout its length, the apex almost flat with slight and shallow indentation, sclerotization on the sides and only in the mid section of the upper part - extending to the top and underneath. DESCRIPTION Holotype male. Body: measurements (mm): TL 2.0 PL 0.6 PW 1.3 EL 1.4 EW 1.7 SL 1.1 SW 1.5. Body convex, rounded, integument brown, finely punctured, covered by broadly oval scales. Scales: two times as long as width, their surface mostly with 14 complete, linear ribs; the apex of the scale is truncated or rounded and an apical lappet is not present (Figs 8-9). Head: is characterized by large convex eyes. Eye with median side broadly and deeply emarginated at about anterior 1/3. Median ocellus present on the frons. Antennae with 11 antennomeres, with 3-segmented antennal club (Fig. 5); anten- nomeres dark-brown. Antenna occupies whole cavity of antennal fossa. Antennal club in both sexes occupies less than half length of the antenna, and is distinctly longer than length of two basal segments combined. Antennal fossa is completely open along lateral margin of the pronotum and occupies 1/3 of the length of lateral margin. Dorsal margin of antennal cavity not visible from above. Dorsal surface covered with white, light- and dark brown scales (Figs 1-2) as follows: pronotum With mixed and irregular light-dark brown and white scales; small white patch near lateral angles enclosing light brown patch. Elytra: covered with one oval white patch on each side of the suture at its base, with three white, transverse, irregularly interrupted patches: first in basal third, second close the middle, and third in apical fourth or fifth. Dark brown patches are present near humeri, in the central part of disc close to suture, and in basal third near the lateral margin. The remaining areas between bands are covered with light brown scales mixed with single white scales. Ventral surface: with all scales white except for visible ventrites II-V covered with mix of yellowish (light brown) and white scales; first abdominal ventrite has stria (Figs 3-4). Legs: brown and covered with white scales on dorsal surface. Tibiae without tibial spines. Tarsus with two slightly curved claws. Claws of the third pair of legs with one small denticle in the half of its length. Male genitalia: as in figure 6. Parameres are deeply U-shaped, covered with few short setae on the lateral margins as well as in the central and inner areas. Median lobe in lateral view straight, with a distal end of aedeagus pointing up; in the frontal NEW SPECIES OF ANTHRENUS FROM OMAN 655 FIGS 5-9 Anthrenus (Anthrenus) ardoi (holotype). (5) Male antenna. (6) Male genitalia. (7) 9h abdominal sternite. (8-9) Scales. view, wider posteriorly, with narrow apodemes which occupying 1/3 of median lobe length (Fig. 6). 9th abdominal sternite as in figure 7. Pygidium with sub-basal, transverse, dark, carina-like line. OBSERVED VARIATIONS: The dorsal patterns vary in color intensity (either darker or lighter). White patches occupies bigger or smaller areas on the pronotum and elytra (Figs 1-2). Body measurements for allotype (female): TL 1.9 PL 0.6 PW 1.2 EL 1.4 EW 1.5 SL 1.0 SW 1.4; for paratypes varying from (mm): TL 2.7-3.3 PL 0.6-0.7 PW 1.4-1.6 EL 2.1-2.4 EW 1.9-2.4. SHORT KEY TO THE RELATED SPECIES AND TO THE KNOWN ANTHRENUS SPECIES FROM OMAN: l na mn 0=lliantennomercs ares. Loos Vous en 2: la. Fee WIN) "ANIÉnNOMEres 7 RE URL OR, TRS NET Eva se 3: 2, Antenna with 10 antennomeres . 4. (Anthrenodes) malkini Mroczkowsk1, 1980 2a. RE D ARTE NOOIITÉS 22 à 2e codae ie MAN e he cu. dé ve | 4, 3. Antennal club with two antennomeres, male antennomere 9 at least 4x longer than antennomere 8 ......... À. (Anthrenops) coloratus Reitter, 1881 3a. Antennal club with three antennomeres, male antennomere 9 at least 2 lon ia amMéennOmMerC Ti, AN AUTO R, FEU RE 2 LT AO EETERRE À. (Anthrenops) cervenkai Hâva & Herrmann, 2006 4. EE TOO ONS CHARDINALEC., 0 eu maso en sim aia «me eme eee e >. A HR PINAP IR OfCyE OVAl : : :...,..,.,.:... À. (Nathrenus) jakli Hâva, 2001 656 M. KADEJ & J. HAVA 5. First pair of legs with tibial spines ........ A. (s. str.) namibicus Häva, 2000e Sa... ‘Firstipair.of legs-without tibial SDINES ER 6. 6. Ventrites II-V covered with mix of yellowish (light brown) and white scales; 9h abdominal segment without distinct waist, with same width throughout its length, apex of segment with cordate, deep indentation; apex of paramers slightly curved in to middle, paramers with apex and median lobe slender and narrower, median lobe with distinct extension in 1/2:0f.1ts length ee RP À. (s. Sstr.) ardoi sp. n. 6a. Ventrites I-V covered with white and light-brown scales; light-brown scales cover posterior margin of ventrites I-V and middle section of ven- trite V; 9th abdominal sternite spatulate, with the apex narrow and flat, three times wider posteriorly than the apex width, distinct waist in a mid section present; parameres curved toward middle, setae present on apex and in inner margin of parameres, median lobe without distinct extension ns do ne OU OC À. (s. str.) flavipes flavipes LeConte, 1854 DISCUSSION: Many of the species of the genus Anthrenus are similar externally. However, there are some species that have varying scales color and patterns (4. la - vipes flavipes LeConte, 1854, À. lepidus LeConte, 1854, À. pimpinellae pimpinellae (F., 1775), À. scrophulariae scrophulariae (L., 1758) , À. verbasci (L., 1767); compare with Beal 1998, Hinton 1945, Kade] 2005a, b). Thus specimens of particular species can be often more similar to other distinct species than to typical form. As a result iden- tification should be based on the general morphology of the male genitalia and details in structure of median lobe, shape of the 9th abdominal segment and respectively galea with lacinia in all cases. ACKNOWLEDGEMENTS We would like to thank to Prof. L. Borowiec (Zoological Institute, Wroctaw University, Poland) for help in improving this manuscript, to R. Danielsson (MZLU) for the loan of material and J. Cooter (OXUM) for help with translation of the manuscript. This work was supported by funding (project no 1244/M/17/2011) from the Institute of Zoology, University of Wroclaw, Poland. REFERENCES BEAL, R. S. 1998. Taxonomy and Biology of Nearctic Species of Anthrenus (Coleoptera: Dermestidae). Transactions of the American Entomological Society 124: 271-332. CASEY, T. L. 1900. Review of the American Corylophidae, Cryptophagidae, Tritomidae and Dermestidae with other studies. Journal New York Entomological Society 8: 51-172. CHOBAUT, A. 1898. Description de quelques espèces et variétés nouvelles de Coléoptères algériens. Revue d'Entomologie Caen 17: 74-88. FABRICIUS, J. C. 1775. Systema Entomologie, sistens Insectorum classes, ordines, genera, species, adiectis synonymis, locis, descriptionibus, observationibus. Flensburgi et Lipsiae: Officina Libraria Kortii, xxx + 832 pp. GEOFFROY, E. L. 1762. Histoire abrégée des Insectes qui se trouvent aux environs de Paris, dans laquelle ces animaux sont rangés suivant un ordre méthodique. 1. Paris: Durand, xxvii + 523 pp. + 10 pl. NEW SPECIES OF ANTHRENUS FROM OMAN 657 HAVA, J. 2000. New interesting Dermestidae (Coleoptera) from the world with descriptions of ten new species. Verofjentlichungen Naturkundemuseum Erfurt 19: 161-171. HAVA, J. 2001. Anthrenus (Nathrenus) jakli sp. n. (Coleoptera: Dermestidae) from the Sultanate of Oman, and distributional notes on some other Anthrenus species. Acta Societatis Zoologicae Bohemiae 65: 1-3. HAVA, J. 2003. World Catalogue of the Dermestidae (Coleoptera). Studie a Zprävy Oblastniho Muzea Praha-vyÿchod v Brandÿse nad Labem a Staré Boleslavi, Supplementum 1: 1-196. HAVA, J. 2007. Dermestidae (pp. 57, 299-320). In: Lôbl, I. & Smetana, A. (eds). Catalogue of Palaearctic Coleoptera. Volume 4. Elateroidea, Derodontoidea, Bostrichoidea, Lymexy- loidea, Cleroidea and Cucujoidea. Apollo Books, Stenstrup, 935 pp. HAVA, J. 2008. Description of a New Subgenus Sertapeacockia subgen. nov. of the genus Anthrenus Geoffroy, 1762 (Coleoptera: Dermestidae: Anthrenini) from Central Asia. Litvijas Entomologos 45: 43-45. HAVA, J. 2010. Dermestidae of the World (Coleoptera). An interactive manual (opened in 2004), available from: hftp://www.dermestidae.wz.cz (last modification 18.01.2010) HAVA, J. & HERRMANN, A. 2006. Additional notes on some Dermestidae and new species from Yemen — Part I. Mitteilungen des Internationalen Entomologischen Vereins 31:1-10. HINTON, H. E. 1945b. À Monograph of the Beetles Associated with Stored Products. Volume 1. London: Order of the Trustees of the British Museum, vi + 443 pp. KADEJ, M. 2005a. Morphological variability of pimpinellae-group (Coleoptera: Dermestidae: Anthrenus) with a key to identification. Part 1. Polskie Pismo Entomologiczne 74: 117-135. KADEJ, M. 2005b. Morphological variability of Anthrenus (Coleoptera: Dermestidae). Part 2. scrophulariae-group with a key to identification. Polskie Pismo Entomologiczne 74: 509-521. LAWRENCE, J. F. & SLIPINSKI A. 2010. 6.1. Dermestidae Latreille, 1804 (pp. 198-206). /n: LESCHEN, KR. A. B., BEUTEL, KR. G., & LAWRENCE, J. F. (eds). Coleoptera, beetles. Volume 2: Morphology and systematics (Elateroidea, Bostrichiformia, Cucujiformia partim). Handbook of Zoology, a natural history of the phyla of the animal kingdom. Volume IV. Arthropoda: Insecta. Part 38 (KRISTENSEN, N. P. & BEUTEL, KR. G., eds). Walter de Gruyter, Berlin, Germany. LECONTE, J. L. 1854. Synopsis of the Dermestidae of the United States. Proceedings of the Academy of the Natural Sciences of Philadelphia 7: 106-113. LINNAEUS, C. von 1758. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, refor- mata. Tomus 1. Holmiae: Laurenti Salvii, IV + 824 + 1 pp. LINNAEUS, C. von 1767. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima tertia, ad Editionen duodecimam reformatam. 7om 1. Pars 11. Holmiae: Laurentii Salvii, pp. 353-1327 34:pp: MROCZKOWSKI, M. 1980. Insects of Saudi Arabia. Coleoptera: Fam. Dermestidae. Part 2, Description of three new species. Fauna Saudi Arabia 2: 124-126. REITTER, E. 1881. Bestimmungs-Tabellen der europäischen Coleopteren. IIT Heft. I. Auflage. Enthaltend die Familien: Scaphidiidae, Lathridiidae und Dermestidae. Verhandlungen der Kaiserlich-Kôniglichen Zoologisch-Botanischen Gesellschaft in Wien 30: 41-94. LT : Didrn. seen mecs RS à bat, (DTA ten sat dit 0e soil AP: » LeRbeher SAS 2 à of 2 svt RÉ d PA pa” æ'à e. se ‘A DA CARE Mg dû + nt RAI MCALUr. mm SL fr etratréihe Dec M mçnat à fins Fe 'e0S rar Rhodes - 4 PT REVUE SUISSE DE ZOOLOGIE 118 (4): 659-666; décembre 2011 First record of the subfamily Pycnocheiridiinae from South America, with the description of Leptocheiridium pfeiferae sen. n., Sp. n. (Arachnida:Pseudoscorpiones: Cheiridiidae) Volker MAHNERT* & Jürgen SCHMIDL** * Muséum d’histoire naturelle, Ville de Genève, case postale 6434, 1211 Geneva 6, Switzerland. E-mail: volker.mahnert(@wanadoo.fr ** Friedrich-Alexander-Universität Erlangen-Nürnberg, Department Biologie, AG Okologie, Staudtstr. 5, 91058 Erlangen, Germany. E-mail: jschmidl(@biologie.uni-erlangen.de First record of the subfamily Pycnocheiridiinae from South America, with the description of Leptocheiridium pfeiferae gen. n., sp. n. (Arachnida: Pseudoscorpiones: Cheiridiidae). - The cheiridiid subfamily Pycnocheiridinae is recorded for the first time from South America, Leptocheiridium gen. n. and its type species pfeiferae sp. n. are described and 1llustrated. The new genus shares with Pycnocheiridium Beier, 1964 (type genus of the Pycnocheiridiinae Beier, 1964) the same morphology of walking legs, eight trichobothria on the fixed chelal finger, only two on the movable finger, and the presence of strongly clavate vestitural setae. Leptocheiridium gen. n. is well characterized by the shape of its slender pedipalps, the presence of three sub-equal setae in the rallum, the presence of five setae on the cheliceral hand, the morphology of the female galea, and the presence of a well-developed transverse furrow on the carapace. Keywords: Taxonomy - ecology - neotropical region - Ecuador - bark dweller. INTRODUCTION Beier (1964) erected, within the family Cheiridiidae, the subfamily Pycnoche: - ridiinae Beier, 1964 for the monotypical genus Pycnocheiridium Beier, 1964 (with its type species mirum Beier, 1964). This genus.takes an intermediate position between Cheiridiidae Hansen, 1893 and Pseudochiridiidae Chamberlin, 1923 in having clearly separated femur and patella on walking legs [/IT, femur+patella of legs III and IV with a distinct, nearly vertical suture (Judson, 1992), eight trichobothria on fixed chelal finger, and 5 setae on the cheliceral hand (pseudochiridnid characters), whilst having normal, unmodified coxae IV also in females, normal-shaped tergites (not chevron- shaped), a rallum with three setae, of which the anterior one 1s larger and dentate (Judson, 1992), and only two trichobothria on the movable chelal finger (cheiïridrid characters). Only a few specimens of Pycnocheiridium mirum are recorded from South Africa (Eastern Cape, Kwazulu-Natal). Nothing is known on its biology (one female was "sifted, (from) very humid rain forest, very thick humus"), and it "appears to be one of the rarest South African species (of pseudoscorpions)" (Judson, 1992). Manuscript accepted 16.09.2011 660 V. MAHNERT & J. SCHMIDL Harvey (1992) placed Cheiridiidae (without mentioning the subfamily Pycnocheiridiinae), together with four other families, in the superfamily Garypoidea. Murienne ef al. (2008) re-established the superfamily Cheiridioidea, including Cheiri- diidae, Pseudochiridiidae and Sternophoridae, by using molecular sequence data. However, their analysis is based on a very low number of genera and species (three) studied. The phylogeny of Cheiridioidea and its validity as a superfamily has also been discussed by Judson (2000). In the course of the worldwide research project (MACAG - Monitoring of Arthropods along Climate and Altitude Gradients: see Schmidl, 2009) conducted by the junior author (JS), 339 samples from tree bark were collected in 2007, 2008 and 2009 at 1000 m (warm rainforest), 2000 m (cloud forest) and 3000 m (Paramo tran- sition forest) in the Podocarpus National Park in Ecuador, to analyze species compo- sitions of arthropods in regard to patterns of diversity between trees, plots and altitu- dinal levels, and for the evaluation of the monitoring performance (application) of the corticolous arthropod fauna. Samples were taken using the barkspray method (protocol according to Schmidl, 2007). A defined area of a living, unbroken tree trunk bark is sprayed with a pyrethroid insecticide spray. Pyrethroids are contact poisons which are highly specific to arthropods and block voltage-sensitive sodium channels in the nerve membrane. Within a few minutes the arthropods lose the ability of coordinated move- ments and tumble down from the substratum. Pyrethroids show an immediate effect on all arthropods, but are relatively harmless for higher vertebrates. Before spraying, a plastic sheet 1s tightly pinned to the bottom of the sampled trunk to collect all affected arthropods, which after 15 minutes are brushed into a small, tree-wise labelled plastic bag (Whirl Paks) containing 70% ethanol. Among the 287 pseudoscorpion specimens, representing seven families and 15 morphospecies, collected during this field survey, we found three specimens of a genus that we place with hesitation in the cheiridnid subfamily Pycnocheiridiinae. À new genus and species are described and figured below. It is well differentiated from Pycnocheiridium, the only other known genus in this subfamily. DESCRIPTION Leptocheiridium gen. n. DIAGNOSIS: Member of Cheiridiidae, ‘subfamily Pycnocheiridiinae. Tegument normally sclerotized, vestitural setae of carapace, tergites and pedipalps strongly broa- dened, leaf-like. Carapace subtriangular, with one pair of eyes, two distinct transverse furrows, the subbasal one flattened, metazone narrow. Most tergites divided, with unmodified lateral borders. Manducatory process rounded, with two marginal setae. Sternites IV to X divided, XI undivided; anal cone with 2 dorsal and 2 acute ventral setae and located between tergite XI and sternite XI; male genitalia of typical cheiri- diid morphology (lateral sacs long, well developed, atrium of ejaculatory canal well developed, median genital sac short), female genitalia consisting of three major cribrate plates, the lateral ones more sclerotized than the larger median one. Pleural membrane striate, partly covered with granula. Chelicera with 5 setae on hand; galea short, distinct apical rami present only in female, subgaleal seta near galea base, rallum LEPTOCHEIRIDIUM GEN. N. FROM ECUADOR 661 with three setae, the distal one with a few fine denticles on anterior margin. Pedipalps coarsely granular, hand nearly parallel-sided; teeth on both fingers mostly acute and contiguous; fixed finger with 8 trichobothria (all situated in basal half of finger), movable finger with 2 trichobothria (b and f); long venom duct present in both fingers. Coxae IV much wider than coxae I. Leg I: femur distinctly longer than patella and with articulation, tarsal segments fused, without visible suture. Leg IV: femur much shorter than patella, suture nearly vertical, tarsal segments fused, without visible suture, no tactile tarsal seta; undivided arolia shorter than smooth claws. TYPE SPECIES: Leptocheiridium pfeiferae sp. n. ETYMOLOGY: The genus name 1s a combination of Cheiridium with the Greek adjective “leptos” (thin, slender), referring to the slender pedipalps. AFFINITIES: Leptocheiridium gen. n. 1s distinguished from Pycnocheiridium by its slender pedipalps, the distinct transverse furrows on its carapace, the presence of 5 setae on its cheliceral hand, the female galea with 6 apical/subapical rami, and the three sub-equal setae of its rallum (in Pycnocheiridium the anterior seta 1s distinctly larger: Judson, 1992). REMARKS: As currently defined, the subfamily Cheiridiinae of the Cheiridiidae, is characterized by having a reduced number of trichobothria on fixed (at most 7) and movable (at most 2) chelal fingers, by the fused femur and patella of leg I and the at least partly fused femur+patella of leg IV; coxa of leg IV also in females unmodified, not enlarged; carapace with two distinct transverse furrows, with depressed metazone (except in Apocheiridium), and two eyes; cheliceral hand with 4 setae; first seta of rallum enlarged. The subfamily Pycnocheiridiinae differs from the Cheiridiinae by having femur and patella of leg I well separated and articulated, femur+patella of leg IV with dis- tinct, nearly vertical suture, and eight trichobothria on the fixed chelal finger. Considering these differences, the Pycnocheiridiinae may even deserve familiar rank. The new genus ZLeptocheiridium gen. n. confirms the main subfamiliar characters, but differs by the following: number of setae on cheliceral hand (4 vs 5) and morphology of rallum (3 setae, the first one not enlarged), which are probably of generic, but not of subfamiliar importance. Leptocheiridium pfeiferae Sp. n. Figs 1-10 HOLOTYPE: MHNG, without registration number; d; Ecuador, Prov. Zamora, env. Estacion Cientifica San Francisco (ECSF), 2000m asl, cloud forest, on tree bark by bark spraying; 31.VIIL.2009, Ig. T. Pfeifer (tree # EL6BOS). PARATYPES: MHNG, without registration number; 1 1 deutonymph; same locality and method, 29.VIII.2009; Ig. T. Pfeifer (tree # EQSB07). REMARK: The three specimens were collected on two different trees several hundred metres apart. Tree EL6B0S had a dimension at breast height (dbh = diameter at 130 cm height) of 26 cm, a medium coverage of climbing plants, a dense cover of epiphytes (mosses, lichens, algae), and a medium rough bark. Tree EQ5SB07 had a dbh of 23 cm, differing from the previous tree by a very smooth bark. Both trees had an intact, unbroken bark. 662 V. MAHNERT & J. SCHMIDL FIGS 1-7 Leptocheiridium pfeiferae gen. n., sp. n., holotype. (1) Carapace, with granules enlarged. (2) Left chelicera & , with rallum and galea of & and © (paratype) enlarged. (3) Left pedipalp. (4) Tricho - bothrial pattern. (5) Tergal seta, enlarged. (6) Left leg I. (7) Right leg IV. Scale unit 0.1 mm. LEPTOCHEIRIDIUM GEN. N. FROM ECUADOR 663 FIGS 8-10 Leptocheiridium pfeiferae gen. n., sp. n. (8) Sternites IT/III of male. (9) Sternites IT/IIT of female, with cribrate plates. (10) Trichobothrial pattern of deutonymph. DESCRIPTION OF ADULTS: Pedipalps, and carapace brown, tergites yellowish brown, legs including coxae reddish brown. Carapace (Fig. 1) subtriangular, broader than long (length/breadth = 0.9), with two transverse furrows, median furrow broad and laterally distad directed, granular, the subbasal one distinct only laterally; two distinct eyes with flat lenses; anterior margin straight, without epistome; coarsely gra- nular; with 42-46 strongly clavate, leaf-like setae (4 along anterior, 7-9 along posterior margin). All tergites visible dorsally, I-X divided, coarsely granular, setae leaf-like (Fig. 5), chaetotaxy of half-tergites: 5-6/6/6/5-6/6/5-6/5-6/4-5/5/3-4, on IV-X one seta anterolaterally placed, tergite XI with 6 setae (no tactile setae). Manducatory process scaly, with 2 marginal setae; pedipalpal coxa coarsely granular, with 8-9 clavate setae, 0-2 smooth setae and one distal tactile seta, coxa I with 6-8 setae (2 slightly clavate), Il with 8-13 smooth setae, III with 15-20, IV with 32-35 smooth setae; genital opercu- lum of & with 28 long setae (Fig. 8), 2/2 short entrance setae, genital organs similar to that of other cheiridiid species (e. g. Dumitresco & Orghidan, 1981: Judson, 1992); genital operculum of ? (Fig. 9) with 15 marginal and discal setae, medial cribrate plate apparently divided into one big and one small part, lateral cribrate plates small (Fig. 9); 664 V. MAHNERT & J. SCHMIDL sternites IV-X divided, the anterior ones scaly, the posterior ones ctenoid-scaly sculptured, some lyrifissures present in front of and between setae, XI undivided, ctenoid-scaly; setae on anterior sternites smooth, acute, on posterior ones slightly clavate (except lateral setae and those of XI), chaetotaxy of half-sternites: 6-8, 0 supra- stigmal setae/7-9+1 suprastigmal seta/8-9/7-8/6-7/5-7/5-6/4, lateral seta on V-X short. Anal cone situated between tergite and sternite XI, with 2 smooth dorsal and 2 (3 in holotype) ventral setae. Pleural membrane shortly granular-striate. Chelicera (F1g. 2) with 5 long acute setae, b distinctly shorter than sb, es as long as sb; fixed finger with 3 retrorse teeth and 2-3 subapical granula, lamina exterior thin; movable finger of 4 with a few thin, thorn-like projections on lateral side, one also at base of galea, with a tooth-like subapical lobe, subgaleal seta reaching clearly beyond tip of galea in d , reaching almost to tip of galea in ®, galea of 4 short (Fig. 2) and rounded, with 2 tiny rounded apical tubercles, galea of ® long, with 6 short sub- apical/apical rami; serrula exterior with 12 lamellae; rallum (Fig. 2) with 3 setae, anterior margin of distal one with 2-3 denticles. Pedipalp (Fig. 3) coarsely granular, vestitural setae leaf-like, chelal finger with fine curved setae; trochanter with small dorsal hump, 2.0-2.1 times longer than broad; femur distally slightly broadened, with a distinct pedicel, 4.3 times longer than broad; patella 3.3 times longer than broad; hand parallel-sided, with pedicel 2.7 times (é ) or 2.3 times (© ) longer than broad and 1.6 times (4 ) or 1.5 times (9) longer than finger; chela with pedicel 4.4 times (4 ) or 3.9 times (®), without pedicel 4.2 times (4 ) or 3.6 times (®) longer than broad; teeth of chelal fingers acute, slightly retrorse (in distal third), then becoming flattened and rounded in basal part, fixed finger with 33 (é ) or 29 (S ) teeth, movable finger with 27 (& ) or 21 (9) teeth; long venom duct present in both fingers, nodus ramosus situated in middle of finger; 10 trichobothria (Fig. 4), all 8 on fixed finger situated in basal half, 2 on movable finger situated in basal third (b and f, following Harvey, 1990); sensillum distal to b. Leg I (Fig. 6): femur and patella articulated, femur 1.6 times (4 ) or 1.4 times (®) longer than patella, femur 2.4-2.5 times longer than deep, patella 1.6 times, tibia 2.3-2.6 times, tarsus 4.3-4.4 times longer than deep, tarsus 1.4-1.5 times longer than tibia; leg IV (Fig. 7): femur much shorter than patella, distinct suture nearly vertical, femur+patella 4.4-4.6 times longer than deep, tibia 4.9 (4 ) or 4.5 (®) times, tarsus 5.2 (S)or 5.6 (9) times longer than deep, no tactile setae present; undivided arolia shorter than smooth claws. MEASUREMENTS (in mm) & (9): Total length 1.4; carapace length/breadth 0.43/0.49 (0.44/0.51); pedipalp length/breadth: trochanter 0.23/0.11 (0.24/0.12), femur 0.44/0.10 (0.47/0.11), patella 0.40/0.12 (0.44/0.13), hand with pedicel 0.36/0.13 (0.37/0.12), length of pedicel 0.03, finger length 0.22 (0.24), chela length with pedicel 0.58 (0.62); leg I length/depth: femur 0.19/0.08 (0.19/0.08), patella 0.12/0.08 (0.14/0.08), tibia 0.16/0.05 (0.16/0.07), tarsus 0.22/0.05 (0.23/0.05); leg IV length/depth: femur+patella 0.40/0.09 (0.39/0.09), tibia 0.31/0.06 (0.32/0.07), tarsus 0.28/0.05 (0.31/0.06). DESCRIPTION OF DEUTONYMPH: Carapace 0.80 times broader than long, co - arsely granular, medial transverse furrow distinct, only lateral depressions of subbasal LEPTOCHEIRIDIUM GEN. N. FROM ECUADOR 665 7 » œ re) De a e ar" : = t — 4 p t _. : os L = L . a à L » | : 4 24080 ‘ É À . 1 1: L EAU Fr ET On | l = A A + a »'é Cr: D 42, LM Di Ébuéh, & LOUE " - td A C4 (rer Fa “à SN DR 2 DATE pan ? à - s Later ; We XP ds à rt: de # 4 k WE: tite “+ .… Fo La À J æ 4 Tr! s0e 4 E LE ct n £ ir ge SAULT | <É Le EMA IS CNRS RS es Le LR DR EE . — 7 Fe + %. ce Ë “se { FL on da re. C8 l REVUE SUISSE DE ZOOLOGIE 118 (4): 723-758; décembre 2011 Taxonomic emendations in the genus Liocheles Sundevall, 1833 (Scorpiones, Liochelidae) Lionel MONOD Muséum d’histoire naturelle, Département des arthropodes et d’entolomogie I, C. P. 6434, CH-1211 Genève, Switzerland. E-mail: lionel.monod@ville-ge.ch Taxonomic emendations in the genus ZLiocheles Sundevall, 1833 (Scorpiones, Liochelidae). - Examination of an extensive material that includes the type series and recently collected specimens enable a precise reassessment of the status of several taxa belonging to the genus ZLiocheles Sundevall, 1833 (Scorpiones, Liochelidae). Two species previously in syno- nymy, 1. e. Liocheles boholiensis (Kraepelin, 1914) and Liocheles neocale- donicus (Simon, 1877), are revalidated. Liocheles australasiae longimanus (Werner, 1939) is elevated to species rank and Liocheles australasiae brevi- digitatus Werner, 1936 is synonymized with Liocheles australasiae (Fabricius, 1775). L. boholiensis (Kraepelin, 1914), L. neocaledonicus (Simon, 1877) and Z. longimanus (Werner, 1939) are thoroughly redes- cribed, diagnosed and illustrated, and their distribution ranges are accurately mapped. Keywords: boholiensis - longimanus - neocaledonicus - taxonomy - Australasia - Indonesia, Sumatra - New Caledonia - Philippines - Bohol. INTRODUCTION The genus Ziocheles Sundevall, 1833 is currently divided into two species groups which are easily distinguishable by the trichobothrial pattern on the retrolateral side of the pedipalpal chela, 1. e. the Ziocheles waigiensis (Gervais, 1843) species group and the ZLiocheles australasiae (Fabricius, 1775) species group (Monod, 2000; Monod & Volschenk, 2004). Three species and two sub-species are recognised within the australasiae-group and six species within the waigiensis-group. However, an ongoing global taxonomic revision of the genus reveals an unsuspected specific diver- sity. Preliminary results of this work were first presented in an unpublished master thesis (Monod, 2000) carried out at the University of Geneva (Switzerland). The present contribution intends to formally present part of this work. Over the years, several authors attempted to revise the genus but were unable to clearly assess the taxonomic status and position of most species, and several valid taxa are currently considered as junior synonyms owing to the paucity of diagnostic characters and to the scarcity of material available in museum collections, especially mature males that are essential for diagnoses of species. Examination of an extensive material, including types, and recent field surveys conducted by the author during which fresh specimens were collected, allowed a precise reassessment and subsequent Manuscript accepted 30.09.2011 724 L. MONOD reinstatement of two species in synonymy, 1.e. Liocheles boholiensis Kraepelin, 1914, and Liocheles neocaledonicus Simon, 1877. Besides, Liocheles australasiae longima- nus (Werner, 1939) is elevated to species rank, while Liocheles australasiae brevidigi- tatus Werner, 1936 is synonymized with Liocheles australasiae australasiae (Fabricius, 1775). Redescriptions, diagnoses, illustrations of important characters and distribution maps are provided for the three re-established taxa. Liocheles boholiensis described by Kraepelin in 1914, was placed into the synonymy of ZL. australasiae by L. E. Koch (1977). However, the species clearly belongs to the waigiensis group. Unlike in most species of the waigiensis group, the anterior area of the carapace 1s almost completely smooth and devoid of a coarse granulation, resembling the carapace of the australasiae group. This feature probably led Koch to erroneously considered the species as a junior synonym of L. australasiae. Liocheles neocaledonicus Was first described as Zschnurus neocaledonicus by Simon (1877). Kraepelin (1894) synonymized it with Hormurus caudicula L. Koch, 1867, but subsequently (Kraepelin, 1914) restored it as a valid species in the genus Hormurus Thorell, 1837. Kopstein (1921) accepted Kraepelin’s decision to split H. caudicula into several distinct species, including Æ. neocaledonicus, and Giltay (1931) cited Kraepelin (1914) as the reference for taxonomic identification of Hormurus species, but contrary to Kopstein he considered the different species of the waigiensis group as subspecies of Æ. caudicula, listing H. caudicula neocaledonicus from central and sou- thern New Caledonia and describing Hormurus caudicula sarasini from the north of the island. Following Karsch (1880) who first synonymized Hormurus with Liocheles, Takashima (1945) recognized L. neocaledonicus and L. sarasini as subspecies of Z. caudicula. L. E. Koch (1977) subsequently treated Æ. caudicula and most of its sub- species (including ZL. caudicula neocaledonicus and L. caudicula sarasini) as junior synonyms of Liocheles waigiensis. The present work confirms that ZL. neocaledonicus is a valid species while ZL. sarasini is a junior synonym of the former. Two subspecies of L. australasiae, 1.e. L. australasiae brevidigitatus and L. aus- tralasiae longimanus were described by Werner in 1936 and 1939, respectively. These taxa remained widely unknown to subsequent authors before Fet (2000) reported them in the Catalog of the scorpions of the world. Significant morphological differences from L. australasiae warrant the elevation of L. australasiae longimanus to species level. However, L. australasiae brevidigitatus cannot be separated from L. australasiae based on morphology and that subspecies is thus abolished. MATERIAL AND METHODS EXAMINATION: Specimens were examined with a ZEISS Stemi SV8 stereo- microscope. Morphological examination is sometimes hampered by mud and soil aggregates adhering to the tegument of specimens and these can obscure fine details. Ultrasonic treatment (sonication) was used to render the specimens free of obstructing particles (Nowak er al., 2008). The specimens were placed for 15 to 90 seconds in a digital ultrasonic cleaner filled with soapy water. This step was repeated until a satis- tactory degree of cleanliness was achieved. For smaller and older specimens more numerous but shorter rounds of sonication were carried out in order to avoid damage. Following sonication, the specimens were rinsed thoroughly with distilled water. LIOCHELES SCORPIONES, LIOCHELIDAE 725 DISSECTION: Mature male specimens were dissected using microsurgical scissors and forceps in order to study their hemispermatophores. Paraxial organ tissue was removed either manually with forceps or chemically with Proteinase K (Qiagen, Venlo, The Netherlands) diluted with water to 50%. Hemispermatophores were emerged in the solution and then placed in an oven at 45-S0°C for 15 minutes to an hour depending on the size and sclerotization of the structure. Once the soft tissue of the paraxial organ was sufficiently digested, hemispermatophore were retrieved from the solution and thoroughly rinsed with water. TERMINOLOGY follows Vachon (1956, 1963) for cheliceral dentition, Stahnke (1970), Stockwell (1989), Sissom (1990) and Hjelle (1990) for pedipalp segmentation, Prendini (2000) for pedipalp carinae patterns, Vachon (1974) for trichobothrial patterns, Couzjin (1976) for leg segmentation, Prendini (2000) for metasomal carinae, and Lamoral (1979) and Monod & Volschenk (2004) for hemispermatophore morpho- logy. Measurements follow Stahnke (1970) and were recorded in mm using an occular micrometer or a digital caliper. ILLUSTRATION: Drawings were produced using a camera lucida mounted on the stereomicroscope. The sketches were subsequently drawn with ink pens and scanned for further processing and editing. Photographs and illustrations were subsequently edited in Photoshop CSS (Adobe Systems, San Jose, CA, USA) (background removal and contrast adjustment) and plates were prepared with Illustrator CSS (Adobe Systems, San Jose, CA, USA). Colour drawings were produced as digital media based on scientific illustrations, photographs of live specimens served as reference to accurately represent the body colours of the animals. MaAPPING: Distribution maps were produced using ArcGIS version 9.3 (Environmental Systems Research Institute, Redlands, CA, USA), by superimposing point locality records on SRTM 90 m (3 arc-second) digital elevation data (Jarvis ef al., 2008) and SRTM 1 km (30 arc-second) global bathymetry dataset (Becker ef al., 2009). Geographical coordinates for records without GPS data were traced by reference to gazetteers, the GEOnet Names Server (http://earth-info.nga.mil/gns/html/index.html), or detailed route maps of various earlier expeditions. MATERIAL LIST: Following Chapman & Grafton (2008), GPS coordinates that may be used by potential collectors seeking to conduct mass harvesting in order to supply the arachnid pet trade were purposely removed from the list of material. These data may be supplied to researchers upon request. ACRONYMS: Depositories of material examined are abbreviated as follows: AMNH, American Museum of Natural History (New York, USA); BPBM, Bernice P. Bishop Museum, Honolulu (Hawaïi, USA); CAS, Californian Academy of Sciences (San Fransisco, USA); LM, Lionel Monod personal collection, to be deposited in the collection of the MHNG; MHNG, Muséum d'histoire naturelle (Genève, Switzerland); MM, Manchester Museum, University of Manchester (Manchester, United Kingdom); MNAN, Muséum national d'Histoire naturelle (Paris, France); MRHN, Musée Royal d'Histoire Naturelle de Belgique (Bruxelles, Belgium); NHML, Natural History Museum (London, United Kingdom); NMB, Naturhistorisches Musem (Basel, 726 L. MONOD Switzerland); NNHM, National Natuurhistorisch Museum (Leiden, The Netherlands); QM, Queensland Museum (Brisbane, Australia); USNM, United States National Museum of Natural History, Smithsonian Institution (Washington, DC, USA); WAM, Western Australian Museum (Perth, Australia); ZMAK, Zoologisches Forschungs- institut und Museum Alexander Koenig (Bonn, Germany); ZMH, Zoologisches Staatsinstitut, Zoologisches Museum Hamburg (Hamburg, Germany). Besides, the following abbreviations were used: DDEE, Direction du Dévelopement Economique et de l’Environnement, Province Nord, Koné, New Caledonia); DRN, Direction des Ressources Naturelles, Province Sud, Nouméa, New Caledonia; IAC, Institut Agro- nomique Néo-Calédonien, Pocqueureux, La Foa; MIS, misidentification. RESULTS Family Liochelidae Fet & Bechly, 2001 Genus Liocheles Sundevall, 1833 Liocheles australasiae (Fabricius, 1775) Liocheles australasiae brevidigitatus Werner, 1936 syn. n. Liocheles boholiensis (Kraepelin, 1914) Figs 1-2, Table 1 Hormurus boholiensis Kraepelin, 1914: 333; Weidner, 1959: 101. Hormurus caudicula boholiensis: Giltay, 1931: 12, 13, 18, 19. Liocheles caudicula boholiensis: Takashima, 1945: 96. Liocheles australasiae: L. E. Koch, 1977: 161 (misidentification). Liocheles boholiensis: Monod, 2000: 72-76, pls 22-24, map 8. DISTRIBUTION RANGE: This species is known only from Bohol Island (Central Visayas), Philippines. However, 1t is most likely present on other nearby Philippine islands. MATERIAL EXAMINED: ZMH, without registration number; 1 ® syntype (here designated lectotype), 1 subadult © syntype (here designated paralectotype); Philippines, Bohol; X.1863. DIAGNOSIS: L. boholiensis can be distinguished from Z. waigiensis by the following characters: (1) prolateral process of pedipalp patella forming 2 distinct spiniform processes (bifid), these sometimes fused but not forming a large median spine; (2) ventral inter- carinal surface of pedipalp chela granular along prolateral and retrolateral edges only, smooth medially; (3) retrolateral ventral intercarinal surface of pedipalp femur smooth or nearly so; (4) ventral intercarinal surface of pedipalp femur with granulation absent or vestigial and limited to the retrolateral edge; (5) ventrosubmedian carinae of meta- somal segment I with 1-3 pairs of well developed medial granules; ventrosubmedian carinae of metasomal segment II with 1 pair of well developed subposterior spiniform granules and 1-3 pairs of well developed medial spiniform granules. DESCRIPTION OF ADULT FEMALE: Coloration: The poor state of preservation of the type specimens did not allowed to accurately describe the coloration of the animals. Ornamentation of cuticle: Non-granular surfaces of mesosoma, metasoma, pro- soma, legs and pedipalp finely punctated. Chelicerae (Fig. 1C): Medial and basal teeth of fixed finger fused into a bicusp. Movable finger: dorsal margin with four teeth (one subdistal and one basal); distal dor- sal tooth smaller than distal ventral tooth; ventral margin smooth. LIOCHELES SCORPIONES, LIOCHELIDAE 127 FIG. 1 Liocheles boholiensis Kraepelin, 1914, female holotype (ZMH). (A) Carapace, dorsal aspect. (B) Left tarsus IV, ventral aspect. (C) Left chelicera, dorsal aspect. (D) Pectines and genital opercula. (E) Metasoma, lateral aspect. Scale lines, 2 mm (A, E), 1 mm (C, D), 0.5 mm (B). 728 L. MONOD TABLE 1] Meristic data for the types of Liocheles boholiensis Kraepelin, 1914. Lectotype, female Paralectotype, subadult female Carapace Length 8.8 6.7 Anterior width 5.8 4.8 Posterior width 9.6 7.6 Metasomal segment I Length 32 22 Width 2.0 1.5 Metasomal segment V Length 4.5 3.6 Width 1.6 1.0 Height 1.6 1.2 Telson vesicle Width 1.8 1.3 Height 19 13 Pedipalp Femur length 8.4 529 Femur width 3.4 2.3 Patella length 8.5 6.2 Patella width 3.4 25 Chela length 17.4 12.0 Chela width 12 4.6 Chela height 3.6 22 Chela movable 8.8 6.0 finger length Total length 53.0 46.0 Pedipalp: Femur as long as or shorter than carapace. Chela virtually asetose. Chela fingers morphosculpture: Denticulate edge even distally, with a double row of primary denticles often fused at the base, with inner accessory denticles (IAD), without enlarged granules. Chela fingers linear, without lobes and notches. Trichobothriotaxy (Fig. 2): Chela manus without accessory trichobothria; Dr tri- chobothrium midpalm or slightly less than midpalm:; four V trichobothria; V; and , separate: Esb trichobothrium more distal than Æ£b group, close to Esf; Eb; trichobo- thrium close to Eb; ;; Est trichobothrium midpalm or nearly so. Chela fixed finger: db trichobothrium on dorsal surface; esb, eb, est, et equidistant (distance est-esb similar to distance esb-eb) or est closer to esb; eb trichobothrium at base of finger, behind point of articulation between fixed and movable fingers, in line with esb-ef axis; esb tricho- bothrium in line with est-eft axis, in proximal region of fixed finger; two i trichobothria. Patella: d, trichobothria distal to patellar spur; ef series with three trichobothria; est series with one trichobothrium: em series with two trichobothria; esb series with two trichobothria; em-esb series with four trichobothria in a single group or in two groups esb} and esb;/em, ;; eb series with five trichobothria in two groups eb} and eb; ; or eb}/eb, ; and eb, ;: three v trichobothria. Carinae: Chela manus: dorsal secondary carina obsolete: digital carina distinct, costate or granular, stronger than external secondary carina; external secondary carinae absent or obsolete; ventroexternal carina granular or crenulate; ventromedian and ventrointernal carinae absent or obsolete; internomedian carina present, granular (at least sparsely). Patella: prolateral process present (distinct), strongly developed, forming two distinct spiniform processes (bifid), these sometimes fused but not LIOCHELES SCORPIONES, LIOCHELIDAE 729 FIG. 2 Liocheles boholiensis Kraepelin, 1914, female holotype (ZMH), pedipalp with trichobothrial pattern. (A) Chela, dorsal aspect. (B) Idem, retrolateral aspect. (C) Idem, ventral aspect. (D) Trochanter, femur and patella, dorsal aspect. (E) Idem, retrolateral aspect. (F) Idem, ventral aspect. Scale line, 2 mm. 730 L. MONOD forming a large median spine; internodorsal and dorsomedian carinae distinct; dorso- external carina absent or obsolete; externomedian carinae granular or costate; ventroexternal carina present and distinct, costate or granular. Femur: dorsoexternal carina present and distinct, at least as a ridge, usually more distinct in proximal half, equally developed than dorsointernal carina; dorsointernal carina present and distinct; internomedian dorsal carina absent, without spines or granules (or with a single basal spine); internomedian ventral carina absent, without spines, or at most vestigial, with a single spine at proximal extremity; ventromedian carina absent or obsolete; ventro- internal carina present, granular. Macrosculpture: Chela fingers granular (at least sparsely in proximal half). Chela fixed finger: area around db/dsb/dst trichobothria granular (at least in the most proximal part), db/dsb/dst trichobothria in a single large smooth depression. Chela: dorsal intercarinal surface entirely and densely granular, with medium-sized spiniform granules; retrolateral intercarinal surface granular; ventral intercarinal surface granular along prolateral and retrolateral edges only, smooth medially; prolateral intercarinal surface at least sparsely granular, granulation less distinct along prolateral ventral edge. Patella: dorsal and ventral intercarinal surfaces entirely granular, at least a reticulated network of granules present; retrolateral intercarinal surface at least sparsely granular; proximal half of prolateral intercarinal surface at least sparsely granular, distal half usually less granular. Femur: dorsal intercarinal surface densely granular except at distal end; dorsal and prolateral intercarinal surfaces with small to medium-sized spiniform granules; retrolateral dorsal intercarinal surface at least sparsely granular; retrolateral ventral intercarinal surface smooth or nearly so; ventral intercarinal surface smooth or with vestigial granulation limited to the retrolateral edge; prolateral inter - carinal surface at least sparsely granular. Carapace (Fig. 1A): Anterior margin with shallow median notch. Anterior furcated suture/sulcus present and distinct. Median ocular tubercle situated anterome- dially, small, occupying about 1/9 of carapace width, at least slightly raised; super- ciliary carinae present, without granules. Median ocelli separated by at least half diameter of median ocellus, at least twice the size of lateral ocelli. Three pairs of lateral ocelli; ocelli equal in size, equidistant, close together, almost touching each other. Margin behind lateral ocellus without spines. Carapace surfaces with minute spiniform granules, evenly and sparsely distributed; anteromedian surface with frontal lobes smooth, granular along median longitudinal sulci and anterior furcated sulci; rest of carapace at least sparsely granular. | Mesosoma: Tergites I-VII with posterior margins straight or nearly so, without distinct prominence; posterior margins of pretergite smooth, without spines of granules; posttergite entirely smooth or nearly so. Posttergites III-VII with distinct reticulated network of ridges and dimples, surface uneven; I-VI with lateral transversal sulcus. Ventral morphology: Anterodistal tip of coxa III without swelling or bulge. Sternum (Fig. 1D) equilateral pentagonal (anterior width slightly greater than posterior width); length less than or equal to posterior width. Genital operculum (Fig. 1D) oval to semi-oval, as wide as long (or wider than long), approximately same width as sternum, sclerites partly fused, median suture distinct; posterior notch present, at least LIOCHELES SCORPIONES, LIOCHELIDAE 731 weakly developed. Pectines (Fig. 1D) short (distal edge not reaching distal edge of coxa of leg IV), with fulcrae and three marginal lamellae; six pectinal teeth, long, straight, covered with sensory papillae only in distal portion. Stigmata (spiracles) half- moon shaped (with a distinct curve), short (less than 1/3 of sternite width). Sternite VII without longitudinal carinae. Metasoma (Fig. 1E) not flattened laterally. Segments I-IV with dorsomedian furrow shallow, weak to absent or only visible on segments I-III; dorsosubmedian carinae absent or obsolete; dorsolateral, ventrolateral and paired ventrosubmedian carinae present and distinct on at least some segments. Segment I: width less than or equal to height; median lateral carina present and distinct; dorsosubmedian carinae with posterior spiniform granules weak to absent (not noticeably larger than preceding granules); dorsomedian posterior spiniform granules weak to absent; ventral surface without posterior spiniform granules; ventrosubmedian carinae with 1-2 pairs of moderate to strong subposterior spiniform granules and 1-3 pairs of well developed medial granules. Segment Il: dorsosubmedian carinae with posterior spiniform granules weak to absent (not noticeably larger than preceding granules); dorsomedian posterior spiniform granules weak to absent; ventrolateral carinae without posterior spiniform granules; ventrosubmedian carinae without posterior spiniform granules, with one pair of well developed subposterior spiniform granules and 1-3 pairs of well developed medial spiniform granules. Segment III: dorsosubmedian carinae with posterior spiniform granules weak to absent (not noticeably larger than preceding granules); ventrolateral carinae with ridges indistinct or weak and smooth; ventro- submedian carinae with ridges indistinct or weak and smooth (sometimes with a few reduced spiniform granules), without subposterior spiniform granules. Segment IV: dorsosubmedian carinae with posterior spiniform granules weak to absent (not noti- ceably larger than preceding granules); ventrolateral carinae with no ridges visible or with weak smooth ridges; ventrosubmedian carinae with no ridges visible or with weak smooth ridges (sometimes with few reduced spiniform granules in posterior half), without subposterior spiniform granules. Segment V: dorsal surface smooth, without smooth shiny depression in posterior half between ventrolateral carinae; dorsolateral carinae absent or obsolete; ventrolateral carinae present and distinct, smooth or nearly so (rarely one pair of vestigial granules posteriorly); ventromedian carina absent or obsolete, ridges absent or weak and smooth; anal arch crenulate, at least with few reduced teeth. Telson as long as or slightly longer than metasomal segment V; vesicle smooth, without granules, unmodified. Legs: Femora I-IV with ventromedian surfaces bicarinate (prolateral carinae often weakly developed). Femur IV with ventromedian carinae vestigial (only expressed distally) or indistinct (only scattered granules). Tibiae I-Il: retrolateral margins Without spiniform macrosetae. Basitarsus I: prolateral margin with 1-4 spini- form macrosetae; retrolateral margin with 1-5 spiniform macrosetae. Telotarsi I-IV (Fig. 1B): two ventrosubmedian rows of secondarily setiform macrosetae; ventro - median row of spinules absent or vestigial; basal spinules present (at least one, usually in a short row); terminal ventromedian spinules absent; ungues shorter than telotarsus. Telotarsi I-IV: prolateral/retrolateral rows with 3/4, 3/4, 3-4/4 and 4/4 spiniform macro- setae respectively. 192 L. MONOD Liocheles longimanus (Werner, 1939) stat. n. Figs 3-9, Table 2 Hormurus australasiae longimanus Werner, 1939: 362. Liocheles australasiae longimanus: Locket, 1997: 331; Fet, 2000: 397; Monod & Volschenk, 2004: 686. Liocheles longimanus: Monod, 2000: 97-101, pls 36-40, map 12. DISTRIBUTION RANGE AND HABITAT: Indonesia, north-western Sumatra (FIG. 3). In humid tropical forests. MATERIAL EXAMINED: ZMAK, alte Trockenpräparate 111: 1 & syntype (here designa- ted lectotype), 1 ® syntype (here designated paralectotype); Indonesia, Sumatra, Sumatera Utara Province, Montes Battak; H. Fruhstorfer. - MNHN, RS 3477; 2 9; Indonesia, Sumatra, Kenandam”?; 111.1913; Buxton?. —- MNHN-RS 3471; 3 d, 5 9%, 4 juv.; Indonesia, Sumatra, Kenandam”?; 1913; Buxton?. - MHNG, Sum-06/11; 1 &; Indonesia, Sumatra, Sumatera Barat Province, Harau Canyon, N of Payakumbuh, 750 m, old secondary forest; 7.VI.2006; P. Schwendinger. - NNHM, without registration number; 1 à ; Indonesia, Sumatra, Sumatera Barat Province, Lubuk Sikaping (Sumatra’s Westkust), 450 m; 1926; E. Jacobson. — CAS, without registration number; 1 &, 1 ©, 1 subadult d; Indonesia, Sumatra, Sumatera Barat Province, Mangani, mine near Kota Tinggi, 700 m; 21.VIL.1983; E. S. Ross. - MNAHN, RS 7417; 1 9,1 juv.; Indonesia, Sumatra, Sumatera Barat Province, Ngalau Kamang Cave, near Bukittinggi; 20.VIL.1979; J. Balazuc. DIAGNOSIS: L. longimanus can be distinguished from Z. australasiae by the following characters: (1) pectinal tooth count slightly higher (Z. longimanus: 7-10 in males and 6-8 in females; L. australasiae: 6-7 males and 5-6 in females); (2) sexual dimorphism pronounced, pedipalps more elongated in males (femur longer than cara- pace); (3) ventral surface of pedipalp chela manus: V3-V4 close together, in proximal part of manus; (4) prolateral process of pedipalp patella forming a single large spine; (5) patella: dorsal intercarinal surface smooth or nearly so, prolateral edge and proxi- mal end weakly granular in some specimens; retrolateral ventral intercarinal surface smooth or nearly so; (6) femur: dorsal intercarinal surface with granulation absent or limited to proximal and retrolateral edges; retrolateral intercarinal surface smooth or nearly so; ventral intercarinal surface with granulation absent or vestigial and limited to retrolateral edge; (7) carapace entirely smooth or nearly so; (8) metasoma: dorso- submedian carinae of segment II with posterior spiniform granules weak to absent (not noticeably larger than preceding granules); ventrosubmedian carinae of segment I with medial granules weak to absent. DESCRIPTION OF ADULT MALE (habitus see Figs 4, SA, B): Coloration: As in figure 4. Pedipalps black to reddish brown. Chelicerae: dorsal surface light brown to yellow; fingers slightly infuscated in some specimens. Carapace dark brown. Tergites dark brown, slightly lighter than carapace. Coxapophyses, sternum, genital operculum, pectines and sternites light brown to yellow. Metasoma dark brown, slightly lighter than carapace. Telson yellow. Legs light brown to yellow (paler than tergites). Ornamentation of cuticle: Non-granular surfaces of mesosoma, metasoma, prosoma, legs and pedipalp finely punctated. Chelicerae: Medial and basal teeth of fixed finger fused into a bicusp. Movable finger: dorsal margin with four teeth (one subdistal and one basal); distal dorsal tooth smaller than distal ventral tooth; ventral margin smooth. Pedipalp: Femur slightly longer than carapace, Chela virtually asetose. LIOCHELES SCORPIONES, LIOCHELIDAE 733 FIG. 3 Localities of Liocheles longimanus (Werner, 1939) on Sumatra. Chela fingers morphosculpture: Denticulate edge even distally, with double row of primary denticles often fused at base, with inner accessory denticles (IAD), without enlarged granules. Chela movable finger: suprabasal lobe well developed, gently rounded dorsally and lacking a sharp conical tooth, wider than high, not overlapping fixed finger; suprabasal lobe and corresponding notch contiguous or at most with a reduced gap; basal lobe absent or at most 2-3 small spiniform granules present. Chela fixed finger: suprabasal notch distinct and deep; suprabasal lobe well developed and conical; basal lobe absent. Trichobothriotaxy (Fig. 6): Chela manus without accessory trichobothria; Df tri- chobothrium midpalm or slightly less than midpalm; four V trichobothria; V; and PV, close together, in proximal part of manus; Esb trichobothrium basal, in same axis as Eb group; Eb; trichobothrium close to £b,.,; Est trichobothrium midpalm or nearly so. Chela fixed finger, db trichobothrium on dorsal surface; esb, eb, est, et equidistant (dis- tance est-esb similar to distance esb-eb) or est closer to esb; eb trichobothrium at base of finger, behind point of articulation between fixed and movable fingers, above esb- et axis,; esb trichobothrium below est-et axis, at base of finger, behind point of articu- lation between fixed and movable fingers; two i trichobothria. Patella: d, trichobothria distal to patellar spur; ef series with three trichobothria; es series with one tricho - bothrium; em series with two trichobothria; esb series with two trichobothria; em-esb series with four trichobothria in two groups esb, , and em, ,; eb series with five trichobothria in two groups eb, and eb, ; or eb,/eb, ; and eb; ;; three v trichobothria. Carinae: Chela manus: dorsal secondary carina obsolete; digital carina distinct, costate or granular, stronger than external secondary carina; external secondary carinae 734 L. MONOD absent or obsolete; ventroexternal carina costate; ventromedian and ventrointernal ca- rinae absent or obsolete; internomedian carina present, granular (at least sparsely or faintly). Patella: prolateral process present (distinct), strongly developed, forming a single large spine; internodorsal and dorsomedian carinae distinct; dorsoexternal carina absent or obsolete; externomedian carinae granular or costate; ventroexternal carina present and distinct, costate or granular. Femur: dorsoexternal carina present and distinct, at least as a ridge, usually more distinct in proximal half, less strongly deve- loped than dorsointernal carina; dorsointernal carina present and distinct; interno- median dorsal carina absent, without spines or granules (or with a single basal spine); internomedian ventral carina vestigial, with two large spines (one proximal and one midway); internomedian carinae oriented parallel to dorsointernal and ventrointernal carinae; ventromedian carina absent or obsolete; ventrointernal carina present. Macrosculpture: Chela fingers granular (at least sparsely in proximal half). Chela fixed finger: area around db/dsb/dst trichobothria smooth (trichobothria not in distinct depressions). Chela: dorsal intercarinal surface with vestigial granulation (at most low granules present) limited to prolateral and retrolateral edges, absent from median part; retrolateral intercarinal surface granular; ventral intercarinal surface without granules; prolateral intercarinal surface at least sparsely granular, granulation less distinct along prolateral ventral edge. Patella: dorsal intercarinal surface smooth or nearly so, prolateral edge and proximal end weakly granular in some specimens; ven- tral, retrolateral intercarinal surface smooth or nearly so; proximal half of prolateral intercarinal surface at least sparsely granular, distal half usually less granular. Femur: dorsal intercarinal surface with granulation absent or limited to proximal and retro- lateral edges, when present, granulation dense; dorsal and prolateral intercarinal surfaces with small to medium-sized spiniform granules; retrolateral intercarinal surface smooth or nearly so; ventral intercarinal surface with granulation absent or vestigial and limited to retrolateral edge; prolateral intercarinal surface at least sparsely granular. Carapace: Anterior margin with shallow median notch. Anterior furcated suture/sulcus present, distinct. Median ocular tubercle situated anteromedially, small, occupying about 1/9 of carapace width, at least slightly raised; superciliary carinae present, without granules. Median ocelli separated by at least half diameter of median ocellus, at least twice the size of lateral ocelli. Three pairs of lateral ocelli; ocelli equal in size, equidistant, close together, almost touching each other. Margin behind lateral ocellus without spines. Carapace entirely smooth or nearly so. Mesosoma: Tergites I-VII with posterior margins straight or nearly so, without distinct prominence; posterior margin of pretergites smooth, without spines of granules. Posttergites I-VI smooth medially, lateral areas at least sparsely granular posteriorly; VII with anteromedian area smooth, lateral and posteromedian areas granular; granular areas of posttergites I-VII with minute spiniform granules, uni - formly and sparsely distributed. Posttergites III-VITI without reticulated network of ridges and dimples, surface even; I-VI with lateral transversal sulcus present. Ventral morphology: Anterodistal tip of coxa III without swelling or bulge. Sternum (Fig. 7A) subpentagonal (anterior width approximately equal to or shightly less than posterior width); length less than or equal to posterior width. Pectines LIOCHELES SCORPIONES, LIOCHELIDAE 735 FIG. 4 Liocheles longimanus (Werner, 1939). Male, dorsal aspect. Reconstruction based on scientific illustrations and photographs of live specimens. Scale line, 5 mm. (Fig. 7A) moderately long (distal edge reaching, but not surpassing distal edge of coxa of leg IV), with fulcrae and three marginal lamellae; 7-10 pectinal teeth, long, straight, entirely covered by sensory papillae. Stigmata (spiracles) half-moon shaped (with a distinct curve), short (less than 1/3 of sternite width). Sternite VII without longitudinal carinae. 736 L. MONOD Metasoma (Fig. 8C): Segments I-V, length similar to or slightly longer than in female (Fig. 8D), not flattened laterally. Segments I-IV with median furrow shallow, weak to absent or only visible on segments I-IIT; dorsosubmedian and dorsolateral carinae absent or obsolete; ventrolateral and paired ventrosubmedian carinae present and distinct on at least some segments. Segment I: width less than or equal to height; median lateral carina present and distinct; dorsosubmedian carinae with posterior spiniform granules weak to absent (not noticeably larger than preceding granules); dorsomedian posterior spiniform granules weak to absent; ventral surface with 1-2 pairs of well developed posterior spiniform granules; ventrosubmedian carinae with 1-2 pairs of moderate to strong subposterior spiniform granules, medial granules weak to absent. Segment IL: dorsosubmedian carinae with posterior spiniform granules weak to absent (not noticeably larger than preceding granules); dorsomedian posterior spini- form granules weak to absent; ventrolateral carinae with well developed posterior spiniform granules; ventrosubmedian carinae with 2-3 pairs of posterior, one pair of subposterior and 1-3 pairs of medial well developed spiniform granules. Segment III: dorsosubmedian carinae with posterior spiniform granules moderate to strong (distinctly larger than preceding granules); ventrolateral carinae with ridges indistinct or weak and smooth; ventrosubmedian carinae with ridges indistinct or weak and smooth (sometimes with a few reduced spiniform granules), without subposterior spiniform granules. Segment IV: dorsosubmedian carinae with posterior spiniform granules moderate to strong (distinctly larger than preceding granules); ventrolateral carinae with no ridges visible or with weak smooth ridges; ventrosubmedian carinae with no ridges visible or with weak smooth ridges (sometimes with few reduced spini- form granules in posterior half), without subposterior spiniform granules. Segment V: dorsal surface smooth, with smooth shiny depression in posterior half between ventro- lateral carinae; dorsolateral carinae absent or obsolete; ventrolateral carinae present and distinct, anterior half smooth or nearly so, posterior half with strong conical tooth:; ventromedian carina expressed only in posterior half, ridges absent or weak and smooth; anal arch crenulate, at least with few reduced teeth. Telson as long as or sligh- tly longer than metasomal segment V; vesicle smooth, without granules, unmodified. Legs: Femora I-IV with ventromedian surfaces bicarinate (prolateral carinae of- ten weakly developed). Femur IV with ventromedian carinae vestigial (only expressed distally) or indistinct (only scattered granules). Tibiae I-Il: retrolateral margins without spiniform macrosetae. Basitarsus I: prolateral margin with 1-4 spiniform macrosetae; retrolateral margin with 1-5 spiniform macrosetae. Telotarsi I-IV (Fig. 8B): two ventrosubmedian rows of secondarily setiform macrosetae; ventromedian row of spinules absent or vestigial; basal spinules absent; terminal ventromedian spinules ab- sent; ungues shorter than telotarsus. Telotarsi I-IV: prolateral/retrolateral rows with 4/4, 4/4, 4/4 and 4/4 spiniform macrosetae respectively. Hemispermatophore (Fig. 9): Distal lamina at least slightly curved, approxima- tely same size as basal part or slightly longer; distal crest absent; single lamellar hook, basal (in basal 1/3 of distal lamella or below, at least less than mid-length); basal extrusion absent; transverse ridge present and distinct, merging with anterior edge abo- ve base of lamellar hook, at approximately same level as base of lamellar hook. Lamella thin, folded only proximally and unfolded to flattened distal extremity (tip and LIOCHELES SCORPIONES, LIOCHELIDAE 734 FIG. 5 Liocheles longimanus (Werner, 1939). (A) Male (MNHN-RS 3471), dorsal aspect. (B) Same, ventral aspect. (C) Female (MNHN-RS 3477), dorsal aspect. (D) Same, ventral aspect. Scale lines, 5 mm. 738 L. MONOD FIG. 6 Liocheles longimanus (Werner, 1939), male (MNHN-RS 3471), pedipalp with trichobothrial pat- tern. (A) Chela, dorsal aspect. (B) Idem, retrolateral aspect. (C) Idem, ventral aspect. (D) Trochanter, femur and patella, dorsal aspect. (E) Idem, retrolateral aspect. (F) Idem, ventral aspect. Scale line, 2 mm. LIOCHELES SCORPIONES, LIOCHELIDAE FIG. 7 739 Liocheles longimanus (Werner, 1939), pectines and genital opercula, ventral aspect. (A) Male (MNEN-RS 3471). (B) Female (MNHN-RS 3471). Scale line, 1 mm. 740 L. MONOD FIG. 8 Liocheles longimanus (Werner, 1939); male (MNHN-RS 3471) (A-C), female (MNHN-RS 3471) (D). (A) Carapace, dorsal aspect. (B) Right tarsus IV, ventral aspect. (C) Metasoma, lateral aspect. (D) Posterior part of metasoma, lateral aspect. Scale lines, 2 mm (A, C, D), 05 mm(B). LIOCHELES SCORPIONES, LIOCHELIDAE 741 Liocheles longimanus (Werner, 1939), male (CAS), left hemispermatophore. (A) In toto, dorsal aspect, Lh (lamellar hook), Tr (transverse ridge). (B) Detail of the capsular region, ental aspect, Ld (distal lobe), Ld (distal lobe). (C) Idem, ventral aspect, La (lamella), Lb (basal lobe). Scale lines, 1 mm. base approximately of same width); longitudinal ‘spine”, accessory hook and accessory lobe absent; lamellar tip at approximately same level as base of lamellar hook, above tip of distal lobe. Distal lobe well developed as a distinct hump, without distinctive hook-like shape, without accessory hook, carinae or crest. Basal lobe well developed (‘spoon’ shaped), merging with accessory anterior basal lobe; distal edge without accessory fold toward ectal part, forming a 90° angle with lamella; basal edge without accessory fold (no groove), forming a 90° angle with lamella. DESCRIPTION OF ADULT FEMALE (habitus see Fig. 5C, D): Same characters as in male except as follows. Pedipalp markedly shorter and bulkier than in male (see Fig. 5). Denticulate edge of chela fingers linear or nearly so (without pronounced lobe and notch). Mesosoma: Posttergites I-VII entirely smooth or nearly so. Ventral Morphology: Genital operculum (Fig. 7B) oval to semi-oval, as wide as long (or wider than long), approximately same width as sternum; sclerites partly 742 L. MONOD TABLE 2 Meristic data for adult males and females of Liocheles longimanus (Werner, 1939). males females Carapace Length 6.8 6.4 6.5 8.2 712 6.9 Anterior width 4.6 4.3 4.5 32 4.7 5.0 Posterior width fée 12 F2 9.0 7 F5 Metasomal Length 2:7 2.6 21 2.8 2.8 PA segment | Width 17 1.6 1.6 2: 1.8 (7 Metasomal Length 4.2 4.2 4.1 4.3 39 4.0 segment V Width {22 12 13 1.6 1,3 l3 Height ES F5 1.5 14 15 1.4 Telson vesicle Width 1.4 15 5 1.6 1.4 1.4 Height 1.4 1.4 1.6 1.8 1.4 1.4 Pedipalps Femur length 8.3 7.4 8.0 7.8 6.8 6.6 Femur width 2:6 29 2.6 31 2.6 2.8 Patella length 8.0 7.6 7.6 8.0 6.8 6.9 Patella width 3.0 au A 36 31 32 Chela length 153 13.6 15.0 15.6 13.6 13.8 Chela width 4.5 4.6 4.2 5.6 4.9 4.8 Chela height 2.8 4 | 2.8 SRE 2.6 2.6 Chela movable 6.6 5.4 6.5 7:1 6.0 6.4 finger length Total length 46 42 42 56 47 46 fusedmedian suture distinct; posterior notch present, at least weakly developed. Six to eight pectinal teeth (Fig. 7B), covered with sensory papillae only in distal portion. INTRA-SPECIFIC VARIABILITY: As in other scorpions, the number of pectinal teeth varies: 7-10 in males and 6-8 in females. Liocheles neocaledonicus (Simon, 1877) Figs 10-16, Table 3 Ischnurus neocaledonicus Simon, 1877: 237, 238. Liocheles neocaledonico: Simon, 1887: 113 (misspelling of species name). Hormurus caudicula: Kraepelin, 1894: 135 (part)(misidentification); Kraepelin, 1901: 272 (part)(misidentification). Hormurus neocaledonicus: Kraepelin, 1914: 330, 332, 334, 335; Lampe, 1917: 201; Weidner, 1959: 101. Hormurus sarasini Kraepelin, 1914: 332, 335, 336 (considered a subspecies of Hormurus caudicula by Giltay, 1931: 13); Weidner, 1959: 101; Forcart, 1961: 49 syn. n. Hormurus caudicula neocaledonicus: Giltay, 1931: 13, 18, 19. Hormurus caudicula sarasini: Giltay, 1931: 13, 18, 19. Liocheles caudicula neocaledonicus: Takashima, 1945: 96. Liocheles caudicula sarasini: Takashima, 1945: 96. Liocheles waigiensis: L. E. Koch, 1977: 166-172, figs 18, 47, 82, 83, maps 9a, b (part)(mis- identification); Kamenz & Prendini, 2008: 11, 43, pl. 134 (misidentification). Liocheles neocaledonicus: Monod, 2000: 102-107, pls 41-45, map 13; Monod, 2005: 5. LIOCHELES SCORPIONES, LIOCHELIDAE 743 FIG. 10 Localities of Liocheles neocaledonicus (Simon, 1877) on New Caledonia. DISTRIBUTION RANGE AND HABITAT: Endemic to New Caledonia, widely distri- buted across the island (Fig. 10). In rocky habitats (scree slopes, boulders) of tropical humid forests. TYPES: The male holotype of Zschnurus neocaledonicus Simon, 1877 could not be found in the collections of the MNHN, it is considered lost. - ZMH, without regis- tration number; ® paratype of /schnurus neocaledonicus (examined); New Caledonia; 20.VIIL.1900; Bougier. —- NMB, 56-a; d lectotype of Hormurus sarasini Kraepelin, 1914 (designated by Forcart, 1961: 49) (not examined); New Caledonia, Tchalabel; V.1911; F. Sarasin & J. Roux. —- NMB, 56-a; 6 4, 27 © paralectotypes of Hormurus sarasini (not examined); same data as for lectotype. —- ZMH, without registration number; 1 d, 5 © and 1 juv. paralectotypes of Hormurus sarasini (examined), Tchalabel; IL.1913; F. Sarasin. OTHER MATERIAL: MNHN, VA 2660; 1 G, 1 %; without locality data; specimens exa- mined by Vachon in 1981.— AMNH, Acc 37523; 1 4,3 %,9 juv.; New Caledonia,; III-IV.1939; L. Macmillan. —- BPBM, without registration number; 1 juv.; J. & M. Sedlacek. - MNEN, without registration number; 19; P. Rougeot. - MNHN, without registration number; 2 9; 14.1.1995, N. & B. Thibaud. - MNHN, RS 0474; 9 G, 11 9, 21 juv.; collection E. Simon. — MNEHN, RS 0488; 1 juv.; Germain. - MNHN, RS 0530; 2 9; M. Bougier; collection Simon n0.1998. —- MNHN, RS 4424; 1 9, 1 juv.; M. Plessis. - MNHN, RS 8253; 1 9. - MREN, without registration number; 1 ©.-— NHML, 1924.11.1.1; 1 9; 1914; P. D. Montague. - QM, S 17097; 1 d; 17.11.1977; B. Jamieson. —- ZMH, without registration number; 1 ©.-— MNHN, RS 8537; 1 juv.; Bogui-Col des Rousettes, Station 26b, 270 m, mesic forest; 4.V.1987; A. Mordan & S. Tillier. - MNHN, RS 3240; 3 4,1 ©, 1 juv.; Bourail; 1902; H. Méray. - AMNH, without registration number; 1 juv.; Col des Rousettes, 490 m, dry forest; 29.V.1987; N. I. Platnick, R. J. Raven. - MNHN, RS 8537; 2 ©, 2 juv.; Col des Rousettes, 500 m, mesic forest; 29.V.1987; S. 744 L. MONOD Tillier. - WAM, 98/1892; 1 9; Col des Rousettes, rainforest; 11.11.1993; M. S. Harvey, N. I. Platnick, R. J. Raven. —- WAM 98/1891; 1 juv.; same data as WAM 98/1892. — LM, without registration number; 2 4,3 ©, 7 juv.; Col des Rousettes, track, 427 m, small patch of rainforest in between niaoulis savannah, in between stones in a scree covered by litter and tree roots, dryer habitat than previous specimens; 21.1X.2004; D. Gaillard, L. Monod. —- AMNH, LP 5544; 1 juv.; Col des Rousettes, track, 427 m, small patch of rainforest in between niaoulis savannah, in between stones in a scree covered by litter and tree roots, dryer habitat than previous specimens; 21.1X.2004; D. Gaillard, L. Monod. - MNHN, RS 8537; 2 9%, 13 juv.; Dothio, Col de Pétchékara, Station 287, 340 m, mesic forest; 27.X.1986; A. & S. Tillier. —- LM, without registration number; 3 4,4 9; Farino, Louis Barbou forest exploitation, 383 m, degraded humid forest, near dry creek, under stones; 7.X.2004; C. Mille, S. Cazeres & T. Nuques (I.A.C. Pocqueureux), L. Monod. —- AMNH, LP 5547; 1 juv.; Farino, Louis Barbou forest exploitation, 383 m, degraded humid forest, near dry creek, under stones; 7.X.2004; C. Mille, S. Cazeres & T. Nuques (I.A.C. Pocqueureux), L. Monod. — LM, without registration number; 2 4,3 9; Farino, Louis Barbou forest exploitation, 477 m, degraded humid forest, under stones; 7.X.2004; C. Mille, S. Cazeres & T. Nuques (I.A.C. Pocqueureux), L. Monod. —- AMNH, LP 5548; 1 juv.; Farino, Louis Barbou forest exploitation, 477 m, degraded humid forest, under stones; 7.X.2004; C. Mille, S. Cazeres & T. Nuques (I.A.C. Pocqueureux), L. Monod. —- QM, S 23332; 1 juv.; Forêt nord, sud de Noumea, Kwa Neie transmitter station, mixed-dry forest; 25.X.1988; T. C. (?) & R. Raven. — MNEHN, RS 8349; 1 juv.; Goro, mesic forest; 3.1X.1975; P. Bouchet. - MNHN, RS 8537; 1 G, 1 juv.; S of Grand Lac, Station 235, 280 m, mesic forest on peridotites; 26.XI.84; À. & S. Tillier, P. Bouchet (MNHNP, Malacologie). —- BPBM, Acc. No. 1979.380; 2 juv.; Hienghene, 10-50m; 14-17.VII1.1979; G. M. Nishida. —- AMNH, AH 4622-4643; 3 4, 17 ©, 2 juv.; Koyaboa Forest (Poindimié), NE central coast, 16/22.X.1983; Saint Louis (Noumea), on the banks of a creek; 16.X1.1979; A. Renévier. —- WAM, 98/1887-8; 1 d, 1 juv.; Ile des Pins, near Grotte de la 3ème, rainforest; 19.111.1993; M. S. Harvey, N. I. Platnick, R. J. Raven. - MNHN, RS 7333; 2 9; La Foa; IX.1976; M. Droin. - MNHN, without registration number; 1%; Grotte Le Cresson, 18.IX.65. — QM, S 39694; 2 ©, 1 juv.; Chute de la Madeleine, 230 m; 12.XI.2000; P. Bouchard, C. Burwell & G. Monteith (9918). —- NHML, 1927.I1.1.2-5; 4 Q ; Mt Arago; P. D. Montague. — USNM, 00753; 1 ©, 1 juv.; Mt Dor, 15 mi from Noumea; 22.X.1944; W. D. Crabb. — LM, without registration number; 3 4,2 9 ; Mt Humbold, near mine, 505 m, degraded primary forest, vine thicket, in between stones in a scree covered by leaf litter and humus and under larger stones; 4.X.2004; L. Monod. —- AMNH, LP 5546; 1 juv.;: Mt Humbold, near mine, 505 m, degraded primary forest, vine thicket, in between stones in a scree covered by leaf litter and humus and under larger stones; 4.X.2004; L. Monod. — QM, without registration number; 1 9; Mt Koghis, 500 m, rainforest, night collecting; 2-3.X1.2002; Burwell, G. Monteith & S. Wright (11088). —- LM, without registration number; 3 4,3 9; Mt Koghis, start of tracks, 440 m, rain- forest, under rocks; 26.1X.2004; D. Gaillard, L. Monod. —- AMNH, LP 5545; 1 juv.; Mt Koghis, start of tracks, 440 m., rainforest, under rocks; 26.1X.2004; D. Gaillard, L. Monod. - AMNH, LP 6224; 2 juv.; Mt Koghis, ca. 20 km N Noumea; 4.XI1.2004; S. Huber. — CAS, without regis- tration number; 1 © ,2 juv.; S. E. slope of Mt Koyaboa (390 m), Poindimié, elevation less than 500 ft, 3.VI.1985, L. Wishmeyer. - MNHN, RS 8537; 1 9, 20 juv.; Mt Mé Ori, SE slope, Station 29 a, 530 m, forêt humide; 7.V.1987; A. Mordan & S. Tillier. - MNHN, without registration number; 1 4,2 9, 1 juv.; Mt Pouédihi forést; 18.1V.1965. - MNHN, RS 8537; 1 juv.; Barrage de la Néaoua (Ouen Sieu), Station 210, 500 m, mesic forest; 20.XI1.1984; A. & S. Tillier, P. Bouchet, M.-P. Triclot (MNHNP, Malacologie). - AMNH, LP 5542; 1 9 ; Road Nouméa-Rivière Bleue Forest Reserve, 177 m, vine thicket, in rock crevices; 17.1X.2004; D. Gaillard & L. Monod. - MNHN, RS 8350; 5 juv.; Oua Tom, 130 m, mesic forest; 16.1X.1978. —- MNHN, RS 8537; 1 ©, 2 juv.; Vallée de la Ouen Nondoué, Station 205, 70 m, Nothofagus forest; 20.X.1984; A. &sS. Tillier, P. Bouchet, M.-P. Triclot (MNHNP, Malacologie). —- NHM, 1924.11.1.6-7; 1 d,1 Q ; Plaine des Lacs, forest, under bark; 23.11.1914; D. Montague. - MNHN, RS 8537; 1 4,1 9, 2 juv.; Rivière Blanche, Parc Cagous, Station 256, 160/170 m, mesic forest; 1.1X.1986; A. & S. Tillier. - MNHN, without registration number; 1 &, 1 ©, 6 juv.; Rivière Bleue Forest Reserve; 17.VII.1965. —- MNHN, without registration number; 3 juv.; Rivière Bleue Forest Reserve; 20.VIIL.1965. —- MNHN, without registration number; 2 juv.; Rivière Bleue Forest Reserve; 21.X1.1995; J. P. Hugot. - MNHN, without registration number; 1 d; Rivière Bleue Forest Reserve, under bark; 17.VII.1965; Stanniloner. - AMNH, without registration number; 1 juv.; LIOCHELES SCORPIONES, LIOCHELIDAE 745 FIG. 11 Liocheles neocaledonicus (Simon, 1877). Male, dorsal aspect. Reconstruction based on scien - tific illustrations and photographs of live specimens. Scale line, 5 mm. Rivière Bleue Forest Reserve, 280 m, wet forest; 21.V.1987; N. IL. Platnick, R. J. Raven. — QM, S 23328; 3 ©, 3 juv.; Rivière Bleue Forest Reserve, N rainforest; 21.V.1987; R. Raven. - WAM, 98/1885-6; 1 4, 1 9 ; Rivière Bleue Forest Reserve, 240 m, rainforest; 9.111.1993; M. S. Harvey, N. L. Platnick, R. J. Raven. — LM, without registration number; 3 4,3 %; Rivière Bleue Forest Reserve, trail between Pont Pérignon and Kaori gèant, 116-169 m, rainforest, under stones and in rock crevices; 18.1X.2004; D. Gaillard, L. Monod. —- AMNH, LP 5543; 1 juv.; Rivière Bleue 746 L. MONOD Forest Reserve, trail between Pont Pérignon and Kaori gèant, 116-169 m, rainforest, under stones and in rock crevices; 18.1X.2004; D. Gaillard, L. Monod. - AMNH, LP 4323; 2 juv.; Rivière Bleue Forest Reserve, Pic du Grand Kaori, 400 m, under dead wood branches; 11.V.2005; J. Murienne. - MNHN, RS 8537; 1 ©, 1 juv.; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250, parcelle VI M, 160 m; 1.VIIL.1986; À. & S. Tillier. - MNEN, RS 8537; 1 juv.; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250 a, parcelle VI H, 160 m; 15.1X.1986; Y. Letocart, A. & S. Tillier. - MNEN, RS 8537; 2 juv.; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250 b, parcelle VI I, 160 m; 2.X.1987; A. & S. Tillier. — MNEHN, RS 8537; 1 juv.; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250 d, par- celle VI I, 160 m, 4.XIL.1987; À. & S. Tillier. - MNHN, RS 8537; 1 juv.; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250 e, parcelle VI J, 160 m; 5.1.1987; A. & S. Tillier. — MNAN, RS 8537; 1 d, 1 juv.; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250 g, parcelle VI O, 160 m, 6.IIL.87; À. & S. Tillier. - MNHN, RS 8537; 1 9; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250 h, parcelle VI O, 160 m,; 6.1V.1987; À. & S. Tillier. — MNAN, RS 8537; 1 juv.; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250 :i, parcelle VI W, 160 m; 30.V.1987; A. Mordan, A. & S. Tillier. - MNHN, RS 8537; 1 d ; Rivière Bleue Forest Reserve, mesic forest/alluvia, station 250 j, parcelle VI G, 160 m; 12.VI.87; À. & S. Tillier. - MNHN, RS 8537; 2 6, 2 juv.; Rivière Bleue Forest Reserve, mesic forest, station 251, parcelle VII V, 170 m; 15.VIIL. 1986; À. & S. Tillier. - MNAN, RS 8537; 2 juv.; Rivière Bleue Forest Reserve, mesic forest, station 251 a, parcelle VII U, 165 m,; 22.IX.1987; À. & S. Tillier. - MNHN, RS 8537; 1 d,1 %, 1 juv.; Rivière Bleue Forest Reserve, mesic forest, station 251 b, parcelle VII O, 170 m; 13.X.1986; À. & S. Tillier. - MNHN, RS 8537; 1 juv.; Rivière Bleue Forest Reserve, mesic forest, station 251 d, parcelle VII U, 170 m; 11.XI1.1986; À. & S. Tillier. - MNHN, RS 8537; 4 juv.; Rivière Bleue Forest Reserve, mesic forest, station 251 e, parcelle VII U, 170 m; 13.1.1987; A. & S. Tillier. - MNEN, RS 8537; 1 4,2 9; Rivière Bleue Forest Reserve, mesic forest, station 251 f, parcelle VII L, 170 m; 12.11.1987; À. & S. Tillier. — MNAHN, RS 8537; 1 juv.; Rivière Bleue Forest Reserve, mesic forest, station 251 g, parcelle VII K, 170 m; 16.111.1987; A. & S. Tillier. - MNEN, RS 8537; 1 d ; Rivière Bleue Forest Reserve, mesic forest, station 251 h, parcelle VII R, 170 m; 14.1V.1987; À. & S. Tillier. - MNHN, RS 8537; 1 4,1 ®, 2 juv.; Rivière Bleue Forest Reserve, mesic forest, station 251 i, parcelle VII G, 170 m; 14.V.1987; A. Mordan, A. & S. Tillier. - MNHN, RS 8537; 1 à, 4 juv.; Rivière Bleue Forest Reserve, mesic forest, station 251 K, parcelle VII, 170 m; 16.VIL.1986; À. & S. Tillier. — MNEN, RS 8537; 2 juv.; Rivière Bleue Forest Reserve, Station 255, Mt Kaala, S slope, dry - forest; 27.VIII.1986; A. Chazeau & S. Tillier. —- QM, S 39696; 1 à, 2 juv.; Rivière Bleue Forest, Kaori géant; 11-12.X1.2000,; P. Bouchard, C. Burwell & G. Monteith (9957). — QM, S 23331; 2 juv.; Rivière Bleue Reserve, near Kaori géant, 120 m; 25.V.1984; G. Monteith & D. Cook. — QM, S 39711; 1 ®, 1 juv.; Rivière Bleue Forest Reserve, Panoramic track; 12.X1.2000; P. Bouchard, C. Burwell & G. Monteith (9951). - QM, without registration number; 1 d ; Rivière Bleue Forest Reserve, Pic du Grand Kaori, 250 m; 16-18.X1.2002; S. Wright (11198). — QM, S 59098; 1 juv.; Rivière Bleue Forest Reserve, Pic du Grand Kaori, 250 m; 29.1.2002; G. B. Monteith (8922). —- QM, S 23339; 1 ©, 3 juv.; Rivière des Pirogues (headwaters), 350-400 m; 22.V.1984; G. Monteith & D. Cook. - MNHN, without registration number; 1 à, 1 juv.; Thi Forest Reserve:"5.VIIL6S. = OM, S 23330: 95 FhPForest Reserve, 150=m: 2 1/VAI08 G. Monteith & D. Cook. - AMNH, without registration number; 2 juv.; Touaourou, under stones; 20.X.1959; B. Malkin. — USNM, 00753; 1 % ; Touaourou; 25.XIL.1960; J. P. E. Morrison (3927, gto VI). —- WAM, 98/1889-90; 1 ?, 1 juv.; 2 km N of Troulala; 14.11.1993; M. S. Harvey, N. I. Platnick, R. J. Raven. DIAGNOSIS: L. neocaledonicus can be distinguished form ZL. waigiensis by the following characters: (1) femur as long as or shorter than carapace length; (2) male pedipalp chela: movable finger with basal lobe present as a low hump with 2-3 larger conical teeth; fixed finger with suprabasal lobe low/reduced; (3) prolateral process of pedipalp patella low, forming two distinct spiniform processes (bifid), these sometimes fused but not forming a large median spine; (4) prosoma with coarse spiniform granules at least on anteromedian surface, rest of prosoma usually with sparse smaller spiniform granules; (5) metasoma: dorsosubmedian carinae of segments III-IV with LIOCHELES SCORPIONES, LIOCHELIDAE 747 Fée 12 Liocheles neocaledonicus (Simon, 1877). (A) Male (MNHN-RS 8537, Rivière Bleue Forest Reserve), dorsal aspect. (B) Same, ventral aspect. (C) Female (MNHN-RS 8537, Rivière Bleue Forest Reserve), dorsal aspect. (D) Same, ventral aspect. Scale lines, 5 mm. moderate to strong posterior spiniform granules (distinctly larger than preceding granules); ventrosubmedian carinae of segment I with 1-2 pairs of strong subposterior spiniform granules and 1-3 pairs of well developed medial granules; ventrosubmedian carinae of segment II with one pair of subposterior and 1-3 pairs of medial well deve- 748 L. MONOD loped spiniform granules; ventrosubmedian carinae of segment III with one pair of well developed subposterior spiniform granules. DESCRIPTION OF ADULT MALE (habitus see Figs 11, 12A, B): Coloration: As in figure 11. Pedipalps black to reddish brown. Chelicerae: dorsal surface light brown to orange, fingers slightly infuscated. Carapace black to dark brown. Tergites black to dark brown, slightly lighter than carapace. Coxapophyses, sternum, genital operculum, pectines and sternites light brown to orange. Metasoma black to dark brown. Telson light brown to orange (paler than metasoma). Legs light brown to orange (paler than tergites). Ornamentation of cuticle: Non-granular surfaces of mesosoma, metasoma, pro- soma, legs and pedipalp finely punctated. Chelicerae: Medial and basal teeth of fixed finger fused into a bicusp. Movable finger: dorsal margin with four teeth (one subdistal and one basal); distal dorsal tooth smaller than distal ventral tooth; ventral margin smooth. Pedipalp: Femur as long as or shorter than carapace. Chela virtually asetose. Chela fingers morphosculpture: Denticulate edge even distally, with double row of primary denticles often fused at base, with inner accessory denticles (IAD), without enlarged granules. Chela movable finger: suprabasal lobe well developed, gently rounded dorsally and lacking a sharp conical tooth, wider than high, not overlapping fixed finger; suprabasal lobe and corresponding notch contiguous or at most with a reduced gap; basal lobe reduced to low hump, with 2-3 larger conical teeth. Chela fixed finger: suprabasal notch distinct and deep; suprabasal lobe present and distinct, low and reduced; basal lobe absent. Trichobothriotaxy (Fig. 13): Chela manus without accessory trichobothria; D trichobothrium midpalm or slightly less than midpalm; four V trichobothria; V; and PV, separate; Esb trichobothrium more distal than ÆEb group, midway between Eb group and sr, Eb; trichobothrium close to Eb,.;; Est trichobothrium midpalm or nearly so. Chela fixed finger, db trichobothrium on dorsal surface; esb, eb, est, et equidistant (distance est-esb similar to distance esb-eb) or est closer to esb; eb trichobothrium at base of finger, behind point of articulation between fixed and movable fingers, above esb-et axis; esb trichobothrium below est-et axis, at base of finger, behind point of articulation between fixed and movable fingers; two i trichobothria. Patella: d, tricho- bothria distal to patellar spur; ef series with three trichobothria; est series with one trichobothrium; em series with two trichobothria; esb series with two trichobothria; em-esb series with four trichobothria in two groups esb;_, and em;_;; eb series with five trichobothria in two groups eb} and eb,., or eb;/eb, ; and eb;, ;; three v tricho - bothria. Carinae: Chela manus: dorsal secondary carina obsolete; digital carina distinct, costate or granular, stronger than external secondary carina; external secondary carinae absent or obsolete; ventroexternal carina granular or crenulate; ventromedian and ventrointernal carinae absent or obsolete; internomedian carina present, granular (at least sparsely or faintly). Patella: prolateral process present (distinct), low, forming two distinct spiniform processes (bifid), these sometimes fused but not forming a large median spine; internodorsal and dorsomedian carinae distinct; dorsoexternal carina LIOCHELES SCORPIONES, LIOCHELIDAE 749 c < « “ fes Crett cer cr £ D HAN CT TT TER ; : Pr S < c € € « RC CCE i LA (ML “° v Ty Le 2 EIG M3 Liocheles neocaledonicus (Simon, 1877), female paratype (ZMH), pedipalp with trichobothrial pattern. (A) Chela, dorsal aspect. (B) Idem, retrolateral aspect. (C) Idem, ventral aspect. (D) Trochanter, femur and patella, dorsal aspect. (E) Idem, retrolateral aspect. (F) Idem, ventral aspect. Scale line, 2.5 mm. 750 L. MONOD absent or obsolete; externomedian carinae granular or costate; ventroexternal carina present and distinct, costate or granular. Femur: dorsoexternal carina present and distinct, at least as a ridge, usually more distinct in proximal half, equally developed than dorsointernal carina; dorsointernal carina present and distinct; internomedian dorsal carina absent, without spines or granules (or with a single basal spine); interno- median ventral carina absent, without spines, or at most vestigial, with a single spine at proximal extremity; ventromedian carina absent or obsolete; ventrointernal carina present, granular. Macrosculpture: Chela fingers granular (at least sparsely in proximal half). Chela fixed finger: area around db/dsb/dst trichobothria granular (at least in the most proximal part), db/dsb/dst trichobothria in three distinct smooth depressions (one around each trichobothrium). Chela: dorsal intercarinal surface entirely and densely granular, with medium-sized spiniform granules; retrolateral intercarinal surface granular; ventral intercarinal surface with granulation present along prolateral and retrolateral edges only, smooth medially; prolateral intercarinal surface at least sparsely granular, granulation less distinct along prolateral ventral edge. Patella: dorsal and ventral intercarinal surfaces entirely granular, at least a reticulated network of granules present; retrolateral intercarinal surface at least sparsely granular; prolateral inter- carinal surface: proximal half at least sparsely granular, distal half usually less granu- lar. Femur: dorsal intercarinal surface densely granular except for distal end; dorsal and prolateral intercarinal surfaces with small to medium-sized spiniform granules; retro- lateral and prolateral intercarinal surfaces at least sparsely granular; ventral intercarinal surface granular proximally, distal part without granulation. Carapace: Anterior margin with shallow median notch. Anterior furcated suture/sulcus present and distinct. Median ocular tubercle situated anteromedially, medium-sized, occupying about 1/7 of carapace width, at least slightly raised; super- ciliary carinae present, with at least few granules. Median ocelli separated by at least half diameter of median ocellus, at least twice the size of lateral ocelli. Three pairs of lateral ocelli; ocelli of equal size, equidistant, close together, almost touching each other. Margin behind lateral ocellus without spines. Carapace entirely granular, at least sparsely; large spiniform granules on anteromedian surface, rest of prosoma usually with smaller spiniform granules, evenly and sparsely distributed. Mesosoma: Tergites I-VII with posterior margins straight or nearly so, without distinct prominence; posterior margin of pretergites smooth, without spines of granules. Posttergites I-VII, with minute spiniform granules, uniformly and sparsely distributed; I-VI entirely granular, at least sparsely in posterior half, granulation absent or vestigial on ridges in some specimens; VII entirely granular, at least sparsely. Posttergites III-VII with distinct reticulated network of ridges and dimples, surface uneven; I-VI with lateral transversal sulcus. Ventral morphology: Anterodistal tip of coxa IIT without swelling or bulge. Sternum (Fig. 14A) subpentagonal (anterior width approximately equal to or slightly less than posterior width); length less than or equal to posterior width. Pectines (Fig. 14A) moderately long (distal edge reaching, but not surpassing distal edge of coxa of leg IV), with fulcrae and three marginal lamellae; 7-12 pectinal teeth, long, straight, entirely covered by sensory papillae. Stigmata (spiracles) half-moon shaped LIOCHELES SCORPIONES, LIOCHELIDAE 751 FIG. 14 Liocheles neocaledonicus (Simon, 1877), pectines and genital opercula, ventral aspect. (A) Male (Æ. sarasini paralectotype, ZMH). (B) Female (4. sarasini paralectotype, ZMH). Scale line, I mm. 792 L. MONOD (with a distinct curve), short (less than 1/3 of sternite width). Sternite VII without longitudinal carinae. Metasoma (Fig. 15C): Segments I-V as long as or slightly longer than in female (Fig. 15D), not flattened laterally. Segments I-IV with dorsomedian furrow shallow, weak to absent or only visible on segments I-III; dorsosubmedian and dorsolateral carinae absent or obsolete; ventrolateral and paired ventrosubmedian carinae present and distinct on at least some segments. Segment I: width less than or equal to height; median lateral carina present and distinct; dorsosubmedian carinae with posterior spiniform granules weak to absent (not noticeably larger than preceding granules); dorsomedian posterior spiniform granules weak to absent; ventral surface without posterior spiniform granules; ventrosubmedian carinae with 1-2 pairs of moderate to strong subposterior spiniform granules and 1-3 pairs of well developed medial granules. Segment Il: dorsosubmedian carinae with posterior spiniform granules weak to absent (not noticeably larger than preceding granules); dorsomedian posterior spini- form granules weak to absent; ventrolateral carinae without posterior spiniform granules; ventrosubmedian carinae without posterior spiniform granules, with one pair of subposterior and 1-3 pairs of medial well developed spiniform granules. Segment IT: dorsosubmedian carinae with posterior spiniform granules moderate to strong (distinctly larger than preceding granules); ventrolateral carinae with ridges indistinct or weak and smooth; ventrosubmedian carinae with ridges indistinct or weak and smooth (sometimes with a few reduced spiniform granules), with one pair of well developed subposterior spiniform granules. Segment IV: dorsosubmedian carinae with posterior spiniform granules moderate to strong (distinctly larger than preceding granules); ventrosubmedian carinae with ridges indistinct or weak and smooth (some- times with few reduced spiniform granules in posterior half), without subposterior spiniform granules; ventrolateral carinae with ridges indistinct or weak and smooth. Segment V: dorsal surface smooth, without smooth shiny depression in posterior half between ventrolateral carinae; dorsolateral carinae absent or obsolete; ventromedian carina absent or obsolete, ridges absent or weak and smooth; ventrolateral carinae present and distinct, smooth or nearly so (rarely one pair of vestigial granules posteriorly); anal arch crenulate, at least with few reduced teeth. Telson as long as or slightly longer than metasomal segment V; vesicle smooth, without granules, un - modified. Legs: Femora I-IV with ventromedian surfaces bicarinate (prolateral carinae often weakly developed). Femur IV with ventromedian carinae vestigial (only expressed distally) or indistinct (only .scattered granules). Tibiae I-Il: retrolateral margins without spiniform macrosetae. Basitarsus I: prolateral margin with 1-4 spini- form macrosetae; retrolateral margin with 1-5 spiniform macrosetae. Telotarsi I-IV (Fig. 15B): two ventrosubmedian rows of secondarily setiform macrosetae; ventro - median row of spinules absent or vestigial; basal spinules present (at least one, usually in a short row); terminal ventromedian spinules absent; ungues shorter than telotarsus. Telotarsi I-IV: prolateral/retrolateral rows with 4/4, 4/4, 5/5 and 5/5 spiniform macro- setae respectively. Hemispermatophore (Fig. 16): Distal lamina at least slightly curved, approxi- mately of same size as basal part or slightly longer; distal crest absent; single lamellar LIOCHELES SCORPIONES, LIOCHELIDAE 753 FIG.. 15 Liocheles neocaledonicus (Simon, 1877). (A) Carapace of female paratype (ZMH), dorsal aspect. (B) Left tarsus IV of male (H. sarasini paralectotype, ZMH), ventral aspect. (C) Metasoma of male (4. sarasini paralectotype, ZMH), lateral aspect. (D) Metasoma of female paratype (Z MH), lateral aspect. Scale lines, 2.5 mm (A), 2 mm (C, D), 0.5 mm (B). 754 L. MONOD a} 7 LU TTL] Fr 1) 41LÉ he LIMIT - ETES F3 :. ww ] POLE gr €” LL “# ae & DA L NS E E E | ur TT FIG 16 Liocheles neocaledonicus (Simon, 1877), male (LM, Mount Koghis), left hemispermatophore. (A) In toto, dorsal aspect. (B) Detail of capsular region, ental aspect. (C) Idem, ventral aspect. Scale lines, 1 mm. hook in basal 1/3 of distal lamella or below; basal extrusion absent; transverse ridge present and distinct, merging with anterior edge above base of lamellar hook, at approximately same level as base of lamellar hook. Lamella thin, folded only proximally and unfolded towards flattened distal extremity (tip and base approximately of same width); longitudinal ‘spine’, accessory hook and accessory lobe absent; lamellar tip below base of lamellar hook and above tip of distal lobe. Distal lobe well developed as a distinct hump, without distinctive hook-like shape, without accessory hook, carinae or crest. Basal lobe well developed (spoon shaped), merging with accessory anterior basal lobe; distal edge without accessory fold toward ectal part, forming a 90° angle with lamella; basal edge without accessory fold (no groove), forming a 135-150° angle with lamella. Book lungs (after Kamenz & Prendini, 2008): Lamellar surface with simple trabeculae. Lamellar distal edges with arcuate bow-like structures. Posterior spiracle edge with chisel-like structures. LIOCHELES SCORPIONES, LIOCHELIDAE 755 TABLE 3 Meristic data for adult males and females of Liocheles neocaledonicus (Simon, 1877). males females Carapace Length FE 6.2 6.9 7.8 6.4 8.6 Anterior width 4.2 3.8 4.8 4.7 3.9 5.4 Posterior width 8.6 7.4 8.0 9.4 de? 9.3 Metasomal Length PA 23 2.4 2.4 2:1 2.5 segment | Width 24 1.9 1.9 2.1 1.9 23 Metasomal segment V Length 4.5 4.2 4.4 4.4 3.6 4.8 Width 15 12 1.4 1, 123 1.8 Height 1.6 1.4 55 1.8 1.4 1.8 Telson vesicle Width 1.6 F3 1.5 1.8 13 1.7 Height 1 1.4 1.6 1.8 1.4 1.8 Pedipalps Femur length 6.5 56 6.5 6.6 52 7.4 Femur width 3.0 2.6 2.9 sf 25 3.4 Patella length 6.8 2 6.8 7.4 5.6 8.1 Patella width 322 ET 3.4 3.4 2.6 4.2 Chela length 14.9 12.8 14.3 15.4 12.0 16.9 Chela width 5.8 4.8 5,3 6.2 4.7 6.4 Chela height 3.2 2.6 3.3 35 23 4.2 Chela movable finger length #7, 6.7 Fa F9 6.2 94 Total length 49 40 ——- 51 42 58 DESCRIPTION OF ADULT FEMALE: Same characters as in male except as follows. Pedipalp: Chela fingers linear or nearly so, without lobes and notches. Ventral morphology (Fig. 14B): Genital operculum oval to semi-oval, as wide as long (or wider than long), approximately same width as sternum; sclerites partly fused, median suture distinct; posterior notch weakly developed. Pectines short (distal edge not reaching distal edge of coxa of leg IV). Six to eleven pectinal teeth covered with sensory papillae only in distal portion. INTRA-SPECIFIC VARIABILITY: In males the lobe on the cutting edge of the movable finger and the corresponding notch on the fixed finger of the pedipalp chela can be more or less developed. Some specimens display very low lobes, while others have more pronounced lobes. Moreover, the fitting between the lobe and the notch is also variable; in most specimens there is no gap between them, or at most a very narrow posterior gap. However, some males have slighty larger posterior gaps. Bigger specimens tend to have a more pronounced lobe and notch, and usually also display larger posterior gaps. The posterior spiniform granules on the dorsosubmedian carinae of metasomal segments III and IV are usually distinctly larger than the preceding granules, however there is no clear size difference in some specimens. Pectinal teeth count variation is as follow: 7-12 in males and 6-11 in females. 756 L. MONOD ACKNOWLEDGEMENTS An extramural training at the MHNG in 1998 was partially funded by the Department of Cultural Affairs of the City of Geneva. I am deeply indebted to the following persons: Wilson Lourenço and Volker Mahnert who supervised the present study; the late Jacqueline Heurtault and Yves Coineau who provided work space and infrastructure at the MNHN; Peter Schwendinger, Bernd Hauser and Charles Lienhard who did the same at the MHNG. I also wish to thank the following persons: (1) for granting access to the collections of their respective institutions, or providing and pre- paring loans: Leon Baert (MRHN); Johnathan Coddington and Dana de Roche (USNM); Hieronymus Dastych (ZMH); Paul D. Hillyard and Janet Beccaloni (NHML); Charles Griswold (CAS); Mark S. Harvey and Julianne Waldock (WAM ); Franz Krapp (ZMAK); Erik Van Nieukerken, B. Van Bekkum-Ansari and Leonne Vermond (NNHM); Lorenzo Prendini, Randy Mercurio and Jeremy Huff (AMNH); Robert Raven, Phil Lawless and Owen Seeman (QM); AI Samuelson (BPBM); (2) for providing helful scientific advice, information, comments and/or support: Alain de Chambrier (MHNG), Danièle Decrouez (MHNG), Yvan Lôbl (MHNG), Dmitri Logunov (MM), Lorenzo Prendini (AMNH), Christine Rollard (MNHN) and Peter Schwendinger (MHNG); (3) for providing bibliographic references: the late N. A. Locket, Victor Fet and Matt E. Braunwalder (Arachnodata); (4) for making the habitus color illustrations: Vladimir Timokhanov; (5) for taking the habitus photographs: Claude Ratton (MHNG); (6) for issuing collecting permits in New Caledonia and/or assisting With the permit application process: Jean-Jérôme Cassan (DDEE), Richard Farman (DRN), Cendrine Meresse (DRN); (7) for assistance and hospitality during field work: Delphine Gaillard, Sylvie Cazeres (TAC), Christian Mille (IAC) and Thierry Nuques (IAC); (7) for their precious advice concerning the realisation of various illustrations: Michèle Bertoncini (MNHN) and Jacques Rebière (MNHN); (8) for kindly reviewing the manuscript and providing useful constructive comments: Peter Schwendinger and Erich Volschenk. REFERENCES BECKER, J. J., SANDWELL, D. T., SMITH, W. H. F., BRAUD, J., BINDER, B., DEPNER, J., FABRE, D., FACTOR, J., INGALLS, S., KIM, S-H., LADNER, R., MARKS, K., NELSON, S. PHARAOH, A., TRIMMER, R., VON ROSENBERG, J., WALLACE, G. & WEATHERALL, P. 2009. 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On two Heptathela species from southern Vietnam, with a discussion of copulatory organs and systematics of the Liphistiidae (Araneae: Mesothelae) ........... LANDRY, Bernard, ROQUE-ALBELO, Lazaro & HAYDEN, James E. À new genus and species of Spilomelinae (Lepidoptera, Pyralidae) from the D D PUS IMANAR ECUAMOR.. . . .. ... 1... un seed eus KADEJ, Marcin & HAVAS, Jirf. A new species of Anthrenus Geoffroy, 1762 (Coleoptera: Dermestidae) from Oman, with a key in related species. MAHNERT, Volker & SCHMIDL, Jürgen. First record of the subfamily Pycnocheiridiinae from South America, with the description of Leptocheiridium pfeiferae gen. n., sp. n. (Arachnida: Pseudoscor- OA) ln ee den ces en House nue Ÿ2 SHI, Li, YANG, Ding & GAIMARI, Stephen D. Four new species from China and Southeast Asia (Diptera, Lauxantidae, Homoneurinae) ....... LÔBL, Ivan. On the Scaphisomatini (Coleoptera: Staphylinidae: Scaphi- AO) OMS PNibpDInes LE... ............,44...4.. 4 en MONOD, Lionel. Taxonomic emendations in the genus Liocheles Sonde al 1535 (Scormiones. Liochelidae).... ........:..:42.4s Pages 599-637 639-649 651-657 659-666 667-693 695-721 723-758 REVUE SUISSE DE ZOOLOGIE Volume 118 — Number 4 SCHWENDINGER, Peter J. & ONO, Hirotsugu. On two Heptathela species from southern Vietnam, with a discussion of copulatory organs and systematics of the Liphistiidae (Araneae: Mesothelae) ........... LANDRY, Bernard, ROQUE-ALBELO, Lazaro & HAYDEN, James E. A new genus and species of Spilomelinae (Lepidoptera, Pyralidae) from the Galapagos Islands; Ecuador re SR EE KADEJ, Marcin & HAVAS, Jiff. À new species of Anthrenus Geoffroy, 1762 (Coleoptera: Dermestidae) from Oman, with a key in related species. MAHNERT, Volker & SCHMIDL, Jürgen. First record of the subfamily Pycnocheiridiinae from South America, with the description of Leptocheiridium pfeiferae gen. n., sp. n. (Arachnida: Pseudoscor- piones: Cheiridudae ns M RETRAIT SHI, Li, YANG, Ding & GAIMARI, Stephen D. Four new species from China and Southeast Asia (Diptera, Lauxaniidae, Homoneurinae) . ...... LÔBL, Ivan. On the Scaphisomatini (Coleoptera: Staphylinidae: Scaphi- diinae)-ot the Philippines: PA MoNOD, Lionel. Taxonomic emendations in the genus Liocheles Sundevall 1833 (Scorbiones IOChENAE) EP REE ERER Er Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX Pages 599-637 639-649 651-657 659-666 667-693 695-721 723-758 PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENÈVE CATALOGUE DES INVERTEBRES DE LA SUISSE, NOS 1-17 (1908-1926) ........ série Fr. (prix des fascicules sur demande) VUE RE PALÉOBIOLOGIB 2:14 Le ui nue cn Echange ou par fascicule Fr. LE RHINOLOPHE (Bulletin du centre d’étude des chauves-souris) ....... par fascicule Fr. THE EUROPEAN PROTURA: THEIR TAXONOMY, ECOLOGY AND DISTRIBUTION, WITH KEYS FOR DETERMINATION fe nee dan un en Ba male svale dues à es à à Fr. CLASSIFICATION OF THE DIPLOPODA D nn as mas se ace atqus ronde denis eus s à Fr. LES OISEAUX NICHEURS DU CANTON DE GENÈVE P. GÉROUDET, C. GUEX & M. MAIRE MEN Cane ChHeures, 108902 lo mins dates e ostee o oare + Fr. CATALOGUE COMMENTÉ DES TYPES D'ECHINODERMES ACTUELS CONSERVEÉS DANS LES COLLECTIONS NATIONALES SUISSES, SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ À LA CONNAISSANCE DES ECHINODERMES ACTUELS ae Sete den ie Lie Dee el re étain de 0 e nine 6 à Fr. RADULAS DE GASTÉROPODES LITTORAUX DE LA MANCHE (COTENTIN-BAIE DE SEINE, FRANCE) RE ONE R MAREDA 62, 1991... 40m soeur dose Fr. GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN: SHELLS AND RADULAS ER MAREDA 1992". une nee 2 dosmeneo ces es eue e Fr. O. SCHMIDT SPONGE CATALOGUE D Pr OUEMROUX-FAUNDEZ & SNL. STONE, 190:p.,.1992 :. ........:... 3,8. #UET: ATLAS DE RÉPARTITION DES AMPHIBIENS ET REPTILES DU CANTON DE GENEVE PME NADINE VV. IMARNERT, 48 p.,1993 4. ....:....,.,......1.:3. FE THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: A DOCUMENTED FAUNAL LIST a ce ne 2 ce an move eue nb Case de saea due Fr. NOTICE SUR LES COLLECTIONS MALACOLOGIQUES DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE nn a = a mn nude à à aténgialata one vus ions e aies de Fr. PROCEEDINGS OF THE XIllth INTERNATIONAL CONGRESS OF ARACHNOLOGY, Geneva 1995 (ed. V. MAHNERT), 720 p. (2 vol.), 1996 ...... FE CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE) erramenta Biooiversiatis DL EORBE, x + 190 p., 1997.....:...............: Fr. CATALOGUE SYNONYMIQUE ET GEOGRAPHIQUE DES SYRPHIDAE (DIPTERA) DE LA REGION AFROTROPICALE (nstrumenta Biodiversitatis W), EH. G. DIRICKX, x +187 p., 1998 ..... ............ FE: A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE WEST PALAEARCTIC REGION (Instrumenta Biodiversitatis WI), S. BOWESTEAD, 203 p., 1999................... Fr. THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE SOKOTRA ARCHIPELAGO (Instrumenta Biodiversitatis IV), B. SCHATTI & A. DESVOIGNES, RE NA ils. Fr. PSOCOPTERA (INSECTA): WORLD CATALOGUE AND BIBLIOGRAPHY (Instrumenta Biodiversitatis V), C. LIENHARD & C. N. SMITHERS, LL BAL LR ce CO RP RO Fr. REVISION DER PALAARKTISCHEN ARTEN DER GATTUNG BRACHYGLUTA THOMSON, 1859 (COLEOPTERA, STAPHYLINIDAE) (1. Teil) (Instrumenta Biodiversitatis VI), G. SABELLA, CH. BÜCKLE, V. BRACHAT D DOME LS LR eme es uenitamee es semis ns si peine Fr. PHYLOGENY, TAXONOMY, AND BIOLOGY OF TEPHRITOID ELIES (DIPTERA, TEPHRITOIDEA) Proceedings of the “3rd Tephritoid Taxonomist’s Meeting, Geneva, 19.-24. July 2004” (Instrumenta Biodiversitatis VII). B. MERZ, vi + 274 p., 2006 Fr. 285. — 35.— 35.— 30.— 40.— 160.— 60.— 70.— 180.— 100.— 100.— e à ps EU 2 o NUE # NS Lo — DUC L4 dre os Us VA +. 50 Er. ARE VU ne a ON LOST ap y b:1 ‘GALL f L TE. f ht LM MORE CPE ECT TT Ur), 44 s'Himanautt 4 Lai tir à ur! £. Her FL" F4 et US RULES | rt 2 4j Th Lu 1 MOI rm" va FHMAEMAT AS ET, rh | ATEN est F a: re AL Le: | PLATE CES DRAC A A0 MOD Lin} à ra non Beer LL Ai ES EN DOTE AS ‘ai FL rasta n US VS Ces CRE PI sANQNEL TA AR : CRUEL re 1 Re s'as dr an: EE VA AA NTM M BASE AU ee AS DUR Vrac Re YA LA tie Ana | Ne bé Vi HIER ‘dei del LP we Me 4 eut 180 nf + Art CHA tt, Ve Véirnr UE LES FOURS (AMAR Due > Le { de: M. ARTE | LE Du. Ha EE LT LL + SaUMTA TT EAN | OPA TBE AN HÉIAS à 02 FOUITIOE LUE 11.1] arte ne L NET TRE "#1 CRRALRSI EE UT. FT LA FAN PS [02 AN PES Vues 4 LATE Li VX en Volume 118 - Number 4 - 2011 Revue suisse de Zoologie: Instructions to Authors The Revue suisse de Zoologie publishes papers by members of the Swiss Zoological Society and scientific results based on the collections of the Muséum d'histoire naturelle, Geneva. 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