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Publication subventionnée par:
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VOLKER MAHNERT Directeur du Muséum d'histoire naturelle de Genève
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Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du Muséum de Genève et de représentants des Instituts de zoologie des universités suisses.
Les manuscrits sont soumis a des experts d’institutions suisses ou étrangères selon le sujet étudié.
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REVUE SUISSE DE ZOOLOGIE 109 (1): 3-8; mars 2002
Smodicinodes schwendingeri sp. n. from Thailand and the first male of Smodicinodes Ono, 1993, with notes on the phylogenetic relationships in the tribe Smodicinini (Araneae: Thomisidae)
Suresh P. BENJAMIN Department of Integrative Biology, Section of Conservation Biology (NLU), University of Basel, St. Johanns-Vorstadt 10, CH-4056 Basel, Switzerland.
Smodicinodes schwendingeri sp. n. from Thailand and the first male of Smodicinodes Ono, 1993, with notes on the phylogenetic relationships in the tribe Smodicinini (Araneae: Thomisidae). - A new species of the enigmatic genus Smodicinodes Ono, 1993 is described from a single male from Thailand. Smodicinodes schwendingeri sp. n. is characterised by the presence of strong, sclerotized tubercles on the prosomal crest and by an oval opisthosoma. Relationships within the tribe Smodicinini are dis- cussed; Smodicinodes Ono, 1993 and Parasmodix Jézéquel, 1966 may be regarded as junior synonyms of Smodicinus Simon, 1895.
Key-words: Smodicinodes — Parasmodix — Smodicinus — Smodicinini — Thomisidae — Araneae — tropical montane forests — Thailand.
INTRODUCTION
The Thomisidae, commonly called crab spiders because of their crab-like appearance and ability to move side-ways, is the sixth largest spider family. It in- cludes 2007 described species in 165 genera (Platnick, 2001), with many more spe- cies remaining to be described. Crab spiders are normally sit-and-wait predators and do not build webs. They are mainly active during the day and, with the help of cryptic colour, large body size, strong front legs and potent venom, are very successful pre- dators. Not surprisingly, they are an important component of terrestrial ecosystems (Riechert, 1974). As predators of agricultural pest, thomisids play an important role in natural pests control (Young & Edwards, 1990; Wise, 1993; Uetz et al., 1999).
However, the exact taxonomic limits of this large family remain an unresolved problem. This may be due to the fact that most taxonomic work is based on Holartic species, although thomisid diversity is predominant in tropical habitats (Benjamin, 2000a). See for example the inference of relationships of thomisids in the study on Misumena vatia by Loerbroks (1984). Furthermore, tropical habitats are in the process of being destroyed and there is an immediate need for intensifying taxonomic studies of tropical species.
Ono (1988) erected the tribe Smodicinini [Original name Smodicini, incor- rectly spelled (ICZN Article 32.5.3.1): the stem of Smodicinus Simon, 1895 is Smo- dicin-, together with the tribe suffix -ini, the taxon name should read Smodicinini] to
Manuscript accepted 13.09.2001
4 S. P. BENJAMIN
accommodate the monotypic genera Smodicinus Simon, 1895 and Parasmodix Jézéquel, 1966, to which he added the genus Smodicinodes in 1993 (Ono, 1993). Smodicinini have a rather peculiar habitus characterized by prosomal modifications. The posterior part of the carapace is raised and sclerotized to from a crest, which is furnished with four to six tubercles with strong seta (Jézéquel, 1966: figs 15, 19; Dippenaar-Schoeman, 1980: figs 6, 7; Ono, 1993: figs 1, 2). These spiders also have a clypeus with distal projections (Fig. 1; Jézéquel, 1966: fig. 19) and the male palpal cymbium has a dorsal outgrowth (do in Figs 3, 4; Jézéquel, 1966: fig. 22a, b). Smodicinini are supposed to live in close association with ants (Lessert, 1943; Dippenaar-Schoeman, 1980; Ono, 1993) but no observations on their behaviour are known.
A recent collection in Doi Chiang Dao Wildlife Sanctuary, Chiang Mai Pro- vince, Thailand, revealed the presence of a new species of this enigmatic genus. Un- fortunately only a single male was collected, making a detailed study of character systems other than genital morphology unpractical. The type species of the genus, S. kovaci Ono, 1993, is from Selangor, West Malaysia, and it is only known from a single female specimen.
In this contribution to thomisid taxonomy a second species and the first male of the monotypic genus Smodicinodes Ono, 1993 is described and phylogenetic implications and the monophyly of Smodicinodes is discussed.
METHODS
Structures were examined in temporary mounts embedded in glycerin. All drawings were made with a Nikon Labophot-2 and a Nikon SMZ-U microscope with drawing tube. The methods are described in detail in Benjamin (2000b). Measurements are in mm. The specimen examined is deposited in the “Museum d’histoire naturelle, Genève” (MHNG). Abbreviations used in the text and figures: AER anterior eye row; ALE anterior lateral eyes; AME anterior median eyes; do dorsal extension of the cymbium; e embolus; MOA-WA anterior width of median ocular area; MOA-L length of median ocular area; pe proximal extension of the cymbium; PER posterior eye row; PLE posterior lateral eyes; PME posterior median eyes; rta retrolateral tibial apophysis; vta ventral tibial apophysis.
TAXONOMY
Smodicinodes Ono, 1993
Type species: By original designation, S. kovaci Ono, 1993, from Selangor, West Malaysia.
Diagnosis: See Ono (1993) and discussion.
Composition: Two species; Smodicinodes kovaci Ono, 1993 and S. schwen- dingeri sp. n.
Phylogenetic relationships: Unknown.
Smodicinodes schwendingeri sp. n. Figs 1-4
Holotype 3: Thailand, Chiang Mai Province, Doi Chiang Dao Wildlife Sanctuary, 1060 m, 1995-1996, leg. S. Gardner (MHNG).
Etymology: The specific name is a patronym in honor of my former lecturer Peter Schwendinger who made the type specimen available for study.
SMODICINODES SCHWENDINGERI SP. N. 5
Fics 1-4
Smodicinodes schwendingeri sp. n. 1. Habitus of male holotype, dorsal view. 2. Male palp, ventral view. 3. Ditto, ectal view. 4. Ditto, dorsal view. Abbreviations: do dorsal extension of cymbium; e embolus; pe proximal extension of cymbium; rta retrolateral tibial apophysis; vta ventral tibial apophysis. Scale line: 0.2 mm (2-4), 1.0 mm (1).
6 S. P. BENJAMIN
Diagnosis: Smodicinodes schwendingeri sp. n. is distinguished from S. kovaci Ono, 1993 by the strong, sclerotized tubercles of the prosomal crest. S. kovaci possesses strong setae surrounded by much less developed tubercles, cf. Ono (1993: fig. 2). S. kovaci has an elongated opisthosoma with parallel borders, whereas in S. schwendingeri sp. n. it is oval. S. schwendingeri sp. n. can be distinguished from Smodicinus coroniger Simon, 1895 by the presence of a tubercle between the AME, by the absences of tubercles between AME and ALE and by the oval opisthosoma. Parasmodix quadrituberculatus Jézéquel, 1966 differs from S. schwendingeri sp. n. by the longer dorsal extension of the cymbium, tapering to a fine, sclerotized hook.
Description: Total length 3.0; prosoma length 1.1, width 0.7; opisthosoma length 1.8. Leg I: femur 1.6, patella 0.6, tibia 1.2, metatarsus 1.0, tarsus 0.4. Colour- ation and markings of the specimen preserved in alcohol: prosoma dark brown, sides darker; eyes surrounded by black rings; ventrally invariably brown; palps dark brown. Opisthosoma dark brown to black, centre yellow, folium as in Fig. 1. Legs unpig- mented except for black lateral markings (possibly green in live specimens). Prosoma and opisthosoma with fine colourless hairs. Prosoma ventrally broad-based, projecting upwards. AER recurved, PER slightly recurved. All eyes on distinct tubercles; ALE and PLE tubercles being the largest. PLE tubercles projecting laterally away from prosoma. AME < PME< PLE < ALE, MOA-WA two-times that of MOA-L. Prosoma with four pairs of projections, the two posterior ones bifurcate, and with a single projection between AME (see arrow in Fig. 1). The projections may have carried strong spines, which were presumably lost in the holotype. Lateral sides of clypeus with a pair of anterior-directed projections (Fig. 1). Legs with a single spine on dorsal femur III and IV, otherwise spineless.
Palp (Figs 2-4): Tibia with stout, blunt ventral tibial apophysis (vta), retro- lateral tibial apophysis (rta) bifurcate. Cymbium modified, with a proximal extension (pe) containing the embolus and a dorsal outgrowth (do); retrolateral tibial apophysis covered by proximal cymbial extension. Tegulum disk-shaped, without apophysis. Embolus with a wide base, tapering, winding half way around tegulum (Figs 2, 3).
Female: Unknown.
Distribution: Known only from the type locality.
Natural history: Collected from a tropical montane forest (label: dense evergreen hill forest). The only other known species of the genus was collected from the internode of a bamboo (Ono, 1993).
DISCUSSION
The descriptions of S. schwendingeri sp. n. and S. kovaci are based on single specimens of the opposite sex. The characters used to distinguish S. schwendingeri sp. n. from S. kovaci may thus be attributed to sexual dimorphism. However, the distance between the two type localities, the different habitats in which the types were found and the distinctive natural histories of the species justify the recognition of a new species. Nevertheless, the relationships of S. schwendingeri sp. n. and S. kovaci should be reassessed after the discovery of either the female of S. schwendingeri sp. n. or the male of S. kovaci.
SMODICINODES SCHWENDINGERI SP. N. 7
The differentiation of the “sister” genera Smodicinus Simon, 1895, Paras- modix Jézéquel, 1966 and Smodicinodes is ambiguous. Likewise, the monophyly of Smodicinodes is unclear. Ono (1993) suggested the following diagnostic characters for Smodicinodes: long maxillae and labium, slender legs, presence of a clypeal tubercle, long opisthosoma. However, none of these characters is apomorphic for Smodicinodes.
Although the available information is minimal, it is reasonable to predict that S. coroniger and P. quadrituberculatus do possess elongated maxillae and labium. Both species have anteriorly flattened chelicerae and a clypeus that slopes forward, similar to individuals of Smodicinodes (Jézéquel, 1966; Dippenaar-Schoeman, 1980). Species of all three genera possess slender legs without spines, except for one or two dorsal spines on femora I-IV (Jézéquel, 1966; Dippenaar-Schoeman, 1980; Ono, 1993). The presences of a clypeal tubercle is also known in P. quadrituberculatus; see Jézéquel (1966). The only remaining relevant character is the elongated opisthosoma. If we consider Smodicinodes to be close to the Tamarini of the subfamily Thomisinae as suggested by Ono (1993), then outgroup comparison indicates that an elongated opisthosoma is plesiomorphic for Smodicinodes. Hence, all three genera of Smodici- nini might be ill-founded taxa.
Thus, Smodicinodes is most probably paraphyletic and should be considered as a junior synonym of Smodicinus. The same can be said for Parasmodix. A single genus including all four species is supported by the following apomorphic characters: modified prosoma with spines; long maxillae and labium; slender legs without spines, except for one or two dorsal spines on femora I-IV; presence of a cymbial dorsal outgrowth (do in Figs 3, 4; Jézéquel (1966: fig. 22)). However, only a phylogenetic analysis would reveal the monophyly of that taxon. Thus, I refrain from effecting the new taxonomic combinations.
ACKNOWLEDGEMENTS
Dr Peter Schwendinger (MHNG) is thanked for critical review of the manus- cript and for providing the interesting specimen for examination. This study was supported by the Swiss National Science Foundation (Grant no. 31-55617.98 to Samuel Zschokke) and the University of Basel.
REFERENCES
BENJAMIN, S. P. 2000a. Epidius parvati sp. n., a new species of the genus Epidius from Sri Lanka (Araneae: Thomisidae). Bulletin of the British Arachnological Society 11: 284- 288.
BENJAMIN, S. P. 2000b. Two new species of the genus Suffasia from Sri Lanka (Araneae: Zodariidae). Revue suisse de Zoologie 107: 97-106.
DIPPENAAR-SCHOEMAN, A. S. 1980. The crab spiders of the southern Africa (Araneae: Thomi- sidae), 2. The genera Pherecydes Pickard-Cambridge, 1883 and Smodicinus Simon, 1895. Journal of the Entomological Society of Southern Africa 43: 327-340.
JEZEQUEL, J. F. 1966. Araignées de la savane de Singrobo (Côte d'Ivoire). V. Note complé-
mentaire sur les Thomisidae. Bulletin du Museum National d'Histoire Naturelle, Paris 37: 613-630.
8 S. P. BENJAMIN
LESSERT, R. DE 1943. Araignées du Congo belge (Troisième partie). Revue suisse de Zoologie 50: 305-338.
LOERBROKS, A. 1984. Mechanik der Kopulationsorgane von Misumena vatia (Clerck, 1757) (Arachida: Araneae: Thomisidae). Abhandlungen und Verhandlungen des Naturwissen- schaftlichen Vereins in Hamburg (Neue Folge) 27: 383-403.
Ono, H. 1988. A revisional study of the spider family Thomisidae (Arachnida, Araneae) of Japan. National Science Museum, Tokyo, 252 pp.
Ono, H. 1993. An interesting new crab spider (Araneae, Thomisidae) from Malaysia. Bulletin of the National Science Museum, Tokyo (Zool.) 19: 87-92.
PLATNICK, N. 2001. Catalog of spiders of the world (CD), by RAVEN, R., version 0.9. The American Museum of Natural History, New York.
RIECHERT, S. E. 1974. Thoughts on the ecological significance of spiders. Bioscience 24: 352- 356.
Uetz, G. W., HALAJ, J. & CADY, A. B. 1999. Guild structure of spiders in major crops. Journal of Arachnology 27: 270-280.
Wise, D. H. 1993. Spiders in ecological webs. Cambridge University Press, Cambridge UK, 328 pp.
YOUNG, O. P. & EDWARDS, G. B. 1990. Spiders in the United States field crops and their potential effect on crop pests. Journal of Arachnology 18: 1-27.
REVUE SUISSE DE ZOOLOGIE 109 (1): 9-16; mars 2002
Molecular identification of an endemic Alpine mammal, Apodemus alpicola, using a PCR-based RFLP method
Brigitte A. REUTTER, Eric PETIT! & Peter VOGEL Institute of Ecology, University of Lausanne, BB, IE-ZEA, CH-1015 Lausanne-Dorigny, Switzerland. E-mail: brigitte.reutter@ie-zea.unil.ch ! Ethology — Evolution — Ecology, University of Rennes I - CNRS (UMR 6552), Campus de Beaulieu, Bat. 25, F-35042 Rennes cedex, France.
Molecular identification of an endemic Alpine mammal, Apodemus alpicola, using a PCR-based RFLP method. - The ability of a PCR-based restriction fragment length polymorphism (RFLP) analysis of the cyto- chrome b (mtDNA) to distinguish Apodemus alpicola from two other Apo- demus species was investigated. The partial sequencing of the cytochrome b allowed the identification of one enzyme as being potentially diagnostic. This was supported by an analysis of 131 specimens previously identified using morphometric and/or allozymic data, indicating that the PCR-based RFLP method provides a rapid and reliable tool for distinguishing A. alpi- cola from its two co-occurring congenerics. The method is applicable to samples taken in the field for ecological studies, and could easily be adapted to the identification of museum samples.
Key-words: Apodemus - mtDNA - PCR - RFLP - identification.
INTRODUCTION
The species Apodemus alpicola Heinrich, 1952 is endemic to the Alps, often being found in sympatry with A. sylvaticus (Linnaeus, 1758) and A. flavicollis (Melchior, 1834). A. alpicola was originally considered as a high-altitude subspecies of A. flavicollis (Heinrich, 1951, 1952) and later described as a new species by Storch and Liitt (1989) based on morphological criteria. A biochemical confirmation was given by Vogel et al. (1991) and Filippucci (1992). In certain regions the overlap of the phenotypes is important (Yoccoz, 1992). Thus, the recognition of A. alpicola as a new species with some intermediate characteristics does not facilitate the identi- fication problem, particularly when juvenile individuals are concerned.
Multivariate skull morphometrics separates 97 % of adult specimens (Reutter et al., 1999). While this technique is indeed a good tool to identify museum material, it does not solve the problem of identifying juveniles and living individuals during
Manuscript accepted 11.09.2001
10 B. A. REUTTER, E. PETIT & P. VOGEL
field studies. Protein electrophoresis has proved to be more useful to distinguish the three Apodemus species without the need to sacrifice individuals and is applicable to young specimens (Reutter er al., 2001). However, it requires to use fresh or frozen blood samples.
Although morphologic (Storch & Liitt, 1989; Spitzenberger & Englisch, 1996; Reutter et al., 1999), karyotypic (Reutter et al., in press) and allozymic studies (Vogel et al., 1991; Filippucci, 1992; Filippucci et al., 1996) have provided a new perspec- tive on the systematics of A. alpicola, it does not solve the problem of the iden- tification of living young animals in the field, a point that is important within the frame of population monitoring or ecological studies. Therefore, a technique is needed which is based on non-destructive sampling, small amounts of biological material, and which leads to a reliable identification of adult as well as young ani- mals. A suitable candidate is the PCR-based restriction fragment length polymor- phism technique (RFLP). This technique consists of three steps. First, a suitable part of the genome is amplified through polymerase chain reaction (PCR). Second, the PCR product is digested using endonucleases, which, for the purpose of species iden- tification, should cut the amplified DNA fragment at different sites in the different species. Third, the digested DNA fragments are separated by electrophoresis and visualised (various staining protocols are available), revealing the restriction patterns. It is worth noting here that this technique requires only minute amount of DNA.
Previous surveys of total mitochondrial DNA variability have revealed that A. sylvaticus and A. flavicollis show much higher inter- than intraspecific variation (Tegelstròm & Jaarola, 1989; Michaux et al., 1996, Michaux et al., 1998). Further- more, a recent analysis of cytochrome b sequences indicated that the divergence between A. alpicola and each of the two other species equals the divergence between A. sylvaticus and A. flavicollis, which is approximately 10 % (Martin et al., 2000). Therefore, owing to the availability of cytochtome b sequences for various Apodemus species (Martin et al., 2000), this gene was chosen to investigate the ability of mito- chondrial DNA polymorphisms to discriminate A. alpicola from the other sympatric species. Such a technique could not only be used in the identification of living animals in the field, but also for museum specimens preserved in ethanol.
MATERIAL AND METHODS
Specimens Examined
Tissues samples (frozen liver or toe-clips) were obtained from a total of 131 specimens (46 A. sylvaticus, 45 A. flavicollis, and 40 A. alpicola) from 15 localities in Switzerland, France, Italy, Germany and Austria. The localities sampled (Fig. 1) cover more or less the range of the three Apodemus species in the alpine region. Sample sizes and locality names are indicated in Tab. 1. All specimens were assigned to species using skull morphology (Reutter et al., 1999) and/or protein electrophoresis (Vogel et al., 1991; Reutter et al., 2001).
MOLECULAR IDENTIFICATION OF APODEMUS ALPICOLA 11
Germany ( { EC ee | DZ NE e RA Sta e SETT A Switzerland \\ a BS beg eae Ae | STE ei AO Me ee rr S4 Et, Nes re CRISI CK SI wee) = \ SS SA ) FI Cn SÌ DÀ) Sl 5 N France sarà In F2 Italy / EX il = ! Fic. |
Geographic distribution of the sampled localities.
TABLE |
A survey of material examined (sampling localities and number of individuals). All specimens were previously identified using morphometric ‘ and/or allozymic © methods.
Locality (map symbol) A. sylvaticus A. flavicollis A. alpicola Switzerland
Gordevio TI (S1) 4 (a)
Prosito TI (S2) 3 (a)
Martigny VS (S3) 4 (a)
Sanetsch VS (S4) | (à) 1 @ 6 (a)
Bourg St. Bernard VS (S5) IA) D) 2,0)
Chur GR (S6) 10 (m) 10 m
Haslital BE (S7) 3 (a) 1 @ 2 (m) Germany
Garmisch (G) 6 (a) Austria
Silbertal (A1) | (a.m) 2 (am) 9 (m)
Hohe Tauern (A2) 5 (m) 7 (m) Italy
Valle d’Aosta (11) 7 (am) 1 (a) 3 (am)
Gran Paradiso NP (I2) 7 (m) 7 (m) 2 (m)
Vinschgau (13) 8 (m) a) 5 (m) France
Morzine (F1) | (m)
Mt. Cenis (F2) 5 (m) 4 (m)
12 B. A. REUTTER, E. PETIT & P. VOGEL
DNA Extraction and Amplification
Genomic DNA was isolated from either liver, heart or kidney preserved in 80% ethanol following a salt/chloroform procedure modified from Miller et al. (1988) but with an additional chloroform/isoamylalcohol (24/1) extraction.
Available sequences (GenBank Accession Nos. AF 159395, AF 159392, and AF 159391) of rodent cytochrome b genes (Martin ef al. 2000) were used to design primers that are specific to the genus Apodemus: CB-AF (5’-ATCAGACACAA- TAACAGCATT-3’) and CB-AR2 (5’-GTTCTACTGGTTGACCTC-3’). These pri- mers allowed the double stranded amplification of an 866 bp-fragment. The 25 ul reaction mixture contained the two primers (1 uM each), 2.5 mM MgCl, 0.25 mM each dNTP, 2.5 ul reaction buffer, 5 ul Q solution (Qiagen), and 1 unit Taq poly- merase (Qiagen). The polymerase chain reaction (PCR) was performed on a thermal cycler UNO-Thermoblock (Biometra) and consisted, after 3 min denaturation, of 35 cycles of 93°C for 45 sec, 47°C for 45 sec, and 72°C for 1 min.
DNA Sequencing
The PCR products of 3 individuals from each of the three species were sequenced in order to identify restriction enzymes that would be potentially diag- nostic, 1.e. with interspecific but no intraspecific variability. PCR products were first purified using the Qiaquick purification kit (Qiagen) according to the manufacturer’ s instructions. The purified products were then sequenced with the Dye Mix 2.0 sequencing kit (Perkin Elmer). The reactions were carried out in a 10 ul volume consisting of 0.5 uM primer, 4 ul Dye mix, and 5.5 ul PCR product (which corres- ponds to 30-70 ng DNA). The sequencing reaction consisted, after 3 min dena- turation, of 25 cycles of 96°C for 20 sec, 50°C for 15 sec, and 60 °C for 4 min. Sequences were then precipitated and run on a 6% polyacrylamide gel on an ABI 373 sequencer (Perkin Elmer). Each PCR product was sequenced using the two primers CB-AF and CB-AR2 in order to sequence both strands. The alignment of sequences and search for restriction sites were carried out using Sequencher 3.0 (Gene Codes Corp.).
Digestion with Restriction Enzyme
The enzyme Spel was identified as being potentially diagnostic to discriminate A. alpicola from the other two species (see Results), and tested on 131 individuals. Each of these samples were analysed with Spe/ according to the following protocol: 10 ul of the amplified DNA were digested in a 25 ul reaction mixture comprising 2.5 units Spel (A//CTAGT) (Life Technologies) and 2.5 ul REACT®4, the digestion buffer, according to the manufacturer’s instructions. The digested samples were subsequently run on a 1% agarose (BioRad) gel (120x200 mm, 150 ml) during 1-2 h at 120 V, and stained with ethidium bromide.
RESULTS
For each of the three species, three individuals taken from alpine populations were sequenced in order to identify restriction sites that are fixed within species and variable between species in the area of sympatry. A site cut by the enzyme Spel was
MOLECULAR IDENTIFICATION OF APODEMUS ALPICOLA 13
found to meet these criteria. Spe/ cleaved the amplified part of the cytochrome b gene at the position 723 into two fragments of 311 and 555 bp for A. alpicola (positions are given according to the standard human mtDNA numbering from Anderson et al., 1981), while A. flavicollis, and A. sylvaticus remained both uncut. Hence, all individuals that showed two fragments on the Spel-gel could be attributed to A. alpicola, and those that showed no fragmentation at all to A. flavicollis or A. sylvaticus (Fig. 2). The PCR-based RFLP protocol presented here indeed correctly discriminated 100 % of the A. alpicola individuals from the two other species in a sample of 131 wood mice previously identified using skull morphology and/or protein electrophoresis.
A. sylvaticus A. flavicollis A. alpicola
Marker
Triin
Spel 500 bp
Fic. 2
Representative examples of fragment patterns after Spe! endonuclease digestion of an 866 bp fragment of the cytochrome b gene. Apodemus alpicola showed two fragments after Spel digestions, A. flavicollis, and A. sylvaticus remained uncut.
DISCUSSION
To our current knowledge, the alpine mouse Apodemus alpicola is the only mammal endemic of the Alps. For promoting its conservation, a better knowledge of its status and ecology is needed. However, any progress is currently impeded by the problem of identification, which concerns the three sympatric species A. sylvaticus, A. flavicollis and A. alpicola. Our initial aim was therefore to provide mammalogists and ecologists with an identification technique based on non-destructive sampling that would permit to discriminate A. alpicola from the other two wood mouse species in
14 B. A. REUTTER, E. PETIT & P. VOGEL
the field. The here presented PCR-based RFLP method fulfil this condition, allowing a reliable identification of A. alpicola.
One potential problem of the method is that, as seen on Fig. 2, a faint band of uncut DNA is visible in all cases. Rather than an excess of DNA, this probably results from the coamplification of a nuclear copy of the cytochrome b, which has lost the restriction site used in this study, together with the targeted original mitochondrial cytochrome b (Zhang & Hewitt, 1996). One may wonder whether such a pattern could be reversed in A. alpicola (a strong band of uncut DNA and hardly visible bands of digested DNA), leading to a pattern that would be difficult to interpret. First, this reversed pattern was never observed in our sample of 40 alpine mice. Second, each cell contains only two copies of each nuclear gene for several thousands copies of mitochondrial DNA. Because the PCR mechanism is actually linear (Rameckers et al., 1997), the ratio of nuclear to mitochondrial copies should be of the same order of magnitude before and after PCR. Hence, the restriction pattern of alpine mice indivi- duals should consistently show a faint band of uncut DNA together with two strong bands of cut DNA.
The main limitation of the RFLP methodology applied to species identification is the possibility that the considered polymorphism is not fixed. This problem can be circumvented by multiplying the number of polymorphisms surveyed. However, each endonuclease added to the protocol also adds costs to the analysis. An alternative is to try to validate the method using samples that are representative of the area where one wishes to apply it. We chose the second option, and the results are rather convincing: 100 % of specimens that originate from 15 localities covering about 75 % of the area of sympatry were correctly discriminated. The method is therefore robust to possible geographic variation, and, because it is based on PCR technology of mtDNA variation, it can be applied to slightly invasive (e.g. toe clipping, ear punch) or non- invasive (e.g. hair follicles) ecological samples (Taberlet et al., 1999), as well as to museum specimens (Thomas ef al., 1990). However, because DNA extracted from museum samples is usually degraded, primers that lead to the amplification of a shorter piece of cytochrome b around the restriction site could be designed to increase the probability of successful amplification.
ACKNOWLEDGEMENTS
We are very grateful to Nelly Di Marco, who did most of the DNA extractions and Nevena Basic, who did a part of DNA amplifications and digestions. We also would like to thank Jiirg Paul Miiller from the Biindner Natur-Museum Chur (Switzerland), Maria Jerabek (Salzburg, Austria), Monika Rier (Götzens, Austria), and Elena Patriarca, Paolo Debernardi (Gran Paradiso Nationalpark, Italy) who provided us a lot of tissue samples. Many thanks also to Karen Parker who corrected the English manuscript.
MOLECULAR IDENTIFICATION OF APODEMUS ALPICOLA 15
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FILIPPUCCI, M. G., STORCH, G. & MACHOLAN, M. 1996. Taxonomy of the genus Sy/vaemus in western Anatolia — morphological and electrophoretic evidence (Mammalia: Rodentia: Muridae). Senckenbergiana biologica 75: 1-14.
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MICHAUX, J. R., Fittppucci, M.-G., LiBois, R. M., Fons, R. & MATAGNE, R. F. 1996. Bio- geography and taxonomy of Apodemus sylvaticus (the woodmouse) in the Tyrrhenian region: enzymatic variations and mitochondrial DNA restriction pattern analysis. Heredity 76: 267-277.
MICHAUX, J. R., LiBors, R., RAMALHINHO, M. G. & Maurois, C. 1998. On the mtDNA restriction patterns variation of the Iberian wood mouse (Apodemus sylvaticus). Comparison with other west Mediterranean populations. Hereditas 129: 187-194.
MILLER, S. A., Dykes D. D. & POLESKY, H. F. 1988. A simple salting out procedure for extracting DNA from human nucleated cells. Nucleic Acids Research 16: 1215.
RAMECKERS, J., HUMMEL, S. & HERRMANN, B. 1997. How many cycles does a PCR need? Determinations of cycle numbers depending on the number of targets and the reaction efficiency factor. Naturwissenschaften 84: 259-262.
REUTTER, B. A., HAUSSER, J. & VOGEL, P. 1999. Discriminant analysis of skull morphometric characters in Apodemus sylvaticus, A. flavicollis, and A. alpicola (Mammalia; Rodentia) from the Alps. Acta Theriologica 44: 299-308.
REUTTER, B. A., BRUNNER, H. & VOGEL, P. 2001. Biochemical identification of three sympatric Apodemus species by protein electrophoresis of blood samples. Mammalian Biology 66: 84-89.
REUTTER, B. A., Nova, P., VOGEL, P. & ZIMA, J. 2001. Karyotypic variation between wood mouse species: banded chromosomes of Apodemus alpicola and A. uralensis. Acta Theriologica (in press).
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STORCH, G. & LUTT, O. 1989. Artstatus der Alpenwaldmaus, Apodemus alpicola Heinrich, 1952. Zeitschrift für Säugetierkunde 54: 337-346.
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16 B. A. REUTTER, E. PETIT & P. VOGEL
VOGEL, P., MADDALENA, T., MABILLE, A. & PAQUET, G. 1991. Confirmation biochimique du statut spécifique du mulot alpestre Apodemus alpicola Heinrich, 1952 (Mammalia, Rodentia). Bulletin de la Société Vaudoise des Sciences Naturelles 80: 471-481.
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REVUE SUISSE DE ZOOLOGIE 109 (1): 17-22; mars 2002
Proportionsänderungen beim M, im Gebiss des mitteleuropàischen Dachses Meles meles (Mammalia, Carnivora)
Beatrice BLOCHLINGER & Peter LUPS Naturhistorisches Museum der Burgergemeinde Bern, Bernastrasse 15, CH-3005 Bern, Schweiz.
Changing proportions in M, in the European badger, Meles meles (Mammalia, Carnivora). - We measured M, trigonid- and talonid-length in 62 recent and 110 neolithic badger Meles meles mandibles with fully erupted teeth from the Swiss midlands. The results show an increase in the size of the cutting trigonid and a decrease in the length of the talonid within the last 5000 to 6000 years. This is somewhat in contrast to the overall pattern of evolution of these teeth in badgers (Melinae) in general and within the genus Meles in particular.
Key-words: badger - teeth - Swiss midlands - neolithic - recent - food.
EINLEITUNG
Der erste untere Backenzahn M; der Carnivora besteht in der Regel aus zwei Teilen, einem vorderen, schneidenden (Trigonid) und einem hinteren, mahlend/ kau- enden (Talonid). Das Längenverhältnis zwischen diesen beiden Teilen des M, ermöglicht Hinweise auf die Nahrung einer Tierart: je mehr das Talonid dominiert, desto stärker scheint sich eine Art von der reinen oder dominierenden Aufnahme von Fleisch entfernt zu haben (van Valkenburg, 1989). Bei den Melinae hat sich das Verhältnis von Trigonid zu Talonid im Verlauf der Evolution deutlich zugunsten des Talonid verschoben (Petter, 1971). Auch für die relativ kurze Zeitspanne von maximal 8000 Jahren seit dem frühen Neolithikum lassen sich solche Anpassungen des Gebisses in Richtung “Allesfressertum” feststellen, wie Degerbgl (1933) bei Dachsen aus Dänemark zeigen konnte.
Analog zu Degerbgl bei den dänischen Dachsen, fanden Grundbacher er al. (1990) bei Dachsen aus dem schweizerischen Mittelland für rezente Tieren grössere Werte in mehreren Skelett-, Schädel- und Zahnmassen als für solche aus der Jung- steinzeit (Seeufer-Siedlung Twann). Eine solche Längenzunahme im Verlaufe von rund 5500-6000 Jahren ist auch bei der oberen Prämolarenreihe zu beobachten. Keine Längenzunahme dagegen weist der untere Molar M, auf. Da der P* mit dem M,- Trigonid ein gut harmonierendes Kauwerkzeug bildet, stellt sich die Frage nach allfalligen Verinderungen der prozentualen Anteile von Trigonid und Talonid unter der Voraussetzung des gleich lang gebliebenen Zahnes.
Manuskript angenommen 28.08.2001
18 B. BLOCHLINGER & P. LUPS
Beim Vorliegen solcher Veränderungen Könnten auf der Basis der Kenntnis der heutigen ökologischen Stellung des Dachses allenfalls Hinweise auf seine damalige Ernährungssituation gewonnen werden. Zur Klärung dieser Fragen werden hier die Resultate von Zahnmessungen an jungsteinzeitlichen und rezenten Dachsen verglichen.
BEARBEITETE ZÄHNE
Folgende Unterkiefer aus dem bernischen Mittelland (Raum Biel-Neuenstadt- Bern-Thun-Langnau-Burgdorf) wurden bearbeitet: l. Intakte Kiefer rezenter, als Unfallopfer und Hegeabschüsse in den Jahren 1967-1982 gesammelter Dachse (Sammlung Naturhist. Museum Bern, Lüps, 1984): - je 12 d und © im Alter von 6 - 12 Monaten, d.h. Tiere mit völlig entwickeltem, aber noch wenig abgenutztem Gebiss (Lüps & Roper, 1988a) - 21 d und 17 ®, mindestens 13 Monate alt (vgl. Lüps er al., 1987, Grundbacher et al., 1990)
2. Unterkiefer aus zwei neolithischen Stationen: - Seeberg Burgäschisee-Süd (ca. 3750 - 3700 v. Chr., Schibler & Suter, 1990): 16 Unterkieferäste (s. Jéquier, 1963) - Twann (ca. 3840 - 3530 v. Chr., Suter & Schifferdecker, 1986): 94 Unterkieferäste (vgl. Grundbacher er al., 1990). Dieses Fundgut war durch die Archäologen in drei Gruppen, entsprechend den gefundenen Schichtpaketen, unterteilt worden.
Die Distanz (Luftlinie) zwischen den beiden neolithischen Siedlungen beträgt 40 km, diejenige zwischen Twann und Worb, dem geographischen Herkunfts-Zen- trum der rezenten Tiere, 35 km.
METHODEN
Messungen (Abb. 1): Gesamtlänge des M,, je die auf der buccalen Seite des Zahnes gemessene Länge von Trigonid und Talonid, und die grösste Breite dieser beiden Bereiche. Die buccale Seite wurde gewählt, weil bei alten Tieren mit stark abgekautem M; die Trennung zwischen Trigonid und Talonid lingual kaum mehr erkennbar ist, und weil die Messung von buccal bei rezenten Tieren bei intakter Unterkiefer-Oberkieferverbindung besser durchführbar ist. (Vergleichende Messun- gen auf der lingualen und buccalen Seite des Zahnes haben gezeigt, dass die Resultate in Bezug auf die vorgelegte Fragestellung betreffend Veränderungen der Talonid- und Trigonidlänge nur unwesentlich voneinander abweichen). Jeder Zahn wurde mit einer Schieblehre (Digitalanzeige: 1/100 mm) zweimal gemessen (für die Auswertung wurde der Durchschnittswert verwendet).
Bei den Schädeln der rezenten Tiere wurde durch Zufall festgelegt, welche Kieferhälfte vermessen werden sollte. Bearbeitet wurden die Daten aller neolithischen Kiefer, auch wenn bei einigen linken und rechten Mandibeln nicht ausgeschlossen werden konnte, dass sie von ein und demselben Tier stammen.
Die statistischen Berechnungen wurden mit Hilfe der Statistikprogramme Systat/Sygraph durchgeführt. Für die angewandten t-Tests wurde die Signifikanz- schwelle bei p<0.05 festgelegt.
Miliz PROPORTIONSÄNDERUNGEN BEI MELES MELES 19
ABB. |
Linke Vorbacken- und Backenzahnreihe (Ansicht buccal) eines rezenten männlichen Dachses, mit eingezeichneten Messstrecken am M..
RESULTATE
Weder bei den Zähnen der adulten, noch bei denjenigen der 6 bis 12 Monate alten rezenten Dachse liess sich bei einem der fünf Masse ein Geschlechtsdimor- phismus nachweisen. Somit sind die Voraussetzungen gegeben, männliche und weib- liche Tiere zusammenzufassen, und sie mit den neolithischen, deren Zusammen- setzung hinsichtlich Geschlecht nicht bekannt ist, zu vergleichen. Das Fehlen von Unterschieden zwischen den Zahnmassen juv. und ad. Tiere zeigt, dass beide Serien als Gesamtheit den neolithischen Mandibeln, deren Alterszusammensetzung nicht bekannt ist, gegenübergestellt werden kann.
Signifikante Unterschiede ergaben sich zwischen den Durchschnittswerten der neolithischen und der rezenten Dachse bei der Länge des Trigonids (Zunahme 2.7 %), der Breite des Talonids (2.4 % breiter) und der Länge des Talonids (5.7 % kürzer als bei den neolithischen).
DISKUSSION
Das Fehlen von Unterschieden zwischen den Geschlechtern in allen fünf untersuchten Massen entspricht früheren Befunden am M, (Gesamtlänge, grösste Breite und Hohe: Lüps & Roper, 1988a; Gesamtlänge und Breite: Grundbacher et al.,
20 B. BLOCHLINGER & P. LUPS
TAB. |
Messwerte am M, fiir neolithische und rezente Dachse in mm. (Student T-Test für unabhängige Stichproben)
Dachse Dachse p Zunahme/Abnahme neolithisch rezent % Gesamtlinge n 94 62 0.136 - x 16.45 16.27 S 0.70 0.78 Trigonid Lange n 92 62 0.025 ee x 8.26 8.49 S 0.69 0.45 Trigonid Breite n 93 62 0.165 - x 5123 5:29 S 0.31 0.23 Talonid Länge n 92 62 0.000 -5.5 x 9.01 8.52 S 0.63 0.54 Talonid Breite n 94 62 0.032 +2.4 x 7.40 7.58 N 0.52 0.48
1990). Männliche Dachse aus dem schweizerischen Mittelland sind zwar schwerer und grösser als die Weibchen (auch in den meisten Schädelmassen), Hinweise auf eine unterschiedliche Ernährungsweise der Geschlechter konnten aber bisher weder durch Untersuchungen am Gebiss (Lüps & Roper, 1988b) noch mittels Nahrungs- analysen (vgl. Stocker & Lüps, 1984) gewonnen werden.
Dachse sind seit dem Neolithikum grösser geworden (Daten für Dänemark: Degerbol, 1930, für das Schweizerische Mittelland: Clutton-Brock, 1990, Grund- bacher et al., 1990). Für den M, liess sich nur bei den dänischen Dachsen eine Zunahme nachweisen. Bei den schweizerischen Dachsen hat sich lediglich das Verhältnis zwischen Trigonid und Talonid verändert. Sowohl die zahlenmässig kleine und geographisch heterogene Serie Clutton-Brocks (neolithisch: Yvonand/Lac de Neuchatel, 45 km westlich von Twann, rezent: Frankreich) wie auch die hier vor- gelegten Daten belegen eine Abnahme der mittleren Talonid-Länge und eine leichte Zunahme der Talonid-Breite und der Trigonid-Länge.
Dieser Befund wirft Fragen hinsichtlich der Evolution des Dachs-Gebisses auf, die auf Kosten des P* zu einem grossflächigen M! geführt hat (Petter, 1971, Barysh- nikov & Potapova, 1990). Der Metaconus des reduzierten P* steht dem M, -Trigonid
M, - PROPORTIONSANDERUNGEN BEI MELES MELES 2]
gegenüber, der Metaconus des M! opponiert zum Talonid des M,. Zu dieser länger- fristigen Entwicklung und derjenigen in viel kürzerer Zeitspanne in Dänemark (rund 10°000 Jahre, vgl. Kurten, 1967) steht der hier dargestellte Schritt für die letzten 5-6000 Jahre in Widerspruch. Den Verhältnissen im Oberkiefer (Zunahme der Prä- molarenreihe, Abnahme der M!-Länge) entsprechen die Verhältnisse am M,: Zu- nahme des Trigonids und Abnahme des Talonids.
Das heute aus vielen Teilen Europas bekannte Bild des Dachses als ein an das Leben in der Landwirtschaftszone gut angepassten Nahrungsopportunisten, mit einer hohen Bedeutung von Regenwürmern und pflanzlicher Nahrung, passt schlecht zu den anhand der Trigonid-Zunahme zu postulierenden Zunahme der Carnivorie.
Es bestehen verschiedene Erklärungsmöglichkeiten für diesen scheinbaren Widerspruch. 1) durch ein im Neolithikum noch stärker ausgeprägters Alles- oder Pflanzenfressertum als dies heute der Fall ist. 2) muss berücksichtigt werden, dass das zwischen 1975 und 1999 gewonnene Bild der Dachs-Nahrung im schweizerischen Mittelland (vgl. Stocker & Lüps, 1984, Ferrari, 1997, Fischer, 1997) aus einer Zeit stammt, in welcher der starke Strukturwandel in der Landwirtschaft bereits voll im Gange war (z.B. Verdreizehnfachung der Anbaufläche von Futtermais von 1955 bis 1990). Dieses intensive Ausnutzen eines durch die Landwirtschaft geprägten Angebotes könnte durchaus eine Frage von weniger als 30 Dachs-Generationen sein. 3) ein Hinweis auf eine zunehmende Carnivorie mag auch aus der durch den Ver- gleich von Alveolarmassen postulierten Grössenzunahme des Caninus zu entnehmen sein (Grundbacher er al., 1990). Bei einer solchen Interpretation gilt es aber zu berücksichtigen, dass der Caninus nicht nur in Bezug auf die Ernährung, sondern auch unter dem Aspekt des bei Fähen und Rüden unterschiedlichen Territorial- und Sexual- verhaltens zu betrachten ist (Lüps & Roper, 1988a).
Die Resultate weisen aber auch darauf hin, wie vorsichtig mit verallge- meinernden Aussagen wie „Grössenzunahme“, „Tendenz zum Allesfresser“ u.s.w. umgegangen werden muss: 1) Grössenveränderungen erfolgen oft nicht proportional für alle Körperteile; 2) das in den letzten Jahren gewonnene Bild einer Tierart muss nicht für Jahrzehnte oder Jahrhunderte Gültigkeit haben.
LITERATUR
BARYSHNIKOV, G. F. & PoTAPOVA, O. R. 1990. Variability of the dental system in Badgers (Meles, Carnivora) of the USSR Fauna. Zoologiceskij Zurnal 69: 84-97. (Russisch, englische Zusammenfassung).
CLUTTON-BROCK, J. 1990. Animal remains from the neolithic lake village site of Yvonand IV, Canton de Vaud, Switzerland. Archives des Sciences, Geneve 43: 1-97.
DEGERBOL, M. 1933. Danmarks Pattedyr i Fortiden i Sammenligning med recente Former I (Oversigt; Rovdyr (Carnivora)). Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kobenhavn 96: 357-615.
FERRARI, N. 1997. Eco-éthologie du blaireau européen (Meles meles L., 1758) dans le Jura suisse: comparaison de deux populations vivant en milieu montagnard et en milieu cultivé de plaine. These, Université de Neuchâtel, VI + 210 pp. (unpubliziert).
FiscHER, C. 1997. Ecologie alimentaire et occupation spatiale du blaireau européen (Meles meles) dans un milieu dominé par l’agriculture intensive. Travail de Diplöme, Univer- site de Neuchâtel, VII + 122 pp. (unpubliziert).
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GRUNDBACHER, B., LÜPS, P. & NUSSBAUMER, M.A. 1990. Osteometrische Untersuchungen an neolithischen Dachsen (Meles meles) aus Twann (Kanton Bern, Schweiz) (pp. 101- 113). In: SCHIBLER, J., SEDLMEIER, J. & SPYCHER, H. (Hrsg.). Festschrift fiir Hans R. Stampfli. Helbing & Lichtenhahn, Basel, XVI + 325 pp.
JEQUIER, J.-P. 1963. Der Dachs, Meles meles (Linné) 1758 (pp. 39-43). In: BOESSNECK, J., JEQUIER, J.P. & STAMPFLI, H.R. Seeberg, Burgäschisee-Süd. Die Tierreste. Acta bernensia II (3), Stämpfli, Bern, 215 pp. + XXII.
KurTEN, B. 1967. Some quantitative approaches to Dental Microevolution. Journal of Dental Research 46: 817-828.
Lups, P. 1984. Gewichtsschwankungen beim Dachs (Meles meles L.) im bernischen Mittelland, nebst Bemerkungen zu seiner Biologie. Jahrbuch des Naturhistorischen Museums Bern 8: 273-289.
Lups, P., GRAF, M. & KAPPELER, A. 1987. Möglichkeiten der Altersbestimmung beim Dachs Meles meles (L.). Jahrbuch des Naturhistorischen Museums Bern 9: 185-200.
Lups, P. & Roper, T. J. 1988a. Tooth size in the European badger (Meles meles) with special reference to diet and sexual dimorphism. Acta theriologica 33: 21-33.
Lürs, P. & Roper, T. J. 1988b. Bemerkungen zum Gebissgebrauch beim Dachs. Mitteilungen der Naturforschenden Gesellschaft in Bern, Neue Folge 45: 147-157.
PETTER, G. 1971. Origine, phylogénie et systématique des blaireaux. Mammalia 35: 567-597.
SCHIBLER, J. & SUTER, P. J. 1990. Archäozoologische Ergebnisse datierter neolithischer Ufer- siedlungen des schweizerischen Mittellandes (pp. 205-240). In: SCHIBLER, J., SEDL- MEIER, J. & SPYCHER, H. (Hrsg.). Festschrift fiir Hans R. Stampfli. Helbing & Lichten- hahn, Basel, XVI + 325 pp.
STOCKER, G. & Lups, P. 1984. Qualitative und quantitative Angaben zur Nahrungswahl des Dachses Meles meles im Schweizerischen Mittelland. Revue suisse de Zoologie 91: 1007-1015.
SUTER, P.J. & SCHIFFERDECKER, F. 1986. Das Neolithikum im schweizerischen Mittelland (pp. 34-43). In: Chronologie, Archäologische Daten der Schweiz. Antiqua 15, Schwei- zerische Gesellschaft für Ur- und Frühgeschichte, Basel, 241 pp.
VALKENBURGH, B. VAN 1989. Carnivore Dental Adaptations and Diet. /n: A Study of Trophic Diversity within Guilds (pp. 410-436). Jn: GITTLEMAN, J.L. (ed.). Carnivore Behavior, Ecology and Evolution. Chapman & Hall, London, XIV + 620 pp.
REVUE SUISSE DE ZOOLOGIE 109 (1): 23-46; mars 2002
A revision of the genus Heteroparasitus new status, with the description of Heteroparasitus (Medioparasitus) athiasae subgen. n., sp. n. from Spain and with a key to the genera of Pergamasinae (Acari, Gamasida, Parasitidae)
Ilinca JUVARA-BALS Museum of Natural History, CP 6434, CH-1211 Geneva 6, Switzerland.
A revision of the genus Heteroparasitus new status, with the descrip- tion of Heteroparasitus (Medioparasitus) athiasae subgen. n., sp. n. from Spain and a key to the genera of Pergamasinae (Acari, Gama- sida, Parasitidae). - The subgenus Heteroparasitus Juvara-Bals, 1976 is redefined and raised to genus rank. Heteroparasitus (Medioparasitus) athiasae subgen. n., sp. n. from Spain is described. Taxonomic problems concerning the genera Leptogamasus Trigardh, 1936, Paragamasus Hull, 1918 and Ologamasiphis Athias-Henriot, 1971 are discussed. The genus Ologamasiphis is divided into two subgenera Ologamasiphis s. str. and Holzmannia subg. n. Valigamasus Karg, 1993 syn. n. is a junior objective synonym of Ernogamasus Athias-Henriot, 1971. A key to the genera of Pergamasinae Juvara-Bals is presented.
Key-words: Acari - Gamasida - Parasitidae - Pergamasinae - Hetero- parasitus - new subgenera - new species - taxonomy - key.
INTRODUCTION
The genus Holoparasitus Oudemans, 1936, widely distributed in the Palearctic region, presently includes 32 species. Additional species are likely to be found in areas of the northern hemisphere, which have not been adequately investigated.
Holoparasitus Oudemans was studied and revised by Oudemans (1936), Micherdzinski (1969), Holzmann (1969), Karg (1971, 1993), Juvara-Bals (1975) and Hyatt (1987). Holzmann (1969) erroneously used the name Ologamasus Berlese 1906 instead of Holoparasitus and recognized two subgenera: Ologamasus s. str. and Ologamasiphis Holzmann, the latter with dorsal and ventral shields separated in females. She included two species in Ologamasiphis, i.e.: Ologamasus rotulifer Willmann, 1940 and the new Ologamasus minimus Holzmann, 1969, but failed to designate a type species.
Micherdzinski (1969) followed Oudemans in using the name Holoparasitus and divided this genus into three species groups, i.e: the H. calcaratus group, the H. pollicipatus group and the Ologamasiphis group. He agreed with Holzmann’s sub-
Manuscript accepted 24.08.2001
24 I. JUVARA-BALS
genera but suggested that either Ologamasiphis should be raised to genus rank or the diagnosis of Holoparasitus should be extended. Karg (1971) modified the diagnosis of Holoparasitus to include the subgenus Ologamasiphis Holzmann. Athias-Henriot (1971a) considered Ologamasiphis as a distinct genus rather than a subgenus of Holoparasitus. She pointed out that the subgenus Ologamasiphis of Holzmann is un- available because Holzmann did not designate a type species and designated Perga- masus epigynalis Willmann, 1940 as the type species of this genus.
Juvara-Bals (1975) distinguished two subgenera of Holoparasitus, Holopara- situs s.str. and Heteroparasitus Juvara-Bals, but considered Ologamasiphis as a sepa- rate genus rather than a subgenus of Holoparasitus. Hyatt (1987) reviewed and rede- fined Holoparasitus to include three subgenera, Holoparasitus s.str., Ologamasiphis and Heteroparasitus.
I have identified a new species from Spain in the Athias collection, which possesses a distinctive combination of characters that places it intermediate between Heteroparasitus, Paragamasus Hull, 1918 and Ologamasiphis Athias-Henriot, 1971. Consequently, Heteroparasitus is raised to genus rank and a new subgenus, Medio- parasitus subgen. n., with H. (Medioparasitus) athiasae sp. n. as type species, is defi- ned in the genus Heteroparasitus.
The confusions and the new data require a discussion of the generic and subgeneric concept of Paragamasus and Ologamasiphis.
The generic concept proposed by Juvara-Bals (1975) was the only one based on the differences in idionotal systems such as chaetotaxy, adenotaxy (gland pores), poroidotaxy (poroids), as well as on morphological characters used by other authors. The importance of these characters in taxonomy, as well as the notation employed for adenotaxy and poroidotaxy, was discussed for the first time by Athias-Henriot (1969, 1971b). Her observations have been further considered and the importance of idionotal systems in the taxonomy of the Gamasina and Ixodida has been recognized again (Krantz & Redmond, 1987; Johnston & Moraza, 1991; Klompen et al., 1996; Lindquist & Moraza, 1998).
In the taxonomic part below I update the notation of the idionotal systems as applied to pergamasine mites, so as to redefine the genus Heteroparasitus, and I add additional characters to the previous descriptions of H. tirolensis (Sellnick, 1968), H. coronarius (Karg, 1971) and H. quadratus (Witalinski, 1972). Heteroparasitus and Medioparasitus are integrated in the key of the subfamily Pergamasinae Juvara-Bals (1972) provided below.
MATERIAL AND METHODS
Most of the mites studied (69 specimens on 64 slides) are from the Athias- Henriot collection deposited in the Natural History Museum of Geneva (MHNG). The material was collected by Prof. H. Franz in Spain and Austria. These specimens are mounted in gum chloral and flattened, so that their idiosomal length and width cannot be measured. A few samples are from Germany and Poland (Witalinski leg., deposited in the Zoological Museum, Jagiellonian University, Krakow-ZMJU and in the MHNG) or Romania (Juvara-Bals leg., deposited in the MHNG). Some specimens
A REVISION OF HETEROPARASITUS DS
are from the Willmann collection deposited in the Zoologische Staatssammlung, Munich, Germany (ZSM).
Generally, the morphological terminology and the system of setal notation for the legs and palpi follows that established by Evans and Till (1979), the system of setal notation for the idiosoma follows that of Lindquist and Evans (1965), with modifications for the chaetotaxy of the opisthogaster as given by Lindquist (1994). The notation of adenotaxy and poroidotaxy follows the system of Johnston and Moraza (1991). Measurements of female structures were taken as follows: Epigynium height (h) is the midline from the tip of this shield to its posterior margin; epigynium basal width (b) represents the length of the posterior margin (Fig. 4D). The distance st-st’ is the distance between the two setae of the pair inserted on the sternal and genital shields. Measurements are given in micrometers.
In the description of the opisthogastric shield (ventrianal shield auct.) only the ventral (opisthogastric) pairs of setae were considered, the three circumanal setae were excluded because they are constant among the pergamasine species. The male genital orifice of some genera of Pergamasinae, possesses a structure between the tritosternum and the genital lamina, which I name subgenital sclerite (Fig. 3C, see arrow).
DESCRIPTIONS
Genus Heteroparasitus Juvara-Bals, 1975, status nov.
Diagnosis. Dorsal shield of adults entire, podonotal region with 18-20 pairs of setae, opisthonotal region with 21-23 pairs of setae. Dorsal adenotaxy with 3 —5 pairs of gland pores, poroidotaxy with 15 pairs. Opisthogastric shield with 7 pairs of ventral setae. Peritrematal shield in females fused or not with dorsal shield; opisthogastric shield separated from dorsal shield; digitus mobilis (d.m.) of chelicera with 4 teeth. Holodorsal shield in males fused with opithogastric region. Femur II with setiform axillary process.
Subgenus Heteroparasitus s. str.
Diagnosis. Podonotal region with 19 pairs of setae, lacking z3, s2, s3, rl; poroidotaxy with 7 pairs of poroids (idj1, idj3, idr4, idz4, ids4, idj6 and idz6 which migrated onto opisthonotum); adenotaxy with 5 pairs of gland pores (gdj2, gdz5, gds4, gdr4, gdz6). Peritrematal shield fused with dorsal shield, peritrema with three gland pores along peritrematal groove (gpl, gp2, gp3) and with two poroids (ip1, ip2). Opisthonotal region of dorsal shield with 21 pairs of setae; poroidotaxy with 10 pairs of poroids; adenotaxy with one pair of gland pores. Idiosomal venter with gland pores gvl, gv2 and gv3 present. Gland pore gv2 double (with two glands opening through two pores on an ovoidal sclerotization). Presternal sclerite joined to sternal shield in female, two additional small triangular microsclerites remain separate; male with subgenital sclerite ellipsoidal, flanked by triangular presternal platelets. Tectum trifid. Gnathosomal sclerotization, in male, with cuticular break under hypostomatic seta 3; hypognathal groove large, with 12-14 denticled ridges, corniculi with protuberance on
26 I. JUVARA-BALS
inner face; palptrochanter seta v2 pilose, palpfemur with setae allsimple but with al2 stout and slightly pilose. D.m. of chelicera in females with usually 4 teeth (3 or 4 in H. quadratus according to Witalinski, pers. com.). Femur II in males with setiform axillary process, with an apophysis on genu and tibia and with short, simple setae pdl, pd2 on femur IV.
Type species: Pergamasus tirolensis (Sellnick, 1968) by original designation. Holotype: one female, collected from “Neuleutasch bei Seefeld, Tirol, 1300m über M. Lärchwiese, VIII. 1964” and deposited in Sellnick’s collection, Zoological Institut and Museum, University of Hamburg, Germany.
Other species included: Holoparasitus coronarius (Karg, 1971), Holoparasitus quadratus (Witalinski, 1972).
Heteroparasitus tirolensis (Sellnick, 1968)
Material examined: AUSTRIA: Niederösterreich: A314, 19, 16, Wolfsbach near Admont, riverside forest, back-water of the river Enn, sifting of decayed tree stumps, 22.10.1943; X436, 19, sifting of fir litter under Vaccinium myrtillus, 800 m, road from Mitterdorf im Mürztal to Stallglam, 9.9.1944; X1551, 19, litter from Ericetum near northern slope, Manhartstal near Grossau, 15.9.1960; — Steiermark: T256, 29, fir woods, sifting of moss under Vaccinium, 1400-1500m, Seckhauer Zinken above Jörgerhütte, 4.6.1960; X1576, 14 forest litter, Lassnitzklamm near Deuschlandsberg, 14.7.1964 ; X1585, 12, 1d, beech and fir litter, forest near the road Salba to Gaberl, 26.7.1964; X1626, 1 9, litter, beech woods, Koralpe, southern slope near Urbani chapel, 18.8.1965; X1575, 19, sifting of moss and litter, conifer woods, road to Trahiitte and Glashütte, Weststeiermark, 11.7.1964. All this material is in MHNG-Athias collection. - GERMANY: 72, 6d, 1 deutonymph, litter in a mixed forest (Tilia, Fagus, Fraxinus, Abies, Picea), Petersdorf near Regensburg, 7. 9. 1999, leg. W. Witalinski. Material deposited in ZMJU. - ROMANIA: Meridional Carpathians: 19, 16, deciduous forest litter, Berzeasca, Almaj Mountain, 15.8.1970, leg. I. Juvara-Bals. Two slides deposited in MHNG. — SLOVENIA: Carniola: 14, Radna Cave, 5.3.1918, leg. K. Absolon (no775, Biospeologica Balcanica). One slide deposited in ZSM- Willmann collection.
DESCRIPTION
Only the characters not mentioned in the previous taxa descriptions are noted.
Female
Idiosomal dorsum (Fig. 1B). Podonotal region with 19 pairs of setae ( j1-j6; zl, z2, z4-z6, sl, s3, s4-s6, r3, r4, r5). Pore-like structures including 7 poroids and 5 gland pores. Opisthonotal region with 21 pairs of setae, Z2, S6 absent; adenotaxy with one gland pore, poroidotaxy including 10 pairs of poroids (idJ1, 1dJ2-double, idJ3, idJ4; idZ4, idS4, idZ5, idR1, idS7). Peritrematal region as in figure IA.
Length of setae: j1 =48-54um; other j setae about 48um, setae of s series about 42um, seta r5 =27-30um, z1=17um:; setae on opisthonotum uniform, their lengths about 36-42um.
Idiosomal venter. Sternal shield subrectangular, sternal setae moderately long, stl =48um, st2 =36um, st3 =44um; gland pore gv! present. Paragynial shield with small ventral protrusion; metagynial sclerites short, rounded, with a median trape- zoidal thickening (Fig. 2 B, C, see arrow). Endogynium a denticulate cup supported by metagynial and inner paragynial sclerites. Epigynium heptagonal, its apex roun-
A REVISION OF HETEROPARASITUS DT
00 > a
DO 33 v on
oO
—
Fic. |
Heteroparasitus tirolensis (Sellnick). Female: A-idiosoma, lateral view; B-idiosoma, dorsal view (modified after Juvara-Bals, 1975).
ded, with a little, terminal tip on its top; subapical structure conspicuous, hyaline wing-like, its anterior margin with median concavity and two lateral thickenings (Fig. 2A). Opisthogastric shield with 7 pairs of ventral setae and 3 circumanal setae; JV4, ZV5 lacking, JV5 usually inserted on soft cuticule, sometimes on opisthogastric shield. Adenotaxy with two gland pores, gv2 double and gv3; poroidotaxy including ivo2, ivo3 and ivp on soft cuticule. Length of setae: ZV3, JV5 =48u, JV3 =30um, others about 36um.
Legs. Coxa II with palmate ridge situated anterolaterally; trochanter IV with setae pdl and pd2 short, simple. Measurements: Tarsus I =150-172.5um; tarsus IV = 184-195.5mm. Epigynium: height (h) =179-198um, length of base (b) =172.5-207um, st5-st5 ‘=115-147um, h/b = about 1. Sternal shield: st1-st1°= 69-75um, st2-st2’ =87- 96um, st3-st3’ =97-103um.
28 I. JU VARA-BALS
MAMMA Ps Au, un
aaa.
Ma My aos MA, Yu
0,05mm.
FiG. 2
Heteroparasitus tirolensis (Sellnick): female: A, B, C; male: D, E. H. coronarius (Karg): female: H-I; male: F, G. H. quadratus (Witalinski): J. Female: A, I- apex of epigynium and subapical structure; B, C, H-paragynia and metagynial sclerite. Male: D-gnathosoma, ventral, D’- corniculus, dorsal; E, F-palptrochanter and palpfemur; G, J-corniculus, ventral.
A REVISION OF HETEROPARASITUS 29
Fic. 3 Male: Heteroparasitus tirolensis (Sellnick): A, B. H. coronarius (Karg): C-F; H. quadratus (Witalinski): H. Female: H. quadratus (Witalinski): G. A, D- leg II femur, genu, tibia; C-
genital lamina, anterior margin of sternal shield; H-trochanter IV; B, E, G-chelicera, paraxial view; F-opisthogastric glande pore gv2.
30 I. JUVARA-BALS
Male
Idiosomal dorsum. Idionotal systems as in female, setal lengths: j1 =36-42um, j2 =48um, s setae =30-35um; setae on opisthonotal region 24 - 30um.
Idiosomal venter. Sternogenital shield reticulated, with a marked convex scle- rotized line behind second pair of sternal setae (st2). Genital lamina square, simple, with straight anterior margin and two lateral protuberances. An ellipsoidal sclerite behind genital lamina and between presternal sclerites and base of tritosternum. Genital lamina located in a concavity surrounded by prominent lobes. Sternal setae, st2, st3 length =24um, stl = 40um; opisthogastric setae =24-30um.
Gnathosoma. Tectum with three short prongs. Palptrochanter with proximal tubercle, vl simple, v2 pilose, both setae inserted on small protuberances; palpfemur with elongated protuberance near seta all (Fig. 2E). D.m. of chelicera with one denticle (two in Romanian specimens), digitus fixus (d.f.) with slightly serrated masti- catory ridge; arthrodial cuticule (membrane sensu Evans and Till, 1979) formed by paraxial brush-like and antiaxial setiform processes (Fig. 3B). Corniculi with median protuberance; another protuberance at base of hypostomatic seta 1 (Fig. 2D’); hypo- gnathal groove with extension between corniculi, cuticular break below hypostomatic seta 3 (Fig. 2D).
Legs. Coxa I with denticulate ridge (Fig. 6N). Coxa II with palmate ridge anterolaterally (Fig. 6M). Armature of leg II (Fig. 3A): axillary process of femur with a seta, genu with medially rounded apophysis, and tibia with small blade-like apophysis. Trochanter IV with setae pdl and pd2 simple and short.
Measurements: tarsus I =149.5-161um, tarsus IV =172.5-184um.
REMARKS
H. tirolensis was described by Sellnick (1968) based on female specimens from Tirol (Neuleutasch bei Seefeld) and from Markenstein in the Wiener Wald.
Karg (1971) collected H. tirolensis adults of both sexes in Germany. He compared his material with Ologamasus absoloni Willmann, 1940 described from Slovenia (Carniola) (Biospeologica Balcanica, loc.775) and stated that the male which was originally described under O. absoloni is conspecific with the males of H. tiro- lensis from Germany.The female of O. absoloni, on which the original description is based (Willmann, 1940) belongs to a different species and is the type of Holo- parasitus absoloni (Willmann, 1940). Karg (1971) briefly described the male and gave illustrations of the ventral part of the gnathosoma and the armature of the leg II. Juvara-Bals (1975) added some characters to the original description which supported the distinction of two subgenera in Holoparasitus.
Karg (1993) considered H. tirolensis to be rare in central Europe, with a preference for damp substrates, beech and alder litter. This species was recorded also from other habitats in Austria (Schmölzer, 1995), in Romania (Juvara-Bals, 1975), in Slovenia (Willmann, 1940) and in Poland near Krakow (Witalinski, pers. comm.).
Heteroparasitus coronarius (Karg, 1971)
Only the morphological characteristics not described or illustrated in the pre- vious taxa descriptions are presented.
A REVISION OF HETEROPARASITUS 3]
Material examined: AUSTRIA: Steiermark: A23, 29, 16 sifting of alder litter, near the Mühlauer river close to a waterfall, Mühlau by Admont, 20.4.1940; A278, 12,16, sifting of decayed tree stumps, Leichenberg by Admont, southern slope, 1.11.1942; X582, 12, 1d, sifting litter, beech woods, Damberg near Steyer, northern slope, 27.4.1946; — Burgenland: X1690a, 24, 19, beech litter, Mandelstein by Weitra, 1.10.1966; — Niederösterreich: X1649, 19, litter and decayed tree stumps, Buchenberg by Waidhofen near Ybbs, 18.5.1966.
All the slides are deposited in the MHNG-Athias collection.
DESCRIPTION
Female
Idiosomal dorsum. Length of setae: jl = 48-50mm; podonotal setae 36-42um; opisthonotal setae 24-36um.
Idiosomal venter. Paragynia with small protrusion, metagynial sclerites slen- der, elongated (Fig. 2H). Epigynium heptagonal, its apex long, triangular, subapical structure round, sclerotized, covered by a fine, hyaline cuticle (Fig. 21). Length of sternal setae about 92um; opisthogastric setae about 30um.
D.m. of chelicera usually with four teeth; one specimen (X582/Q272) with three teeth on one d.m. and four on the other.
Legs. Coxa II and femur IV as in male. Measurements: tarsus I = 147-161mm, tarsus IV = 177-184um. Epigynium: h =180.5um, st5-st5’ =122um, b = 149.5um, h/b =1,21. Sternal shield: st1-st1° = 61um, st2-st2’ = 94.3um, st3-st3 = 106um.
Male
Idiosomal dorsum. Length of setae: jl = 48-54um, seta sl =12um, other s setae 24-36um; setae on opisthonotum about 24um.
Idiosomal venter. Genital lamina situated in a slight concavity bordered by two pronounced protuberances; its shape trapezoidal, with lateral corners folded, its anterior margin straight. Ellipsoidal sclerite between genital lamina and tritosternum with ribbon-like base (Fig. 3C). Sternogenital region reticulated, with marked line behind second pair of sternal setae, gland pore gv2 double (Fig. 3F). Length of sternal setae: st] =42um, st2 = 36um, st3 =30um; length of opisthogastric setae about 24um.
Gnathosoma. D.m. of chelicera with 3-6 denticles, d.f. oligodont, some denticles beside pilus dentilis; arthrodial cuticle with brush-like process paraxially (Fig. 3E).
Palptrochanter with seta vl simple and v2 pilose, both situated on rounded protuberances (Fig. 2F). Corniculi with inner protuberance (Fig. 2G).
Legs. Coxa I paraxially with a ridge formed by small and large denticles (Fig. 6K). Leg II: coxa with short rounded denticulated ridge anterolaterally (Fig. 6J); axillary process of femur with seta; genu with small trapezoidal apophysis medioventrally; tibia bearing long blade-like apophysis, its tip reaching distal margin of tibia; base of tarsus humped (Fig. 3D). Femur IV setae pdl and pd2 simple and short, tarsus IV as in H. tirolensis (Fig. 6L).
Measurements: tarsus I =145-152um; tarsus IV =165.6-177um.
I. JUVARA-BALS
o>) 159]
REMARKS
Holzmann (1969) inadvertently indentified specimens found in litter of deci- duous forest near Erlangen (Germany) as Holoparasitus (Ologamasus) rotulifer (Willmann, 1940). On the base of Holzmann’s descriptions and drawings Karg (1971) recognized that they belong to a new species, which he named A. coronarius. Holo- parasitus rotulifer is another, valıd species related to the Holoparasitus s.str. species, whose males are provided with an excipulum. Males of the related A. tirolensis and H. coronarius can be separated by the characteristics of the armature of leg II, the corniculi and the chelicera. The shape of the epigynial apex, which is triangular in H. coronarius and rounded in A. tirolensis, distinguishes females; other differen- tiating characteristics are the shape of the endogynium and of the metagynial sclerites. Karg (1971) thought that A. coronarius probably occurs in the litter of deciduous forest in Europe. Koehler (2000) found it in forest soil contamined with TNT in Harz, Germany. I have identified this species only in samples from Austria. The distribution of this species remains poorly known.
Heteroparasitus quadratus Witalinski, 1972
Material examined: POLAND: 29, 28 Myslenice near Krakow, Southern Poland, type locality, litter in a mixed forest, 10.11.1969, leg. W.Witalinski. Material deposited in MHNG.
REMARKS
I studied the material from Poland, kindly send to me by W.Witalinski. The following observations and some measurements are added to the original description (Witalinski, 1972).
In the original description Witalinski noted only 3 teeth on the movable digit of the chelicera. Witalinski and myself checked this character in a greater number of females and found that the number varies between 3 and 4 teeth (Fig. 3G).
The male has the same sclerotization break under the corniculus as males of the other species included in this genus (Fig. 2J). Trochanter IV (Fig. 3H) bears a flat protuberance on its ventral side. The reticulated ridge of coxa I is formed by 3 large denticles above a line of 9 fine denticles (Fig. 6P); coxa II has a short ridge formed by 6-7 denticles (Fig. 60).
Measurements. Male: tarsus I =138um; tarsus IV =184-195um. Female: tarsus I =145-150um; tarsus IV =196um; epigynium: h =196um, b =176um, h/b =1.1, st5- st5’ =127-136um; length of sternal shield setae 60um.
Subgenus Medioparasitus subgen. n.
Type species: Heteroparasitus (Medioparasitus) athiasae sp. n., by present designation and by monotypy.
Diagnosis: Dorsum generally with long setae, their tips reaching alveoli of following row of setae. Podonotal region apparently with 20 pairs of setae, opistho- notal region with 23. Tectum triangular. Gland pore gvl on sternal shield absent. Peritrematal shield of females united with holodorsal shield anteriorly and free posteriorly. Leg II of males with spurs only on femur and genu, femur IV with setae pdl and pd2 short and pilose; subgenital sclerite with denticles.
A REVISION OF HETEROPARASITUS 33
Description. Characteristics of the idionotal systems cannot be properly obser- ved because the specimens are mounted and the sclerocuticle is folded or crushed.
Idiosomal dorsum. Dorsal shield setae long, except for z1, sl, s2, their tips reaching following row; setae J5, Z4, Z5, S3, S4, S5 three times longer than the other one. Podonotal region, apparently with 20 setae (z3, r4 lacking ); opisthonotal region with 23 setae, S2 and perhaps some submarginal setae (UR) absent. Adenotaxy: gland pores gdz5, gds4, gdZ3 present. Poroidotaxy: podonotum with 5 pairs of poroids, opisthonotum with only 7 poroids observable. Peritrematal shield of females free, gland pore gdr4 and ip2 opening on soft cuticule; two other gland pores, gpl and gp3 located in peritrematal groove.
Idiosomal venter. Female sternal shield lacking gland pore gv1, opisthogastric shield with 7 pairs of ventral setae. Male with platelets surrounding genital lamina and subgenital sclerite.
Gnathosoma. D.m. of chelicera in females with 4 teeth; d.m. in males with row of 6-7 denticles, d.f. oligodont. Hypostome with short, fan-shaped internal malae. Tectum triangular.
Legs. Armature of leg II in males: tibia without spur, genu with one spur, and axillary process of femur very smal, bearing seta. Leg IV: femur with pdl and pd2 small and pilose, tibia with seta pl! stout and pilose, tarsus with seta pd2 very long.
Heteroparasitus (Medioparasitus) athiasae sp. n.
Type material: SPAIN: Sp408 slide J430 14, holotype; paratypes: 32, 36, Montes del Invernardero near Verin (Provincia Orense), Sierra de San Mamed, Campobecerros, sifting of litter near a stream, 24.7.1955 .
Other material: SPAIN: Sp42, 26, Ceiro de Mirador, Sierra de Luna near Algesiras (Andalusia), sifting of litter and humus under Erica sp., 28.2.1951; Sp211, 29, 16, sur- rounding of Pontevedra, Cuesta del Ralo Saluda, sifting of litter under Cyprus,1.7.1952; Sp483, 1d, 19, Isela de Onc, Prov. Pontevedra, sifting of litter under Ulex europaeus, 4.8.1956; Sp485, 19, Near the road Gondomas-Tuy (Sierra de la Grora), Bayona, sifting of litter (Quercus sp. forest), 5.8.1956; Sp500, 29, surroundings of Ezaro, south of Cabo Fisterra, sifting of litter (Quercus robur forest), 15.8.1956; Sp571, 2d, Sierra de Son (Leon) near San Feliz de las Lavanderas, Astorga, sifting of litter under Quercus and Erica arborea, 11.8.1957; Sp567, 32, 34, Rio Duerna valley, Molina Ferreda near Astorga, sifting of litter of Quercus pyrenica and Erica arborea, 10.8.1957; Sp579 16, 42 Near San Saturino (Lugo), sifting of litter under Rubus, 15.8.1957; Sp587 1d, El Fito, West of Aviles, (Oviedo), sifting of litter under Betula sp., Quercus sp., 16.8.1957; Sp496, 66, 19, Sierra, north-west of Outes, near Noia (La Coruna), sifting litter, oak forest, 13.8.1956.
All the material is deposited in MHNG-Athias collection.
DIAGNOSIS
Adults of this species are readily recognized by the characters given for the subgenus. In females the peritrematal shield free posteriorly, the d.m of chelicera with 4 teeth and the ventrianal shield with 7 pairs of setae. The male’s particularities are the shape of the subgenital sclerite, which is rectangular and denticulated, and the leg II with simple triangular spurs on femur and genu.
ng ee li
34 I. JUVARA-BALS
wat ® N
Fic. 4
Medioparasitus athiasae sp. n. Female: A-idiosoma, dorsal view; B-peritrematal region; C- sternal shield, paragynia and endogynium; D- epigynium; E- endogynium, F-opisthogaster.
A REVISION OF HETEROPARASITUS 35
Fic. 5
Medioparasitus athiasae sp. n. Male: A-H. Female: I-M. A-sternogenital region; B-genital region with subgenital sclerite (sg. scl.); C-genital lamina; D, L-chelicera, antiaxial; E, I- tectum; F, J-hypognathal groove; G, M-palptrochanter, palpfemur; H-corniculus; K-internal mala.
36 I. JUVARA-BALS
0,05mm. H-P L AI
Fic. 6 Male. Medioparasitus athiasae sp. n.: A-I: Heteroparasitus coronarius (Karg): J, K, L; A. tirolensis (Sellnick) : M, N: H. quadratus (Witalinski): O, P. Leg II, antiaxial view: A-femur, genu, tibia (Sp 408): B-femur, genu (Sp 496); C-tarsus. Leg IV, ventral view: D-trochanter; E- femur; F-tibia; G, L-tarsus. Denticulated ridge, coxa I-H, K, N, P; idem, coxa II-I, J, M, O.
A REVISION OF HETEROPARASITUS 37
DESCRIPTION Female
Idiosomal dorsum (Fig. 4A). Length of idiosomal setae: podonotum j1 =42- 45um, sl =6-7um; r2, r3, s2 =13-14um; other setae about 42-48um; opisthonotal setae from 48um (J3, S3, S4) to 54 um (S5, Z5).
Idiosomal venter. Peritrematal shields free posteriorly (Fig. 4B). Presternal sclerites fused to sternal shield, except for two little pieces. Sternal shield narrow distinctly reticulated, with prominent arched line between st2 and st3 (Fig. 4C). Length of sternal setae: stl =54um, st2 =60um, st3 =48um. Gland pore gv! absent. Posterolateral protrusion of paragynial shield large and rounded; metagynial sclerite small, ellipsoidal (Fig. 4C). Epigynium septagonal, central apex with small notch, lateral angles with sharp prongs; subapical structure small, rounded, extended on dorsal side by triangular cuticular formation ending in lateral prongs (Fig. 4D). Endogynium apron-like, flanked by two elongated sclerotized strips (Fig. 4E). Opis- thogastric shield with 7 pairs of setae, their lengths 42-48um, and with 3 circumanal setae; gland pore gv2 (double) and gv3 on cuticule (Fig. 4F).
Gnathosoma. Tectum triangular, with simple slender central prong (Fig. SI). Hypognathal groove with 9 oligodent rows; anterior hypostomatic seta and palpcoxal seta slightly pilose, others simple; internal malae fan-shaped, fimbriated; corniculi conical (Fig. 5J, K). Palptrochanter with vi simple, v2 pilose, palpfemur with all pectinate and with small rounded protuberance at base of all (Fig. 5M). D.m. of chelicera with 4 teeth, d.f. with two denticles antiaxially, one paraxially alongside pilus dentilis (Fig. 5L).
Measurements: tarsus I =127-140um, tarsus IV =161-179um. Epigynium: h =173-202um, b =156-177um, st5-st5’ =103-129um, h/b =1.13. Sternal shield: st1- stl’? =46-55um; st2-st2’ =69-87um, st3-st3’ = 92-127um.
Specimen collected near Noia (Sp496): tarsus I =110-127um; tarsus IV =150- 159um.
Epigynium: h =166-170um, b = 170-179um, v5-v5’ =108-115um, h/b =0.96.
Male
Idiosomal dorsum. Chaetotaxy with 20 pairs of setae on podonotal region and 23 pairs on opisthonotal region, setae simple, their lengths: j1 =36um, z1 =12um, s2- s3 =18um, s6 =48um, j3-j4 =54um, other setae of series z =36-42um. Opisthonotal setae of series J and S =48um, setae of series R =42um. Adenotaxy: gland pores gdj2, gdj4, gdz6, gdZ1 and gdZ4 lacking. Poroidotaxy: podonotum with 5 pairs of poroids, opisthonotal poroids not observable.
Idiosomal venter. Holoventral shield reticulated, with prominent procurved line passing behind sternal setae 2 (Fig. 5A). Anterior margin of sternal shield flanked by two protuberances and concave at location of genital lamina. Genital lamina tra- pezoidal, anterior margin folded, its angles appearing rounded (Fig. 5C). Subgenital sclerite rectangular, situated between base of tritosternum and genital lamina, with ventral and dorsal side partially denticulated and with a central triangular opening. This structure flanked on each side by two circular platelets supporting and restric- ting, probably, displacement of rectangular structure and genital lamina when open
38 I. JUVARA-BALS
(Fig. 5B). Opisthogastric shield with 7 pairs of ventral setae, their length from 24 to 36um. Gland pore gv! absent, gv2 double.
Gnathosoma. Hypognathal groove with 9-10 clearly denticulated rows, palp- coxal seta simple (Fig. 5F). Corniculi simple, conical (Fig. 5H). Tectum triangular, with central prong long, slender and with lateral margins rounded (Fig. 5E). Palp- trochanter with vl simple, v2 pilose, palpfemur with small protuberance near base of spatulate and pectinate anterolateral seta (al) (Fig. 5G).
Chelicera (Fig. 5D). D.f. without denticles, only with a rounded protuberance proximally, d.m. with 5-8 denticles and a larger tooth; apex of both digits slightly hooked. Arthrodial cuticule with small brush-like process paraxially and setiform process antiaxially.
Legs. Armature of leg II with simple triangular spurs only on femur and genu; femur with small axillary process bearing seta and with adjacent distal protuberance (Fig. 6A). Coxa I with 7 small denticles on its paraxial, distal ridge (Fig. 6H). Coxa II with serrated denticulated ridge anterolaterally (Fig. 61). Leg IV: trochanter with flattened protuberance on posterolateral side; femur with finger-like prominence posterolaterally on distal margin and with pdl and pd2 setae pilose and short (Fig. 6E); tibia with one stout pilose, posterolateral seta (Fig. 6F); tarsus with pd2 simple and long, reaching base of pdl (Fig. 6G). Measurments: tarsus I =115-129um; tarsus IV =161-173um. Specimens from near Noia (Sp 496) tarsus I =116u; tarsus IV — SOU
ETYMOLOGY
The new species is dedicated to Dr Claire Athias-Henriot who pioneered modern systematics of gamasid mites and stimulated me to carry on with my research on mites.
REMARKS
The male of M. athiasae subgen., sp. n. is characterized by the following combination of characters:
- Armature of leg II with a reduced axillary process bearing a seta; tibia without spurs.
- Genital lamina trapezoidal; rectangular denticulated sclerite situated between genital lamina and basis of tritosternum; genital orifice flanked by platelets.
- Chelicera with d.f. oligodont and d.m. with 7-8 denticles.
The female is unique among the Pergamasinae by having the combination of a peritrematal shield only anteriorly united to the dorsal shield, as in Paragamasus, and the d.m. of chelicera with 4 teeth, as in Heteroparasitus. Other peculiar characteristics are the shape of the endogynium and epigynium.
Specimens from near Noia (La Coruna) differ in some characters from spe- cimens collected from other localities: in males the sternogenital cuticule has a different pattern with two parallel sclerotized lines between st2 and st5, the distal apophysis on femur II is absent (Fig. 6B), the distal spur on femur IV is shorter; in females the proportions of the epigynium are different.
A REVISION OF HETEROPARASITUS 39
The setal shape of the tarsi II (Fig. 6C) and IV (Fig. 6G, L) are different in M. athiasae and H.coronarius. For the moment I cannot evaluete the taxonomic impor- tance of this observation because of insufficient knowledge of the setal pattern of tarsi I-IV in most Pergamasinae.
DISCUSSION
The genus Holoparasitus Oudemans, 1936 was divided by Juvara-Bals (1975) into three genera, Holoparasitus Oudemans, 1936, Heteroparasitus Juvara-Bals, 1975 and Ologamasiphis Holzmann, 1969. This decision was based on the following characters:
- Idiosomal chaetotaxy; idionotal cuticular systems; number of opisthogastric setae in both sexes.
- Females: fusion or separation of ventrianal shield with dorsal and peritre- matal shields; number of teeth on d.m. of chelicera.
- Males: structure of the genital orifice and particularities of the leg II.
Holoparasitus comprises only the species whose characters are similar to those of the type species of Holoparasitus as defined by Hyatt (1987). Ologamasiphis is a valid genus. It differs from Heteroparasitus by the patterns of the idionotal systems and by details of the armature of the leg II (3), the genital orifice (6) and the epi- gynium and endogynium (2).
Heteroparasitus is defined by a combination of features discussed in the pre- sent paper. Medioparasitus subgen. n. possesses a peculiar combination of characters, mainly in the female. Its peritrematal shield is free as in Paragamasus Hull, 1918. The digitus mobilis has 4 teeth as in Heteroparasitus, but unlike in other taxa in- cluded in Paragamasini sensu Juvara-Bals (1975) which all have only 3 teeth. The ventral gland pore gv1 is absent as in Paragamasus (Meriadenogamasus) from Nepal (Athias-Henriot, 1973) or in Ologamasiphis Athias-Henriot, 1971. The placement of Medioparasitus subgen. n. in the genus Heteroparasitus is uncertain at present. It is close to this genus by the number of opisthogastric setae, the structures of the genital orifice and of femur II in the male, and by the characters of the epigynium in the female. As mentioned above, the chaetotaxy and cuticular systems could not be properly studied because the available specimens were folded or crushed under the cover slides. However, the adenotaxy is deficient, lacking the gland pores gdj2, gdj4, gdz6, gdZ4 and gvl.
KEY TO THE GENERA OF PERGAMASINAE JUVARA-BALS, 1972 (Modified after Evans & Till, 1978 and Karg, 1993)
l Tarsus I without claws and pulvillus; holodorsal shield attenuated pos- teriorly, its opisthonotal region with less than 12 pairs of setae; idio- soma length 490 um, male unknown........... Pergamasellus Evans, 1957 Type species: P. delicatus Evans, 1957 - Tarsus I with claws and pulvillus; holodorsal shield not attenuated posteriorly, opistonotal region with more than 12 pairs of setae; idio- somaslenstht+30-I400 mE 2. ee e OI ICONA 2
40
[55]
I. JUVARA-BALS
Holodorsal, peritrematal and opisthogastric shields fused posteriorly in females and males, opisthogastric region with 8-9 pairs of ventral setae; movable digit of female chelicera with three teeth; idiosoma globular, well sclerotized, maximum length 950 um. . . Holoparasitus Oudemans, 1936 Type species: Gamasus calcaratus Koch, 1839 Female holodorsal and peritremal shield fused or separate, opistogastric shield free; in males all shields fused; movable digit of female chelicera with three or four teeth; idiosoma oval-shaped, rarely globular (in this case weakly sclerotized), maximum length 2060um................. 3 Female holodorsal and peritrematal shields fused (except in the subg. Medioparasitus), opisthogastric shield free; male without transverse suture on dorsal shield; movable digit of female chelicera with four teeth . . . 4 Female holodorsal shield anteriorly united with peritrematal shield, the latter fused or not fused with opisthogastric shield; male with or without transverse suture on dorsum idiosoma; movable digit of female chelieera‘with'three or fouriteeth.- i. RME RENE 5 Podonotal region with 18-22 pairs of setae, opisthonotal region hyper- trichous; opistogastric shield with 11-32 pairs of ventral setae, tectum with 3-5 prongs: female with two big triangular presternal sclerites, epigynium triangular or subpentagonal; femur II of male with triangular or different- shaped apophysis; idiosoma oval-shaped, length 880 to AOC OT A MR CR RE Nm ie Pergamasus Berlese, 1904 Type species: Acarus crassipes Linné sensu Berlese, 1906 Podonotal region with 20 pairs of setae, opisthonotal regions with 21- 23 pairs of setae; opisthogastric shield with 7 pairs of ventral setae, tec- tum trifid or triangular; female with presternal sclerites almost complet- ly fused to sternal shield, with small triangular structures remaining, epigynium heptagonal; femur II of male with triangular apophysis and axillary process bearing seta; idiosoma globular, length less than TACO TTT ERE te e SR Heteroparasitus Juvara-Bals, 1975 Type species: Pergamasus tirolensis Sellnick, 1968 a. Setae on dorsal scutum moderately long, not reaching line of the following setal row; female peritrematal shield fused with dorsal shield; gland pore gvl present on sternal shield; tectum trifid; male leg II with one spur on femur, genu and tibia ; subgenital sclerite oval Ro N LE GUIA E RE Subgen. Heteroparasitus s. str. b. Setae on dorsum scutum long, especially on opisthonotum, reaching mid-length of setae of the following setal row; female peritrematal shield free posteriorly; gvl absent on sternal shield; tectum triangular; male leg II with spurs only on femur and genu; subgenital sclerite rectangular with denticles............. Subgen. Medioparasitus subgen. n. Type species: M. athiasae sp. n. Female dorsal and peritrematal shields united anteriorly, peritrematal shield fused with opisthogaster; extension of peritrematal shield behind stigmata discernible in male; male without transverse suture on idiosoma . . . 6
A REVISION OF HETEROPARASITUS 41
Female dorsal and peritrematal shields united anteriorly, peritrematal shield free posteriorly; extension of peritrematal shield behind stigmata not recognizable in male; male with or without transverse suture on CONSUMO CIOSOMAIE EA RO RR RT SE n 9 Podonotal region with 19-20 pairs of setae, opisthonotal region poly- trichous; palpgenual setae all and al2 bifid or fringed or foliaceous; presternal sclerites of female large, triangular, contiguous; movable digit of male chelicera with two teeth, subgenital sclerite present, idio- somatlenstht800-12 00 mts: srr sth eth ee yen ean oe a 7 Podonotal region with 13-20 pairs of setae, opisthonotal region holo- trichous (23-24 pairs of setae) or oligotrichous; palpgenual setae all and al2 truncate, presternal sclerites of female triangular, small, distant from each other or contigous; movable digit of male chelicera with one tooth, subgenital sclerite absent, idiosoma length 350-800um ........... 8 Palpgenual seta all bifid and al2 foliaceous; opisthogastric shield with 23-29 pairs of ventral setae, sclerocuticle wrinkled; movable digit of female chelicera with four teeth; idiosoma large, length 1000-1200um Re Rea ida: addio FILI Mixogamasus Juvara-Bals, 1972 Type species: M. intermedius Juvara-Bals, 1972 Palpgenual setae alland al2 fringed, opisthogastric shield with 11-16 pairs of ventral setae; sclerocuticle smooth; movable digit of female chelicera with three teeth and adjacent denticles between them; idio- soma length of male 787-896um, of female 1028-1300um © Gd Chas se RER ESPRIT aoe Phytiogamasus Juvara-Bals & Athias-Henriot, 1972 Type species: Parasitus primitivus Oudemans, 1904 Podonotal region with 19-20 pairs of setae, opisthonotal region with 23- 24 pairs; opisthogastric shield with 9-10 pairs of ventral setae; movable digit of male chelicera with one tooth, movable digit of female cheli- cera with four teeth; idiosoma length of male 350-800um, of female ZA OOOO MINES eni eee Leptogamasus Tragardh, 1936 Type species: L. suecicus Trägardh, 1936 Podonotal region with 13 pairs of setae, opisthonotal region with 12 pairs; opisthogastric shield with 7 pairs of ventral setae; movable digit of female chelicera with three teeth; idiosoma length of female 450um, malesunknownlt sense Zelogamasus Hennessy & Farrier, 1989 Type species: Z. oligochaetus Hennessy & Farrier, 1989 Podonotal region with 22-23 or 31-45 pairs of setae, opisthonotal region hypertrichous, with about 60 pairs of setae; opisthogastric shield with 11-30 pairs of ventral setae; palpgenual setae alland al2 fringed; movable digit of male chelicera with two teeth; female genital pores iv5 near posterior margin of epigynium; presternal sclerites of female trian- gular, contiguous; idiosoma length of male 700-885um, of female OSES OU MiMi ee er Aan ogee on Aa | Amblygamasus Berlese, 1906 Type species: Gamasus dentipes Koch, 1839
42 I. JUVARA-BALS
- Podonotal region with 16-18 or 20-21 pairs of setae, opistonotal region oligotrichous or holotrichous; opisthogastric shield with 8-11 pairs of ventral setae; palpgenual setae all and al2 always entire, spatulate; male movable digit of chelicera with one or two teeth; female genital pores 1v5 on soft cuticle between epigynium and opisthogaster; prester- nal sclerites of female triangular or ribbon-like, or in the shape of small triangular sclerites with intermediate sclerotizations, or fused with ster- nal shield; idiosoma length of male 420-1100um, of female 450- ZO RR AP EE nS 10
10 Podonatal region with 16-18 pairs of setae, opisthonotal region with 14-
21 pairs of setae, opisthogaster shield with 7-8 pairs of setae; male movable digit of chelicera with 2 teeth, femur II without axillary pro- cess, genital opening without subgenital sclerite but with a sclerified tape (ribbon) linked with the anterior margin of sternal shield; female with presternal sclerites fused to sternal shield, idiosoma globular ROOT aire ro PRO AE € SE AA RARES Ologamasiphis Athias-Henriot, 1971 Type species: Pergamasus epigynialis Willmann, 1940 - Podonotal region with 20-21 pairs of setae, opisthonotal region with 23- 24 pairs of setae, opisthogastric shield with 11pairs of ventral setae; male movable chelicera with one or two teeth, femur with axillary process, genital opening with subgenital sclerites; presternal sclerites of female triangular or ribbon like or in the shape of small sclerites with intermediate sclerotizations; idiosoma oval........ Paragamasus Hull, 1918 Type species: Parasitus robustus Oudemans, 1902
COMMENTS ON THE KEY
During the last twenty years a large number of taxa have been described and a great variety of characters used in the descriptions of species and supraspecific taxa of Pergamasinae. Unfortunately, the descriptions were often inconsistent, resulting in divergent taxonomic concepts.
The aim of the present key is to give a practical tool for identification of the genera in Pergamasinae. No attempt was made to analyse the phylogenetic relation- ships, because important characters of some taxa remain unknown. This is parti- cularly true for the characters common to both sexes, such as the idionotal pore-like systems (glands, poroids) and the leg chaetotaxy, especially that of the legs II and IV, as well as of the sensory field of the leg I. Characteristics of the idionotal systems are very important for the classification of the supraspecific taxa (Johnston & Moraza, 1991; Klompen er al., 1996). These characteristics have been studied for only some of the genera presently placed in the Pergamasinae (Athias-Henriot, 1971 b, 1973; Juvara-Bals, 1972, 1975, 1981). A revision of the genera and subgenera is required for a better understanding of the supraspecific concepts in the subfamily.
Only the genus Heteroparasitus 1s keyed to subgeneric level in order to ac- commodate Medioparasitus in the system. Nevertheless, some clarifications concer- ning the subgenera Leptogamasus Trägardh, 1936 and Paragamasus Hull, 1918, and the genus Ologamasiphis Athias-Henriot, 1971 are given here.
A REVISION OF HETEROPARASITUS 43
The genus Leptogamasus includes three subgenera, 1.e.: Leptogamasus Träg- ardh, 1936 (type species: L. suecicus Trägardh, 1936), Ernogamasus Athias-Henriot, 1971 (type species: Pergamasus leruthi Cooreman, 1951), Tomeogamasus Athias- Henriot, 1971 (type species: Pergamasus falculiger Berlese, 1906).
Valigamasus Karg, 1993 is a junior objective synonym of Ernogamasus, which is based on the same type species (syn. nov.). Diagnoses of the subgenera were given by Athias-Henriot (1971a, 1972) and by Juvara-Bals (1981) who raised the sub- genera to genus rank. It is beyond the scope of this paper to further discuss the generic or subgeneric level of these insufficiently known taxa. I consider, however, for the moment, Leptogamasus as a genus with three subgenera.
Within Paragamasus Hull, 1918 nine subgenera were recognized (Athias- Henriot, 1971a, 1973) corresponding mainly to her “ types d’ organisation ” (Athias- Henriot, 1967). Karg (1971) distinguished two subgenera within Paragamasus 1.e.: Paragamasus Hull, 1918 and Lysigamasus Karg, 1971, which he later raised to gene- ric level (Karg, 1993). The only given character separating the latter taxa is the shape of the presternal sclerites. Paragamasus Hull sensu Karg (1993) includes the sub- genera Paragamasus s.str. and Aclerogamasus Athias-Henriot, 1971. Lysigamasus is a mixture of the subgenera Anidogamasus, Anchigamasus, Dyogamasus and Tany- gamasus as proposed by Athias-Henriot (1971a). Karg’s short diagnosis of this genus it is in need of some more accuracy and details. Lysigamasus is commonly used as valid but according to the International Code of Zoological Nomenclature (ICZN, 1999, art. 21.3) it is a junior subjective synonym of Anidogamasus Athias-Henriot, 1971. Both subgenera were described in 1971, with Athias-Henriot’s paper issued on 17.05. 1971, but an exact date for Karg’s description is unknown. A study of the groupings in Paragamasus is beyond the scope of this paper. The Paragamasus group needs a revision, which takes into account the characters of the idiosoma (chaetotaxy, adenotaxy, poroidotaxy ) and of the leg chaetotaxy, and which finally rearranges the genus into taxa that correspond to natural groups. In the following key Paragamasus is considered as a genus with several subgenera.
The genus Ologamasiphis presents many problems and requires a thorough revision. Holzmann (1969) established Ologamasiphis as a subgenus in Holoparasitus for two species, O. minimus Holzmann, 1969 and O. rotulifer Willmann, 1940, but she did not designate a type species. Karg (1971) considered O. rotulifer sensu Holzmann 1969 as a new species that he named Holoparasitus coronarius and he supported Ologamasiphis as a subgenus of Holoparasitus, with three species, H. (O.) coronarius, H. (O.) tirolensis and H. (O.) minimus but he also did not designate a type species. According to the International Code of Zoological Nomenclature (ICZN, 1999, art. 13.3) the name Ologamasiphis Holzmann, 1969 is thus unvailable. Athias- Henriot (1971a) considered Ologamasiphis as a separate genus in which she included four species belonging to her “type d’organisation epigynalis”. She designated Per- gamasus epigynalis Willmann, 1940 as the type species and considered O. minimus Holzmann, 1969 synonymous with O. disfistulatus (Athias-Henriot, 1967). This synonymy has to be verified because the respective descriptions differ in some cha- racters. It is impossible, for the moment, to compare O. disfistulatus with O. minimus
44 I. JUVARA-BALS
because the latter species was deposited in the Holzmann collection, which was unfortunately mislaid. More research has to be carried out when this material becomes available again.
The species included by Athias-Henriot (1967,1971a) in Ologamasiphis be- long to two groups, to which I attribute subgeneric status:
1 - Ologamasiphis sensu str., type species P. epigynalis (Willmann, 1940).
The following species are included: O. parnethortus (Athias-Henriot, 1967), O. bulgatulus (Athias-Henriot,1967), O. turdetanus (Athias-Henriot,1967), O. judae- ortus (Athias-Henriot,1967).
Diagnosis: Podonotal region with18-20 pairs of setae, opisthonotum with 21- 22 pairs of setae, adenotaxy with 4 pairs of podonatal and 3 pairs of opisthonotal gland-pores; on ventral idiosoma gv! absent; in females peritrematal shield anteriorly fused with the margin of dorsal shield and posteriorly free, d.m. of chelicera with 3 teeth; in males a ribbon-like structure, instead of a subgenital sclerite, linked with anterior margin of sternal shield.
2 - Ologamasiphis (Holzmannia) subgen. n., type species Pergamasus dis- fistulatus (Athias-Henriot, 1967), syn. O. minimus Holzmann, 1969.
Diagnosis: Podonotal region with 16 pairs of setae, opisthonotum with 20; adenotaxy with 4 pairs of podonotal and 2 pairs of opisthonotal gland-pores; on ventral idiosoma gvl present; in females peritrematal shield fused with dorsal shield, d.m.of chelicera wih 3-4 teeth; in males subgenital sclerite absent.
I also assign to this new subgenus one unidentified specimen found in the Athias collection (slide H3/G 260) and the specimens which Juvara-Bals (1975) identified as O. disfistulatus which probably belong to a new species.
ACKNOWLEDGEMENTS
I thank for providing the specimens used in the above descriptions Drs V. Mahnert and P. Schwendinger (Geneva, Switzerland) and Dr W. Witalinski (Krakow, Poland). For reviewing the manuscript, for their critical comments and helpful suggestions I kindly thank Drs E. E. Lindquist (Ottawa, Canada), I. Lobl, P. Schwen- dinger and C. Lienhard (Geneva, Switzerland). I am indebt to Drs Axel Christian (Gorlitz, Germany) and L. Tiefenbacher (Munich, Germany) for their kind assistance in trying to obtain some type material. Dr C. Vaucher (Geneva, Switzerland) kindly provided working space and laboratory facilities.
REFERENCES
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ATHIAS-HENRIOT, C. 1969. Les organes cuticulaires sensoriels et glandulaires des gamasides. Poroidotaxie et adénotaxie. Bulletin de la Société Zoologique de France 94 (3): 485- 492.
ATHIAS-HENRIOT, C. 1971a. Paragamasus (Tanygamasus) probsti (Oudemans) (Systématique, géographie), avec quelques mises au point synonymiques. (Arachnides, Gamasides tocospermiques, Parasitidae). Zoologische Mededelingen 45, 16: 169-179.
A REVISION OF HETEROPARASITUS 45
ATHIAS-HENRIOT, C. 1971 b. La divergence néotaxique des Gamasides (Arachnides). Bulletin Scientifique de Bourgogne 28: 93-106.
ATHIAS-HENRIOT, C. 1972. Espèces frangaises du sous-genre Leptogamasus s. s. (Arachnida, Gamasida, Parasitidae: genre Leptogamasus). Annales de la Société entomologique de France 8 (1): 189-204.
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SELLNICK, M. 1968. Zwei neue Pergamasus Arten aus Osterreich. Berichte des Naturwissen- schaftlich-Medizinischen Vereins in Innsbruck 56: 463-472.
WILLMANN, C. 1940. Die Acari der Höhlen der Balkanhalbinsel. Studien aus dem Gebiete der allgemeinen Karstforschung, der wissenschaftlichen Hohlenkunde, der Eiszeitforschung und den Naturgebieten, Biologische Serie, Brno, 8: 1-79.
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REVUE SUISSE DE ZOOLOGIE 109 (1): 47-113; mars 2002
A taxonomic revision of the family Oncopodidae III. Further new species of Gnomulus Thorell (Opiliones, Laniatores)
Peter J. SCHWENDINGER! & Jochen MARTENS?
! Muséum d’histoire naturelle, case postale 6434, CH-1211 Genève 6, Switzerland.
? Institut für Zoologie, Johannes Gutenberg-Universität Mainz, Saarstr. 21, D-55099 Mainz, Germany.
A taxonomic revision of the family Oncopodidae III. Further new species of Gnomulus Thorell (Opiliones, Laniatores). - Twenty-one new Gnomulus species are described and placed in nine species groups. The new taxa are: G. carinatus (Kalimantan), G. claviger (Philippines), G. crassipes (Philippines), G. exsudans (Sarawak, Sabah), G. hamatus (Phi- lippines), G. hutan (Sarawak), G. javanicus (Java), G. latoperculum (Sula- wesi), G. leofeae (Myanmar), G. lomani (Borneo), G. marginatus (Thai- land), G. matabesar (Halmahera), G. monticola (peninsular Malaysia), G. obscurus (Sarawak), G. pilosus (peninsular Malaysia), G. rostratoideus (peninsular Malaysia), G. ryssie (Thailand), G. sinensis (southern China), G. spiniceps (Vietnam), G. tuberculatus (Sumatra) and G. tumidifrons (Halmahera). Additional specimens of G. armillatus (Thorell) and G. laruticus Martens & Schwendinger are reported and illustrated, respec- tively. Relationships and zoogeography are discussed.
Key-words: Opiliones - Oncopodidae - Gnomulus - new species - taxo- nomy - zoogeography - Asia.
INTRODUCTION
In our preceding paper on the Oncopodidae (Schwendinger & Martens, 1999b) we have re-examined the 27 known species of Gnomulus without taking new taxa into account. Here we add 21 new species to this genus. Most of them were collected fairly recently by means of leaf litter sifting and soil extraction; a few others were found in old collections, where they had been misidentified by previous taxonomists who did not examine the genitalia of these specimens. The present number of 48 nominal Gnomulus species is remarkable when considering that oncopodids were long regarded as being extremely rare. Prior to a partial revision by Schwendinger in 1992, merely 21 species (including G. thorelli Sorensen, which was then overlooked) were known for the whole family.
In an attempt to keep some degree of order within this species-rich and presumably further expanding genus, 11 preliminary species groups are distinguished.
Manuscript accepted 03.10.2001
48 P. J. SCHWENDINGER & J. MARTENS
This grouping has no nomenclatural relevance; it is done for purely practical reasons and does not represent the results of a thorough phylogenetic analysis. Some (hopefully the majority) of our species groups may actually correspond with mono- phyletic lineages, others may not. A few species, which stand isolated and do not fit in well with any group of related species, are placed in monotypical species groups. It is hoped that additional new species will be found, which will either link these outsiders to other groups or will prove that they belong to distinct lineages (as in the case of the sumatranus-group 1n here).
Four of the six groups distinguished by Schwendinger & Martens (1999b: 979) are re-evaluated and five additional groups are added. The aborensis-group (with three species from central Nepal, northeastern India and northern Thailand) is excluded, because no new material has become available. The rostratus-group (with two described species from peninsular Malaysia) is not treated here either. Several new species have meanwhile been discovered in peninsular Malaysia and Thailand, which also belong to this very distinct species group. They will be treated separately.
MATERIALS AND METHODS
External structures were studied and drawn with a ZEISS SVII stereomicro- scope, the penes with a NIKON Optiphot compound microscope (each with a drawing tube). The penes were expanded by placing them in hot lactic acid and then in destilled water. Expansion is reversed when the penes are transferred to 70% alcohol.
Body measurements refer to the dorsal scutum. Leg articles were measured on their dorsal side, from joint to joint. All measurements are given in mm. Terminology of penis morphology follows that of Martens & Schwendinger (1998: fig. 1).
Abbreviations used in the text: AMNH American Museum of Natural History, New York; BMH Bishop Museum, Honolulu; MAR collection of J. Martens, Mainz; MSNG Museo Civico di Storia Naturale, Genova; NHML Natural History Museum, London [formerly British Museum (Natural History)]; MHNG Muséum d’histoire naturelle, Geneve; NSMT National Science Museum, Tokyo; SMF Naturmuseum und Forschungsinstitut Senckenberg, Frankfurt; ZMB Museum für Naturkunde der Humboldt-Universität, Berlin; ZMC Zoologisk Museum, Kgbenhavn; ZMH Zoolo- gisches Institut und Museum, Universitàt Hamburg.
TAXONOMY
Gnomulus Thorell, 1890
Synonymy and diagnosis: See Martens & Schwendinger (1998: 526) and Schwendinger & Martens (1999b: 946).
Type species: Gnomulus sumatranus Thorell, 1891. Designated by ruling of the International Commission on Zoological Nomenclature (2001), following an application by Schwendinger & Martens (1999a). See also Schwendinger & Martens (1999b: 946).
FURTHER NEW SPECIES OF GNOMULUS 49
Fic. |
Records of Gnomulus species treated in this paper. - 1 Omei Shan (G. sinensis sp. n.), 2 Cuc Phuong (G. spiniceps sp. n.), 3 Tham Pu Lub (Gnomulus sp.), 4 Khao Yai N. P. (Gnomulus sp.), 5 Nam Tok Phliu N. P. (G. marginatus sp. n.), 6 Ko Chang N. P. (G. marginatus sp. n.), 7 Kaeng Krachan N. P. (G. ryssie sp. n.), 8 Malewoon (G. leofeae sp. n.), 9 Ko Siray (Gnomulus sp.), 10 Jeram Pasu (Gnomulus sp.), 11 Maxwell Hill (G. laruticus Martens & Schwendinger; Gnomulus sp.), 12 Chenderiang (Gnomulus sp.), 13 Cameron Highlands (G. monticola sp. n.), 14 Taman Negara (G. pilosus sp. n.), 15 Kota Tinggi (G. rostratoideus sp. n.), 16 Bukit Timah (G. rostratoideus sp. n.), 17 Ketambe (G. tuberculatus sp. n.), 18 Bukit Lawang (Gnomulus sp.), 19 Deli (Gnomulus sp.), 20 Gunung Kerinci (G. armillatus (Thorell)), 21 Mt. Gede (G. Javanicus sp. n.), 22 Bandjermasin (G. carinatus sp. n.), 23 Santubong (Gnomulus sp.), 24 Kuching (G. obscurus sp. n.), 25 Kapit (G. hutan sp. n.), 26 Gunung Mulu N. P. (G. exsudans sp. n.), 27 Sepilok (G. exsudans sp. n.), 28 Sapagaya (G. exsudans sp. n.), 29 Mt. Kinabalu (Gnomulus sp.), 30 Tiger Hill (Gnomulus sp.), 31 Nunukan Island (Gnomulus sp.), 32 Sagada (Gnomulus sp.), 33 Quezon N. P. (Gnomulus sp.), 34 Mt. Banahaw (G. hamatus sp. n., G. claviger sp. n., G. crassipes sp. n.), 35 Mt. Makiling (G. hamatus sp. n., G. claviger sp. n.), 36 Baybay (Gnomulus sp.), 37 Morotai (Gnomulus sp.), 38 Tobelo (G. matabesar sp. n.), 39 Buli (G. tumidifrons sp. n.), 40 Waigeo (Gnomulus sp.), 41 Dumoga - Bone N. P. and Gunung Tongara (G. latoperculum sp. n.).
50 P. J. SCHWENDINGER & J. MARTENS
THE SINENSIS-GROUP (new)
Diagnosis: Medium-sized (4.3-5.5 mm) species with robust chelicerae; no ventrobasal process on palpal femur; stigmatic pit without tubercle on posterior margin; dorsal scutal areas only indistinctly elevated, not medially divided by a pronounced longitudinal furrow as in the aborensis-group. Glans penis with distally hook-shaped, outwards-bent lateral sclerites and a pair of subterminal ventral teeth on the stylus; stylus base bulbous. This species group is very close to the aborensis- group (Schwendinger & Martens, 1999b: 948); it comprises two species, G. sinensis sp. n. and G. spiniceps sp. n., from the northeast of the known distribution area of Gnomulus.
Gnomulus sinensis sp. n. Figs 2-9
Material: CHINA, Sichuan Province, Omei Shan, Wannian, 1050 m, 1 d holotype (MHNG), leg. W. Schawaller, 19.IIT.1999.
Etymology: Latin: sinensis (adjective of sina) = chinese.
Diagnosis: Close to G. aborensis (Roewer) but distinguished by: Body smal- ler; lateral tubercles on posterior carapace region wider; dorsal scutal areas less elevated; chelicerae less robust, with only a low ventral mound on proximal article; proximal palpal femur without dorsal boss or ventral process; palpal trochanter with strong ventral process: truncus penis fairly stout, distally truncate; glans penis short, wide; lateral sclerites strongly convex, distally narrow, basally elevated; median plate short, completely covering membraneous tubes.
Description: 3 (holotype). Coloration: Body light amber, with dark brown reticulation on carapace and on proximal articles of pedipalps and chelicerae. Legs mostly brown, interspersed with dark areas; leg tarsi II, IV, palpal tarsi and cheliceral hand light amber, leg tarsi I, II cream. Dorsal scutum with dark patches on lateral margin; dorsal and ventral scutal areas with dark margin around amber central portion; anterior dorsal scutal areas medially divided by a shallow amber furrow, ventral areas undivided. Genital operculum dark. Fig. 9a-c.
Carapace short, with indistinct low, widely rounded eye tubercle and a pair of broadly rounded lateral tubercles below wide, undivided carapace-abdomen bridge (Fig. 9a, c). Dorsal scutum with anterior areas only slightly elevated, posterior ones more distinctly so; ventral scutal areas only moderately swollen (Fig. 9c), bearing “encrusted” hairs (see Schwendinger & Martens, 1999b: fig. 69a, b). Palpal coxa with large ventral process; ventral side of leg coxa I without anterolateral process; ventral side of leg coxa II with small anteroproximal process (no posteroproximal one), coxa III without process. Genital operculum quite large, somewhat triangular in shape, slightly wider than long; posterior margin of stigmatic pit without tubercle (Fig. 9b).
Chelicerae (Fig. 6) fairly robust; proximal article with distinct dorsodistal to dorsomedian boss; ventral side with low, wide mound.
Palps (Fig. 7): Ventral side of femur without proximal process; trochanter with strong, slightly distad-inclined ventral process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus of leg II 2.2 times longer than wide (Fig. 8).
FURTHER NEW SPECIES OF GNOMULUS 5]
Fics 2-8
Gnomulus sinensis sp. n., d holotype. - Penis, dorsal (2) and lateral view (3); apex of penis, dorsal (4) and lateral view (5). Left chelicera, retrolateral view (6); left palp, retrolateral view (7); distal part of left leg I, retrolateral view (8). - Scale lines 0.1 mm (4, 5), 1.0 mm (others).
Penis (Figs 2-5): Truncus penis fairly stout, its distal margin widely rounded and slightly invaginated. Glans penis short, wider than truncus at that point; lateral sclerites strongly convex, in proximal portion elevated above median plate, with wrinkles on lower side and with narrow, outward-bent, strongly hook-like tips; mem- braneous tubes completely covered by a short, broadly triangular median plate; stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
2. Unknown.
Measurements: (3): Body 4.32 long, 3.29 wide; carapace region 1.06 long, 2.00 wide. - Palp and legs:
Ahr Fe Pa ID Mt Ta Total Palp 0.67 0.96 0.62 0.44 - 0.91 3.60 Leg I 0.52 1.58 Oa 0.82 1.43 0.74 5.86 Leg II 0.59 2.03 1.01 1.26 2.03 IIS 8.03 Leg III 0.52 1.48 0.79 0.86 1.63 0.59 SION Leg IV 0.64 1.93 1403 12311 DEAD. 0.74 8.07
Relationships: Gnomulus sinensis sp. n. is closest to G. spiniceps sp. n.
Distribution and bionomics: Known only from Mount Omei (3079 m) in southern China. The specimen was sifted from the forest floor of a subtropical broad- leaf-forest. This is so far the northernmost record for the family Oncopodidae BES.
Gnomulus spiniceps sp. n. Figs 10-19
Material: VIETNAM, Ninh Binh Province, Cuc Phuong National Park, 450 m, about 40 km NW of Ninh Binh, 3 holotype (NSMT-Ad 174), leg. S. Nomura, 15.X.1995.
52 P. J. SCHWENDINGER & J. MARTENS
Fic. 9
Gnomulus sinensis sp. n., d holotype. - Body, dorsal (a), ventral (b) and lateral view (c). - Scale line 1.0 mm.
Etymology: Latin: spina = thorn, spine, ceps (from caput) = head; noun in apposition. The specific epithet refers to the long and pointed eye tubercle of the holotype.
Diagnosis: Closest to G. sinensis sp. n., distinguished by: Eye tubercle pro- nounced, pointed; posterior scutal areas less elevated; distitarsus II longer; penis more slender, with a narrower glans and a widely truncate median plate.
FURTHER NEW SPECIES OF GNOMULUS 53
Se f 15
DEN
ur
SA
IN
Fics 10-18
Gnomulus spiniceps sp. n., 6 holotype. - Penis, dorsal (10) and lateral view (11); penis with expanded glands, lateral view (12); apex of penis, dorsal (13) and lateral view (14); expanded glans, dorsal view (15). Left chelicera, retrolateral view (16); left palp, retrolateral view (17); distal part of left leg II, retrolateral view (18). - Scale lines 0.1 mm (13-15), 1.0 mm (others).
Description: 3 (holotype). Coloration: Body light amber, with dark brown reticulation on carapace and on proximal articles of pedipalps and chelicerae. Dorsal scutum with dark pattern on scutal elevations and with dark patches on light lateral and posterior margin (Fig. 19a, c). Ventral side of body light amber, ventral scutal elevations pale, with dark fringes; genital operculum darkened in its centre (Fig. 19b). Trochanters and femora of palps and trochanters to metatarsi of legs darkened (on posterior legs most distinctly so in proximal portion), palpal tarsi and cheliceral hand light amber, leg tarsalia II cream.
Carapace with distinct, acutely pointed eye tubercle; carapace-abdomen bridge wide, undivided, with very wide, low tubercles below. Dorsal scutal areas slightly elevated, medially indistinctly broken by a shallow longitudinal furrow; ventral scutal areas moderately swollen, without modified hairs (Fig. 19a, c). Palpal coxa with large ventral process; leg coxa I with small anterolateral process; leg coxa II with distinct anteroproximal and indistinct posteroproximal processes; coxa III without process. Genital operculum anteriorly rounded, slightly wider than long; posterior margin of stigmatic pits without tubercle (Fig. 19b).
Chelicerae (Fig. 16) fairly robust; proximal article with distinct, forward- inclined dorsodistal to dorsomedian boss and knob-shaped proventral subdistal process.
Palps (Fig. 17): Ventral side of femur without proximal process; trochanter with long, slightly distad-inclined ventral process.
Legs 1342, tarsal formula 2-2-3-3. Distitarsus II about 2.9 times longer than wide (Fig. 18).
54 P. J. SCHWENDINGER & J. MARTENS
Fic. 19
Gnomulus spiniceps sp. n., d holotype. - Body, dorsal (a), ventral (b) and lateral view (c). - Scale line 1.0 mm.
Penis (Figs 10-15): Truncus penis relatively slender, its anterior margin widely arched. Glans short, narrower than truncus at that point; lateral sclerites strongly convex, elevated at proximolateral margins, with strongly outward-bent hook-like tips carrying transversal wrinkles on lower side; membraneous tubes completely covered by short, very wide median plate with almost straight distal margin and distinctly
FURTHER NEW SPECIES OF GNOMULUS 55
dentate lateral corners; stylus slender, base bulbous, apex with a small pair of sub- terminal ventral teeth.
2. Unknown.
Measurements: (6): Body 5.46 long, 4.06 wide; carapace region 1.48 long, 2.31 wide. - Palp and legs:
Ihe Fe Pa Ti Mt Ta Total Palp 0.84 123 0.84 0.62 - 1.28 4.81 Leg I 0.64 2:02 0.96 1.03 17/7 0.76 7.18 Leg II 0.79 2.66 1555) 1.67 2:93 1.16 10.14 Leg III 0.59 2402 0.96 1.08 1.94 0.69 128 Leg IV 0.79 2:61 1.28 1.67 2.95 0.81 10.11
Relationships: Gnomulus spiniceps sp. n. is most closely related to G. sinensis sp. n. External morphology of both new species shows a clear relationship with the aborensis-group, but their penis morphology is distinct.
Distribution: Known only from the type locality in northern Vietnam [Fig. 1
Q)]-
THE ASLI-GROUP (see Schwendinger & Martens, 1999b: 956)
Diagnosis: These small (2.3-4.2 mm) species can be further characterized by: Palpal trochanter with a distinctly distad-directed ventral process; tubercle on pos- terior margin of stigmatic pit distinct (G. laruticus, G. monticola sp. n., G. pilosus sp. n.), indistinct or absent (other species); stylus penis with a ventral pair of sub- terminal teeth and a bulbous base.
Species account and distribution: Five species are known from the western and central parts of peninsular Malaysia, i.e. G. asli Martens & Schwendinger, G. hirsutus Martens & Schwendinger, G. laruticus Martens & Schwendinger, G. monticola sp. n. and G. pilosus sp. n.
Gnomulus laruticus Martens & Schwendinger, 1998 Figs 20-22
Gnomulus laruticus Martens & Schwendinger (1998: 539-542, figs 105-113).
New material: MALAYSIA (peninsula), Perak, Maxwell Hill near Taiping (type locality) [Fig. 1 (11)], at 1150 m, 2 8,7 2, 24.- 25.X1.1999, at 1250 m, 1 gd, 23.X1.1999; all specimens leg. G. Cuccodoro & I. Löbl (1 6, 1 £ in MAR, others in MHNG).
Remarks: All newly collected specimens possess the unusual tarsal formula (2-2-2-2), which confirmes that this is a diagnostic character for G. laruticus. The amber coloration and dark markings of the new specimens are more pronounced than in the holotype and they all possess a genital operculum with a dark central zone. Their penes largely correspond with that of the holotype, but one d has a distinctly narrower apex and a lateral glans sclerite with an inwards pointing apex (only on one side, probably deformed; Fig. 22).
The 2 from the same locality (at 1200 m) mentioned in Martens & Schwen- dinger (1998: 549) clearly belongs to a different, presumably undescribed species.
56 P. J. SCHWENDINGER & J. MARTENS
21
iS 2
Fics 20-22
Gnomulus laruticus Martens & Schwendinger. - Apex of penis of three males, dorsal view. - Scale line 1.0 mm.
Gnomulus monticola sp. n. Figs 23-35
Material: MALAYSIA (peninsula), Pahang, Cameron Highlands, near Tanah Rata: Gunung Jasar, 1550 m, trail 11, d holotype (MHNG), 16,19 paratypes, 3 juv., 24.11.1993; G. Jasar, 1720 m, 1 8,2 © paratypes, 25.11.1993; trails 4 and 13 (E of Tanah Rata), 1500 m, 1 3d paratype, 23.III.1993; trail 9 (between Tanah Rata and Robinson Fall), 1400 m, 2 2 paratypes, 27.III.1993, all leg. I. Löbl & F. Calame. - Ringlet, 960 m, 1 juv., leg. T. Jaccoud. 1 3,1 © paratypes in MAR, others in MHNG.
Etymology: Latin: monticola = mountain dweller; noun (male gender) in apposition.
Diagnosis: Close to G. asli, distinguished by: Body larger; colour pattern different; anterior dorsal scutal margin less rounded; teeth of carapace-abdomen bridge longer, more widely separated; glans penis with more rounded median plate; tips of lateral glans sclerites very close to each other.
Description: 3 (holotype). Coloration: Body light amber, ventral scutum more reddish; characteristic dark pattern on dorsal and ventral scuta. Genital operculum reddish amber with dark central zone (Fig. 35a-c). Leg segments (except tarsi) darkened, with light circular distal band on all tibiae and light median bands on metatarsi HI and IV and (less distinct) on all femora. Palps and chelicerae light amber, with a dark reticulation (faint on tarsus and cheliceral hand, respectively).
Carapace with low rounded eye tubercle; no lateral tubercles present. Cara- pace-abdomen bridge distinctly divided, composed of two widely separated opposing pairs of fairly long conical processes. Dorsal and ventral scutal areas only slightly elevated (Fig. 35a, c). Ventral scutum covered with fine short hairs (much denser than on dorsal scutum). Palpal coxa with distinct ventral process; leg coxa I with low, wide anterolateral one; ventral side of leg coxae II and III with small anteroproximal
FURTHER NEW SPECIES OF GNOMULUS 57
Fics 23-34
Gnomulus monticola sp. n., 6 holotype (27, 28, 30, 32, 33), d paratypes (23-26, 29), ? paratype (31, 34). - Penis, dorsal (23) and lateral view (24); apex of penis, dorsal (25, 27, 29) and lateral view (26, 28). Left chelicera, retrolateral view (30, 31); left palp, retrolateral view (32); distal part of left leg II, retrolateral view (33); anterior body and proximal palp (34). - Scale lines 0.1 mm (25-29), 1.0 mm (others).
processes, coxa II also with small posteroproximal one. Genital operculum slightly wider than long; a small but distinct tubercle on posterior margin of stigmatic pit (Fig. 35b).
Chelicerae (Fig. 30): Hand weak, proximal article with distinct dorsodistal and indistinct dorsomedian boss, no ventral tubercle.
Palps (Fig. 32): Ventral side of femur with small proximal process; trochanter with basally wide, distad-inclined ventral process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus II about 2 times longer than wide (Fig. 33).
Penis (Figs 23-28; holotype: 27, 28): Truncus fairly slender, continually widening towards apex, with almost straight distal margin. Glans distinctly remote from tip of truncus, with quadrangular membraneous socket; short, widely rounded
58 P. J. SCHWENDINGER & J. MARTENS
Fic. 35
Gnomulus monticola sp. n., d holotype. - Body, dorsal (a), ventral (b) and lateral view (c). - Scale line 1.0 mm.
median plate covering membraneous tubes; lateral sclerites sickle-shaped, bent away from the truncus, their tips almost touching each other; stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
2. As the male, no external sexual dimorphism discernible.
FURTHER NEW SPECIES OF GNOMULUS 59
Measurements: 3 holotype (2 in parentheses): Body 3.38 (3.60) long, 2.32 (2.54) wide; carapace region 0.79 (0.75) long, 1.29 (1.32) wide. - Palp and legs:
fie Fe Pa Ti Mt Ta Total Palp 0.41 (0.41 0.47 (0.47) 0.38 (0.39) 0.27 (0.27) SÈ 0.56 (0.56) 2.09 (2.10) Leg I 0.38 (0.38 0.86 (0.89) 0.50 (0.53) 0.52 (0.53) 0.75 (0.75) 0.58 (0.58) 3.59 (3.66)
Leg HI 0.38 (0.38) 0.86 (0.91) 0.53 (0.57) 0.58 (0.60) 0.97 (0.97) 0.44 (0.44) 3.76 (3.87)
) ) ) Leg II 0.47 (0.47) 1.16(1.21) 0.69 (0.71) = 0.79 (0.82) 1.16 (1.16) 0.69 (0.69) 4.96 (5.06) ) ) Leg IV 0.44 (0.47) 1.22(1.27) 0.69(0.72) 0.88 (0.39) 1.44 (1.44) 0.50 (0.50) 5.17 (5.29)
Variation: Range of measurements in dd (n=4) and 9 2 (n=5; in paren- theses): Body 3.31-3.38 (3.39-3.75) long, 2.32-2.36 (2.32-2.59) wide, carapace region 0.72-0.79 (0.72-0.79) long, 1.27-1.29 (1.29-1.37) wide. There is only very little variation in the shape of the eye tubercle and of the ventral processes on palpal trochanter and femur. The penes possess a slightly arched or slightly invaginated distal margin (Figs 25, 27, 29). The holotype has a circular constriction above the base of its truncus penis. This is absent in one d and developed into a small circular fold [as otherwise only observed in the lectotype of G. sumatranus (Schwendinger & Martens, 1999b: figs 70, 71)] in the other d (Figs 23, 24).
Relationships: External and genital morphology show that G. monticola sp. n. is most closely related to G. asli, which occurs in the lowlands at the foot of the Cameron Highlands.
Distribution and bionomics: Known only from the Cameron Highlands in the western part of peninsular Malaysia [Fig. 1 (13)]. The specimens were sifted from leaf litter of a montane rain forest.
Gnomulus pilosus sp. n. Figs 36-45
Material: MALAYSIA (peninsula), Pahang, Taman Negara (= National Park), Tembeling Trail, 90-120 m, 4 holotype, 10./13.IIT.1993, leg. I. Löbl & F. Calame (MHNG).
Etymology: Latin: pilosus = hairy.
Diagnosis: Similar to G. hirsutus, distinguished by: Hair cover less dense; colour pattern on ventral scutum different; anterior dorsal scutal margin less rounded; teeth of carapace-abdomen bridge more widely separated; ventral process on palpal femur smaller; glans penis with a shorter, more V-shaped median plate and more strongly converging lateral sclerites with a wider base.
Description: 3 (holotype). Coloration: Body amber, with dark reticulation in carapax region and with dark pattern on dorsal (shaped as in G. hirsutus) and ventral scuta. Genital operculum dark. Palps and chelicerae light amber, with a dark reticulation (except on palpal tarsus). Legs mostly dark brown, with a light circular median band on metatarsi HI and IV and (less distinct) on femur IV; tarsi light amber, with distitarsus I dorsally darkened.
Carapace with rounded eye tubercle, no lateral tubercles. Left and right processes of carapace-abdomen bridge distinctly separated from each other (Fig. 42). Dorsal and ventral scutal areas moderately elevated (Fig. 40). Palpal coxa with distinct ventral process; leg coxa I with widely triangular anterolateral process; ventral side of leg coxae II and III with conical anteroproximal processes, the latter
60 P. J. SCHWENDINGER & J. MARTENS
36 AE
D a
Fics 36-45
Gnomulus pilosus sp. n., & holotype. - Penis, dorsal (36) and lateral view (37); apex of penis, dorsal (38) and lateral view (39). Body, lateral (40) and ventral view (41); anterior body, dorsal view (42); left chelicera, retrolateral view (43); left palp, retrolateral view (44); distal part of left leg II, retrolateral view (45). - Scale lines 0.1 mm (38, 39), 1.0 mm (others).
overlapped by rounded posteroproximal process on coxa II. Genital operculum slightly wider than long; a distinct tubercle present on posterior margin of stigmatic pit (Fig. 41). Whole body, except for carapace region, covered by fine hairs (Fig. 40).
Chelicerae (Fig. 43): Hand weak, proximal article with distinct dorsodistal and indistinct dorsomedian boss and with indistinct retroventral tubercle.
Palps (Fig. 44): Ventral side of femur with small rounded proximal process; trochanter with distad-inclined ventral process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus II 2.1 times longer than wide (Fig. 45).
Penis (Figs 36-39): Truncus continually widening towards apex, with widely arched, indistinctly invaginated distal margin. Glans quite remote from tip of truncus, with rounded membraneous socket and short, widely V-shaped median plate covering membraneous tubes; lateral sclerites sickle-shaped, their bases wide (as seen in lateral view; Fig. 39), distal parts bent away from the truncus and towards each other; stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
2. Unknown.
Measurements: 3: Body 3.45 long, 2.32 wide; carapace region 0.74 long, 1.28 wide. - Palp and legs:
FURTHER NEW SPECIES OF GNOMULUS 61
Ihe Fe Pa Ti Mt Ta Total Palp 0.38 0.50 0.39 027 - 057 DA Leg I 0.38 0.98 0.55 OS 0.82 0.66 3.96 Leg I 0.49 1932 Oil 0.88 1.34 0.71 5.45 Leg HI 0.39 0.98 0.58 0.61 OF 0.46 4.09 Leg IV 0.49 1.39 (07/7 0.96 1.61 0.53 DIS
Relationships: Externally the new species is very similar to G. hirsutus (especially in hair cover and dorsal colour pattern), but penis morphology indicates a closer relationship with G. monticola sp. n.
Distribution and bionomics: Known only from the type locality in the southern part of Taman Negara, in the centre of peninsular Malaysia [Fig. 1 (14)]. The type specimen was sifted from leaf litter of a lowland rain forest.
THE ROSTRATOIDEUS-GROUP (new)
Diagnosis: Medium-sized (about 5.5 mm) species with strongly forward- inclined, beak-like eye tubercle and distinct carapace-abdomen bridge; ventral process on palpal trochanter distad-directed; dark margin around dorsal scutum unbroken; posterior margin of stigmatic pit without tubercle; stylus penis with invaginated base and without subterminal ventral teeth.
Species account and distribution: This species group is represented only by G. rostratoideus sp. n. from the southern end of the Malay Peninsula.
Gnomulus rostratoideus sp. n. Figs 46-55
Material: MALAYSIA (peninsula), Kota Tinggi Waterfalls, 170 m, at the foot of Gunung Muntahak, ca. 15 km NW of Kota Tinggi, Johor, & holotype, 5.11.2000, leg. P. J. Schwendinger. - SINGAPORE, Bukit Timah Nature Reserve, 1 9 (not a type), 4.VII.1969, leg. D. H. Murphy; same locality, Jungle Fall Valley, 100 m, 1 ® (not a type), 9.VI.2001, leg. P. Schwendinger. All specimens in MHNG.
Etymology: The latinized Greek suffix -oideus refers to similarities with G. rostratus Thorell.
Diagnosis: Externally similar to G. rostratus Thorell, distinguished by a nar- rower eye tubercle (in dorsal view), by a more arched dorsal and ventral scutum and by an unbroken dark margin around the dorsal scutum. Penis very different in shape: Truncus more slender, distal margin widely rounded; glans with convex pincer-like lateral sclerites ending in undivided apices, with a rounded median plate covering short membraneous tubes, and with a thin, tubular stylus without subterminal ventral teeth.
Description: 3 (holotype). Coloration: Body amber, with dark brown reticu- lation on carapace, chelicerae, pedipalps and ventral side of prosoma. Abdominal part of dorsal scutum framed by an unbroken dark margin; dark transversal bands on scutal elevations, medially interconnected in areas III-V (Figs 50, 51). Legs mostly dark brown; tarsi light brown, with reddish grey on proximal part of dorsal distitarsi I and II. Ventral scutal areas with faint dark transversal bands; genital operculum dark reddish brown.
62 P. J. SCHWENDINGER & J. MARTENS
TSE
Sa
NM
IE
Fics 46-55
Gnomulus rostratoideus sp. n., 4 holotype (46-54), 2 (55). - Penis, dorsal (46) and lateral view (47); apex of penis, dorsal (48) and lateral view (49). Body, dorsal (50) and lateral view (51); left chelicera, retrolateral view (52); left palp, retrolateral view (53); distal part of left leg II, retrolateral view (54). Anterior body, chelicera and proximal palp, lateral view (55). - Scale lines 0.1 mm (48, 49), 1.0 mm (others).
Carapace with large pointed, anteriad-inclined eye tubercle and with a rounded hump behind it; no lateral tubercles present; carapace-abdomen bridge composed of two opposing pairs of widely separated teeth (Figs 50, 51). Abdominal part of dorsal scutum moderately arched; ventral and dorsal scutal areas only slightly elevated, the ventral ones bearing “encrusted” hairs (Fig. 51). Palpal coxa with large ventral pro- cess; ventral side of leg coxa I with distinct anterolateral process; ventral side of leg coxa II with pronounced anteroproximal and posteroproximal processes, coxa II with distinct anteroproximal one. Genital operculum as long as wide, anteriorly rounded; posterior margin of stigmatic pit without tubercle but with distinct ledge.
Chelicerae (Fig. 52) weak; proximal article with dorsodistal to dorsomedian boss (distally low) and without ventral tubercle. Cheliceral fingers very long and slender.
Palps (Fig. 53): Ventral side of femur with small proximal process and bulging distal margin; trochanter with distad-directed ventral process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus of leg II 2.2 times longer than wide (Fig. 54).
FURTHER NEW SPECIES OF GNOMULUS 63
Penis (Figs 46-49): Truncus penis fairly slender, continually widening from base to the height of glans, narrower between middle of glans and widely rounded distal margin. Glans penis with large membraneous socket, about as wide as truncus at that point; lateral sclerites convex, sickle-shaped, with acutely pointed tips crossing each other; membraneous tubes short, completely covered by a rounded, U-shaped median plate; stylus long and slender, its base slightly invaginated, its apex without subterminal ventral teeth.
2 (Identification uncertain). As the male, but eye tubercle less elevated and more pointed in dorsal view, region behind eye tubercle less elevated, abdominal part of dorsal scutum less arched, ventral process on palpal trochanter shorter, more knob- shaped (Fig. 55).
Measurements: 3 holotype (9 in parentheses): Body 5.48 (5.43) long, 3.48 (3.33) wide; carapace region 1.70 (1.50) long, 2.06 (1.90) wide. - Palp and legs:
Ihr Fe Pa Ti Mt Ta Total Palp 0.63 (0.51) 1.15 (0.91) 0.79 (0.65) 0.61 (0.53) - - 1.47 (1.31) 4.65 (3.91) Leg I 0.63 (0.53) 1.74 (1.43) 0.83 (0.75) 1.03 (0.87) 1.50(1.27) 0.83 (0.69) 6.56 (5.54) Leg II 0.75 (0.71) 2.30(1.96) 1.19(1.17) 1.66 (1.39) 2.30 (1.90) 1.03 (0.87) 9.23 (8.00) Leg III 0.55 (0.55) 1.72 (1.39) 0.91 (0.75) 1.07 (0.87) 1.72 (1.50) 0.61 (0.55) 6.58 (5.61) RENE 077.067) 7238.98) 121 (07) 1.62.1.39)7 72.65. 228) 0.75.(0:63), 9/58 (8/02)
Variation: The second 2 measures: Body length 5.26, width 3.22, carapace length 1.49, width 1.78. Both © from Singapore are smaller than the d from Malay- sia and differ in some details of external morphology. These specimens may therefore not be conspecific, but the common presence of an unbroken dark dorsal scutal margin, a fairly distinct hair cover and geographical proximity clearly show that they are more closely related with each other than with any of the species of the rostratus- group. Until this uncertainty is solved by the discovery of a d from Singapore, we tentatively place the 9 9 examined in G. rostratoideus sp. n., but we do not designate them as paratypes.
Relationships: Gnomulus rostratoideus sp. n. appears to be the sister taxon of the rostratus-group. Congruence in penis morphology with the species of the good- nighti-group is considered to be due to convergence (see discussion).
Distribution and bionomics: Known from rainforests in the south of peninsular Malaysia [Fig. 1 (15)] and on nearby Singapore Island [Fig. 1 (16)].
THE SUMATRANUS-GROUP (see Schwendinger & Martens, 1999b: 957)
Diagnosis: The new species possesses a distinctly elevated eye tubercle and posteroproximal processes on coxae II, but no transversal keels on its dorsal scutum. Therefore the diagnosis of this group has to be modified. The sumatranus-group is essentially characterized by: Body large (6.8-9.0 mm); chelicerae robust (at least in the S), with a distad-inclined proventral tooth (pointed process) on proximal article; a subdistal process present on ventral side of palpal femur (indistinct in some females); proximal process on ventral side of palpal femur slightly to distinctly distad-directed; distodorsal tubercle present on palpal trochanter; posterior margin of stigmatic pit with tubercle; lateral sclerites on glans penis cylindrical, acutely pointed, only slightly bent; stylus penis with ventral pair of subterminal teeth and bulbous base.
64 P. J. SCHWENDINGER & J. MARTENS
Species account and distribution: This group comprises two species from Sumatra, i.e. G. sumatranus Thorell and G. tuberculatus sp. n.
Gnomulus tuberculatus sp. n. Figs 56-73
Material: INDONESIA, Sumatra, Aceh Province, Gunung Leuser National Park, Ketambe Research Station, 300-500 m, 4 holotype (MHNG), 1 4,3 © paratypes, 3 juv., 23.- 30.X1.1989, leg. I. Lobl, D. Agosti & D. Burckhardt (1 © paratype in MAR, others in MHNG).
Etymology: Latin: tuberculatus = tuberculate. The specific epithet refers to the presence of two tubercles on the dorsal side of leg coxa IV and of one tubercle on the anterolateral side of trochanter III.
Diagnosis: Close to G. sumatranus, distinguished by: Body smaller; carapace shorter in the Sd, with a distinct eye tubercle in both sexes; ventral side of leg coxa II with an anteroproximal process, process on coxa III larger; trochanter HI with a small prolateral tubercle; proximal process on ventral side of palpal femur smaller, not distad-inclined, subdistal process indistinct; distitarsus II shorter; penis without circu- lar fold around subbasal truncus; distal margin of truncus less arched; two setae present on each side of glans penis; tips of lateral sclerites slightly inclined towards each other.
Description: 3 (holotype). Coloration: Body amber, with dark reticulation on carapace, chelicerae and pedipalps; dorsal scutal elevations dark brown, separated by a light median, longitudinal, partly broken stripe in areas I-IV and by pairs of light transversal stripes ending in light paramedian patches (Fig. 73a, c). Legs dark brown; tarsi I, II cream (slightly darkened on dorsal side of distitarsus I), tarsi II, IV light brown.
Carapace with conical, distally rounded eye tubercle and an indistinct pair of lateral tubercles below wide undivided carapace-abdomen bridge. Dorsal scutal areas only slightly elevated; ventral scutal areas distinctly swollen, without hairs (Figs 73a, c). Palpal coxa with distinct ventral process; leg coxa I with long, outwards-inclined anterolateral one; ventral leg coxae II and III with distinct anteroproximal processes, coxa II additionally with low posteroproximal one (Fig. 73b); dorsal side of leg coxa IV with two knob-shaped tubercles (Figs 69-72, 73c). Genital operculum about as long as wide; rounded tubercle on posterior margin of stigmatic pit (Fig. 73b).
Chelicerae (Figs 62, 63): Hand strong, cutting edges of fingers each with a strong subbasal tooth; proximal article with forward-inclined dorsodistal to dorso- median boss and with two distad-inclined proventral teeth (the subbasal one smaller than the subterminal one; Fig. 63).
Palps (Fig. 66): Ventral side of femur with indistinct subdistal and distinct proximal process; trochanter with low dorsal tubercle and with small, slightly distad- inclined ventral process.
Legs 1324, tarsal formula 2-2-3-3. Trochanter of leg III with prolateral tubercle (Fig. 67); distitarsus II about 2.2 times longer than wide (Fig. 68).
Penis (Figs 56-61; holotype: 60, 61): Truncus fairly slender, with very widely arched distal margin; membraneous socket of glans penis laterally bordered by two pairs of setae. Glans with subtriangular median plate partly covering membraneous tubes; distal portion of lateral sclerites cylindrical, acutely pointed, not sigmoid and only moderately bent away from the truncus, tips slightly inclined towards each other: stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
FURTHER NEW SPECIES OF GNOMULUS 65
VA TR
Fics 56-72
Gnomulus tuberculatus sp. n., 4 holotype (60, 61, 62, 63, 66-68), 4 paratype (56-59, 72), 9 paratypes (64, 65, 69-71). - Penis, dorsal (56) and lateral view (57); apex of penis, dorsal (58, 60) and lateral view (59, 61). Left chelicera, retrolateral view (62, 64); proximal article of left chelicera, prolateral view (63, 65); left palp, retrolateral view (66); trochanter of left leg III, dorsal view (67); distal part of left leg II, retrolateral view (68); anterior body and proximal palp, lateral view (69-72). - Scale lines 0.1 mm (58-61), 1.0 mm (others).
2. As the male but coloration generally more reddish; carapace region slightly smaller; chelicerae weaker, subbasal tooth on cutting edge of both cheliceral fingers indistinct, no dorsomedian boss on proximal article (Figs 64, 65); subdistal process on
66 P. J. SCHWENDINGER & J. MARTENS
ventral side of palpal femur reduced, hardly discernible (Figs 69-71); ventral scutal areas not swollen.
Measurements: 3 holotype (© in parentheses): Body 6.83 (6.90) long, 4.80 (4.70) wide; carapace region 1.83 (1.39) long, 2.77 (2.43) wide. - Palp and legs:
Tr Fe Pa Ti Mt Ta Total Palp 0.99 (0.79) 1.63 (1.29) 1.04 (0.84) 0.77 (0.59) -- 1.88 (1.53) 6.31 (5.04) Leg I 0.79 (0.69) 2.57(2.40) 1.14(1.04) 1.29(1.19) 2.28 (2.05) 1.06 (0.99) 9.13 (8.36) LegII 0.94(0.84) 3.32(3.17) 1.56(1.41) 2.18(2.03) 3.42 (3.22) 1.29(1.29) 12.71 (11.96) Leg IN 0.74 (0.69) 2.52 (2.38) 1.19(1.14) 1.44 (1.34) 2.70 (2.57) 0.69 (0.79) 9.28 (8.91) Leg IV 1.01 (0.99) 3.27(3.17) 1.58 (1.48) 2.23 (2.13) 4.11 (3.98) 0.94 (0.89) 13.14 (12.64)
Variation: Range of measurements in Sd (n=2) and 2 2 (n=3; in paren- theses): Body 6.83-7.05 (6.90-7.18) long, 4.80-4.93 (4.60-4.95) wide, carapace region 1.73-1.83 (1.39-1.46) long, 2.77-2.82 (2.43-2.52) wide. Most specimens have a narrowly rounded eye tubercle (Figs 69, 70, 72, 73c), in one @ it is almost pointed (Fig. 71).
Relationships: Genital morphology and certain characteristics of chelicerae, palps and leg coxae I indicate that Gnomulus tuberculatus sp. n. is most closely related to G. sumatranus.
Remark: The eye tubercles of the juvenile specimens and of a female were illustrated in Schwendinger & Martens (1999b: figs 135-137).
Distribution: Known only from the type locality in northern Sumatra [Fig. 1 (17)].
THE ARMILLATUS-GROUP (see Schwendinger & Martens, 1999b: 958)
Diagnosis: The following characteristics are added to the diagnosis of this group of medium-sized to large (5.3-8.6 mm) species: Ventral side of palpal trochanter with ventrad- or slightly distad-directed process; posterior margin of stigmatic pit with distinct (often pronounced) tubercle; stylus with a ventral pair of subterminal teeth and with a bulbous base.
Species account and distribution: 20 species are known at present: One from Myanmar (G. leofeae sp. n.), two from Thailand (G. marginatus sp. n., G. ryssie sp. n.), two from peninsular Malaysia [G. piliger (Pocock), G. pulvillatus (Pocock)], two from Sumatra [G. armillatus (Thorell), G. drescoi (Silhavy)]. two from Java [G. javanicus Sp. n., G. thorelli (Sgrensen) (male unknown, uncertain assignment)] and 10 from Borneo [G. annulipes (Pocock), G. baharu Schwendinger, G. carinatus sp. n., G. conigerus (Schwendinger), G. exsudans sp. n., G. hutan sp. n., G. laevis (Roewer), G. lomani sp. n., G. obscurus sp. n., G. sundaicus (Schwendinger)] and one from Palawan, the Philippines [G. palawanensis (Suzuki), male unknown, uncertain assignment].
Gnomulus marginatus sp. n. Figs 74-91
Material: THAILAND, Chanthaburi Province and District, Nam Tok Phliu - Khao Sabap National Park, near Phliu Waterfall, 50 m, & holotype, 2 d, 1 © paratypes, 12.X1.1998. - Trat Province, Laem Ngop District, Ko Chang (= Elephant Island) National Park, forest above White Sand Beach, 20-50 m, 3 d paratypes, 6.-8.IX.1993, 1 juv., 24.VIII.1992 (all specimens
FURTHER NEW SPECIES OF GNOMULUS 67
E
Fic. 73
Gnomulus tuberculatus sp. n., 3 holotype. - Body, dorsal (a), ventral (b) and lateral view (c). - Scale line 1.0 mm.
leg. P.J. Schwendinger); same island, west side, 12°03’N, 102°18’E, 50-200 m, 1 & paratype, 3.-23.X11.1999 (leg. A. Schulz). 16 paratype in MAR, others in MHNG.
Etymology: Latin: marginatus (adjective of margo) = framed. The specific epithet refers to the conspicuous dark marginal and light submarginal band around the abdominal part of the dorsal scutum.
68 P. J. SCHWENDINGER & J. MARTENS
SAX eh
4 75 74 L N —=S
Gnomulus marginatus sp. n., © holotype (74-77, 85-88), d paratype (78-82, 84, 90), 9 paratypes (83, 89). - Penis, dorsal (74) and lateral view (75); apex of penis, dorsal (76, 78) and lateral view (77, 79); glans penis, dorsal view (80-82). Left chelicera, retrolateral view (83-85); left palp, retrolateral view (86); trochanter and femur of left palp, dorsal view (87); distal part of left leg II, retrolateral view (88); anterior body and proximal palp, lateral view (89, 90). - Scale lines 0.1 mm (76-82), 1.0 mm (others).
Fics 74-90
Diagnosis: Similar to G. armillatus, distinguished by: Light submarginal band on abdominal part of dorsal scutum; retroventral tooth on proximal cheliceral article more distinct and pointed; ventral processes on palpal femur and trochanter smaller; two tubercles on dorsal side of leg coxa IV and one on lateral side of abdominal
FURTHER NEW SPECIES OF GNOMULUS 69
Fic. 91
Gnomulus marginatus sp. n., d holotype. - Body, dorsal (a), ventral (b) and lateral view (c). - Scale line 1.0 mm.
segment II; penis more stout, glans wider, its lateral sclerites more convex and basally more elevated, its median plate longer, tongue-shaped.
Description: 3 (holotype). Coloration: Body brown, dorsal scutum with dark brown pattern and clearly outlined red-brown margin below light brown submarginal
70 P. J. SCHWENDINGER & J. MARTENS
band (Fig. 91a, c); ventral scutal elevations pallid (Fig. 91b); legs dark brown, except for light yellow-brown tarsi (tarsus I dorsodistally darkened); dark reticulation on brown carapace and palps and on light brown chelicerae.
Carapace with conical, pointed eye tubercle and with a pair of lateral tubercles posteriorly below wide, indistinctly divided carapace-abdomen bridge. Dorsal scutal areas only slightly elevated, divided by a shallow median furrow in areas I-IV; ventral scutal areas distinctly swollen, no hairs present (Fig. 91a, c). Palpal coxa with large ventral process; leg coxa I with distinct anterolateral process; ventral side of leg coxae II and III with distinct anteroproximal processes, coxa II also with indistinct postero- proximal one (Fig. 91b); dorsal side of leg coxa IV with two knob-shaped tubercles; one more such tubercle posterior to them, on the lateral side of abdominal segment II (Fig. 91c). Genital operculum somewhat triangular, wider than long; posterior margin of stigmatic pit with a distinct rounded tubercle (Fig. 91b).
Chelicerae (Fig. 85): Hand weak, proximal article fairly strong, with dorso- distal to dorsomedian boss and distinct retroventral tooth.
Palps (Figs 86, 87): Femur with low prodorsal median boss (Fig. 87) and indistinct ventroproximal process; trochanter with low ventral process.
Legs1324, tarsal formula 2-2-3-3. Distitarsus of leg II about 2.3 times longer than wide (Fig. 88).
Penis (Figs 74-82; holotype: 74-77): Truncus fairly stout, slightly constricted below glans, with widely arched distal margin and plenty of subapical setae. Glans with tongue-shaped median plate covering membraneous tubes; lateral sclerites with moderately elevated dorsal ledge in proximal part, distal part slender, sigmoid, cylindrical, pointing away from the truncus, tapering tips widely apart; stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
2. As the male but dark pattern on dorsal scutum less pronounced, ventral scutal elevations not pallid; eye tubercle slightly smaller and less pointed (possibly an individual variation; Fig. 89); proximal article of chelicerae more slender (Fig. 83).
Measurements: 3 holotype (@ in parentheses): Body 6.74 (7.08) long, 4.82 (4.97) wide; carapace region 1.48 (1.38) long, 2.61 (2.58) wide. - Palp and legs:
Tr Fe Pa Hi Mt Ta Total Palp 0.89 (0.89) 1.48 (1.40) 1.08 (1.08) 0.81 (0.79) - - 159153) 5.81 (5.69) Leg I 0.69 (0.64) 2.46 (2.41) 1.16(1.11) 1.25(1.28) 2.16(2.12) 0.98 (0.93) 8.70 (8.49) Keel (0:89'0:86) 3:17 6-15). 1.48:0.48) 221 (2.12) 3257820) 133.029) 7233702209) Leg UI 0.79 (0.74) 2.51 (2.41) 1.21(1.21) 1.43 (1.38) 2.56 (2.48) 0.79 (0.74) 9.29 (8.96) Leg IV 0.93 (0.93) 3.20 (3.15) 1.53 (1.48) 2.19 (2.09) 3.89 (3.81) 0.93 (0.93) 12.67 (12.39)
Variation: Range of measurements in d d (n=6): Body 6.33-6.95 long, 4.65- 4.82 wide, carapace region 1.45-1.56 long, 2.54-2.61 wide. One d paratype with a bifid ventral tooth on proximal cheliceral article (Fig. 84) and with a truncate ventral process on palpal trochanter (Fig. 90).
Relationships: Gnomulus marginatus sp. n. clearly belongs to the armillatus- group; it appears most closely related to G. piliger, G. pulvillatus from the Malayan Peninsula and to G. armillatus from Sumatra. Striking similarities in external and penis morphology are also evident between G. marginatus sp. n. and G. annulipes from Sarawak. These are regarded as convergences.
FURTHER NEW SPECIES OF GNOMULUS 71
Distribution and bionomics: Known from two localities (separated by about 50 km) in southeastern Thailand, one on the mainland [Fig. 1 (5)] and the other on an island ca. 6 km off the coast [Fig. 1 (6)]. The animals were found under decaying wood on the forest floor of a semi-evergreen rain forest (at Nam Tok Phliu) and of a secondary forest adjacent to primary forest (on Ko Chang).
Gnomulus ryssie sp. n. Figs 92-101
Material: THAILAND, Phetchaburi Province, Kaeng Krachan National Park, 300-400 m, 25-30 km W of park headquarters, d holotype, 17.X1.1985, leg. I. Lòbl & D. Burckhardt (MHNG).
Etymology: Ryssie is a forest-dwelling hermit in Thai (originally Hindu) mythology. Noun in apposition.
Diagnosis: Closest to G. marginatus sp. n., distinguished by: Eye tubercle larger; abdominal region higher; no ventral process on proximal article of chelicerae; no prodorsal boss on palpal femur; ventral process on palpal trochanter smaller; no posterior tubercle on dorsal side of coxa IV; lateral tubercles on abdominal segment II indistinct; penis more slender, with fewer subapical setae; glans with longer, sub- triangular median plate; lateral sclerites less distinctly elevated above median plate.
Description: 3 (holotype). Coloration: Body light amber, with dark reticu- lation in carapace region, on proximal article of chelicerae and on femur to tarsus of pedipalps; dorsal scutal elevations dark (Fig. 96), ventral ones pallid; legs and palps dark amber, all tarsi and distal portion of leg tibiae III and IV light brown; dorsal side of distitarsus I darkened.
Carapace with strong conical eye tubercle and a small pair of lateral tubercles below wide, medially divided carapace-abdomen bridge. Abdominal part of dorsal scutum high; scutal areas slightly elevated, medially separated by a shallow longitudinal furrow in anterior part of abdominal region (Figs 96, 97); ventral scutal areas distinctly swollen, without pubescence. Palpal coxa with conical ventral pro- cess; leg coxa I with anterolateral one; ventral side of leg coxae IT and III with distinct anteroproximal processes, coxa II with small posteroproximal one; dorsal side of leg coxa IV with distinct anterior tubercle, posterior one absent; pair of lateral tubercle on abdominal segment II indistinct. Genital operculum wider than long; stigmatic pit with a rounded tubercle on posterior margin.
Chelicerae (Fig. 98): Proximal article with distinct dorsodistal and indistinct dorsomedian boss; no ventral process.
Palps (Figs 99, 100): Femur with indistinct ventroproximal process, no pro- dorsal boss (Fig. 100); trochanter with very small ventral process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus II about 2.2 times longer than wide (Fig. 101).
Penis (Figs 92-95): Truncus slightly constricted below glans, with widely arched distal margin; glans with large subtriangular median plate covering membra- neous tubes; lateral sclerites convex, with slightly elevated ledge in proximal part; distal part slender, sigmoid, tapering, pointing away from the truncus, tips widely apart; stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
2. Unknown.
72 P. J. SCHWENDINGER & J. MARTENS
gi R © NT ORAN 7 pl I)
4, N
=
Fics 92-101
Gnomulus ryssie sp. n., 6 holotype. - Penis, dorsal (92) and lateral view (93); apex of penis, dorsal (94) and lateral view (95). Body, dorsal view (96); anterior body and proximal palp, lateral view (97). Left chelicera, retrolateral view (98); right palp, retrolateral view (99); trochanter and femur of left palp, dorsal view (100); distal part of left leg II, retrolateral view (101). - Scale lines 0.1 mm (94, 95), 1.0 mm (others).
Measurements: (3): Body 6.22 long, 4.38 wide; carapace region 1.13 long, 2.21 wide. - Palp and legs:
the Fe Pa Ji Mt Ta Total Palp 0.69 1.03 0.83 0.59 - IRIS 4.29 Leg I 0.54 1.92 0.89 0.98 1.62 0.79 6.74 Leg II 0.64 Zi 113 ea Dail 0.98 9.54 Leg III 0.57 1.92 0.98 1.18 2.02 0.59 TAS Leg IV 0.74 2:50 1.21 1.80 310 0.69 10.1
Relationships: Gnomulus ryssie sp. n. is closest to G. marginatus sp. n. Distribution: Known only from the type locality in the southwestern part of central Thailand [Fig. 1 (7)].
Gnomulus leofeae sp. n. Figs 102-111
Pelitnus segnipes Loman: Roewer (1935: 13).
Material: MY ANMAR, Tenasserim, Malewoon (= Maliwun), 4 holotype (MSNG, Nr. 10203), leg. L. Fea, VIII-IX 1887.
FURTHER NEW SPECIES OF GNOMULUS 73
Etymology: This species is dedicated to the Italian naturalist Leonardo Fea, who travelled and collected in Myanmar in the years 1885-1888. The genitive ending “-ae” is linguistically correct (and accepted by the International Code of Zoological Nomenclature) for names with a terminal “-a”, also when referring to a man.
Diagnosis: Closest to G. pulvillatus, distinguished by: Eye tubercle lower; lateral tubercles in posterior carapace region present; median dorsal scutal elevations keeled; palpal femur more slender, its ventroproximal process more distad-inclined; leg coxa II without posteroproximal process; penis not constricted at position of glans; glans narrower and less convex in dorsal view, its lateral sclerites longer and more slender; isolated pair of lateral setae on each side of membraneous socket situated more proximally.
Description: 3 (holotype). Coloration: Body and limbs light amber throughout (bleached), except for slightly darker leg tarsi.
Carapace with widely conical, pointed eye tubercle and a pair of small lateral tubercles (Fig. 106) below wide, medially divided carapace-abdomen bridge (Fig. 108). Dorsal scutal areas slightly elevated, the median ones keeled (Fig. 106). Ventral scutum heart-shaped (probably deformed due to preservation), areas swollen, without pubescence (Fig. 107). Palpal coxa with digitiform ventral process; leg coxa I with pronounced anterolateral one; ventral side of leg coxae II and II with distinct anteroproximal processes, no posteroproximal one coxa II; no tubercles on dorsal side of leg coxa IV. Genital operculum somewhat triangular, slightly wider than long; posterior margin of stigmatic pit with pronounced tubercle (Fig. 107).
Chelicerae (Fig. 109): Proximal article with distinct dorsodistal and indistinct dorsomedian boss, ventral side with low, wide hump.
Palps (Fig. 110): Strong, slightly distad-inclined ventral processes on proximal femur and on trochanter.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus II about 2.8 times longer than wide (Fig. 111).
Penis (Figs 102-105): Truncus widest at position of glans, distal margin indistinctly invaginated. Glans with short, widely rounded median plate covering membraneous tubes; lateral sclerites cylindrical, fairly long and slender, their distal parts sigmoid, tapering, pointing away from the truncus; stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
2. Unknown.
Measurements: (4): Body 5.74 long, 4.21 wide; carapace region 1.44 long, 2.13 wide. - Palp and legs:
Ibe Fe Pa Ti Mt Ta Total Palp 0.87 1.24 0.99 0.64 - 1.29 3103 Leg I 0.69 2.45 1.16 1.34 2.08 1.06 8.78 Leg II 0.82 39 1.58 238 DRD. 1.34 12.66 Leg III 072 250 1.29 56 DST 1.14 9.78 Leg IV 0.89 3,32 1.36 2.28 3.86 1.29 13.20
Relationships: Gnomulus leofeae sp. n. is closely related to G. pulvillatus from central peninsular Malaysia and to G. piliger from southern Thailand.
74 P. J. SCHWENDINGER & J. MARTENS
da AS En Ve Dx
- = PEN GES 110 = G \& > Pa CZ LES TS N
Fics 102-111
Gnomulus leofeae sp. n., $ holotype. - Penis, dorsal (102) and lateral view (103); apex of penis, dorsal (104) and lateral view (105). Body, lateral view (106); body, ventral view (107); anterior body, dorsal view (108). Left chelicera, retrolateral view (109); left palp, retrolateral view (110); distal part of left leg II, retrolateral view (111). - Scale lines 0.1 mm (104, 105), 1.0 mm (others).
Remarks: Gnomulus leofeae sp. n., G. piliger and G. pulvillatus are quite simi- lar to each other in external and genital characters, and each species is known only from a single specimen. Additional conspecific material from the Malay Peninsula may show whether these are really three distinct species or just individual or popu- lation-dependent variations of a widely distributed and unusually variable species.
Distribution: Known only from the type locality at the southernmost tip of Myanmar [Fig. 1 (8)].
Gnomulus armillatus (Thorell, 1891) Synonyms: See Schwendinger & Martens (1999: 963).
Remark: A new d specimen was collected on Gunung (= Mount) Kerinci (2160 m, 17.-18.11.2000, leg. P. J. Schwendinger, MHNG), Jambi Province, Sumatra [Fig. 1 (20)], which largely accords with the d from the same locality mentioned by Schwendinger & Martens (1999b: 963, 965, figs 114, 122). The new d also possesses a fairly low, rounded eye tubercle, but its ventral processes on palpal femur and trochanter are larger (more typical for males of this species) and the ventral tubercle on the proximal cheliceral article is indiscernible. The median plate of the glans penis
FURTHER NEW SPECIES OF GNOMULUS 75
is more distinctly V-shaped, with clearly discernible lateral teeth, well-according with conspecific males from close to the type locality (see Schwendinger & Martens, 1999b: figs 110, 112).
Gnomulus javanicus sp. n. Figs 112-124
Pelitnus segnipes Loman. - Loman (1902: 182, partim). - Roewer (1923: 63, partim).
Material: INDONESIA, Java, Mt. Gede, SE of Bogor, d holotype (ZMH, with labels: “Pelitnus segnipes, Loman det. 1901/02, H. Fruhstorfer vend. 18.11.1897” and “Pelitnus segnipes, Roewer det. 1914, No. 1258”). - Java, without exact locality, 2 2 paratypes (SMF 1602, with label: “Pelitnus javanus n. sp. Roewer, typus, 1 4, 1 2, Roewer det. 1929”).
Diagnosis: Similar to G. laevis, distinguished by: Body smaller; palp without prolateral boss and with distinct ventrobasal process on femur; ventral process on palpal trochanter longer; distitarsus II shorter; penis with narrower apex and less strongly arched distal margin; glans narrower, its lateral sclerites less convex, with more distinctly pointed apices; median plate shorter.
Description: 3 (holotype). Coloration mostly dark reddish brown; transversal bands on dorsal scutal elevations slightly darker, medially disconnected in areas I-III; chelicerae and proximal article of pedipalps slightly lighter.
Carapace with low, rounded eye tubercle, without lateral tubercles posteriorly below wide, indistinctly divided carapace-abdomen bridge (Figs 116, 124). Dorsal scutal areas distinctly elevated, anterior ones rounded, posteriors keeled; ventral scutal areas swollen, without modified hairs. Palpal coxa with distinct ventral process; leg coxa I with small anterolateral one; ventral side of leg coxae II and III with distinct anteroproximal processes, coxa II with indistinct posteroproximal one; dorsal side of coxa IV without tubercle. Genital operculum narrow, slightly longer than wide (Fig. 117); posterior margin of stigmatic pit with tubercle.
Chelicerae (Fig. 119) weak, proximal article with distinct dorsodistal and less distinct dorsomedian boss; no ventral tubercle.
Palps (Fig. 124): Femur stout, with strong ventroproximal process and long dorsodistal to dorsoproximal boss; trochanter with long, digitiform, slightly distad- inclined ventral process.
Legs 13(24?), tarsal formula 2-2-3-3. Distitarsi of legs II missing.
Penis (Figs 112-115): Truncus slender, widest below glans, with narrow, slightly arched apex. Glans slightly narrower than truncus at that point; lateral sclerites convex and with a moderately elevated dorsal ledge in proximal half, distal half narrowly paddle-shaped, pointing away from the truncus, both sides parallel to each other; knee between proximal and distal part of lateral sclerites bent at right angles, not bulged towards truncus (Fig. 115); median plate very short, widely rounded, with a pair of rounded lateral teeth; membraneous tubes distally not covered by median plate; stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
2. As the male but with much shorter ventral processes on palpal femur and trochanter; palpal femur without dorsal boss (Figs 122, 123); proximal article of chelicera without dorsomedian boss (Fig. 120); dorsal scutal areas less elevated and
76 P. J. SCHWENDINGER & J. MARTENS
IA:
CAPUA =,
Il dl at x fo ist 120 112 113 Lo
122
Fics 112-124 Gnomulus javanicus sp. n., 3 holotype (112-117, 119, 124), 2 paratypes (118, 120-123). - Penis, dorsal (112) and lateral view (113); apex of penis, dorsal (114) and lateral view (115). Anterior body and chelicerae, dorsal view (116); genital operculum (117, 118); left chelicera, retrolateral view (119, 120); distal part of left leg II, retrolateral view (121); anterior body and proximal palp, lateral view (122-124). - Scale lines 0.1 mm (114, 115), 1.0 mm (others).
ventral scutal areas not swollen. Legs 1324; distitarsus II about 2.1 times longer than wide (Fig. 121).
Measurements: 3 holotype (@ in parentheses): Body 5.86 (5.92) long, 4.05 (4.02) wide; carapace region 1.25 (1.08) long, 2.23 (2.10) wide. - Palp and legs:
‘Abe Fe Pa Bl Mt Ta Total
Palp 0.81 (0.69) 1.12 (1.04) 0.94 (0.74) 0.69 (0.54) - - 1.18 (1.08) 4.74 (4.09) Leg I 0.56 (0.54) 2.24 (1.68) 1.06 (0.89) 1.31 (1.04) 1.93 (1.63) 0.94 (0.84) 8.04 (6.62) Leg II OGIO) 24 PS2) ESS AIG) 23763) 8242820 (1.13) ? (9.20) ) )
Leg Ill 0.69(0.59) 2.43(1.75) 1.18 (0.99) 1.62 (1.16) 2.37(1.92) 1.00(0.84) 9.29 (7.25) LegIV 1.03 (0.89) 3.18 2.42) 1.50 (1.21) 2.24(1.77) 3.58(2.91) 1.18 (0.94) 12.71 (10.14)
Variation: Range of measurements in 2 2 (n=2): Body 5.67-5.92 long, 3.86- 4.02 wide, carapace region 1.08-1.18 long, 1.99-2.10 wide. One 9 has a very low eye tubercle (Fig. 122 ).
FURTHER NEW SPECIES OF GNOMULUS ANG
Relationships: Gnomulus javanicus sp. n. is closest to G. lomani sp. n.
Remarks: The specimens examined are clearly distinct from G. thorelli, which also occurs on Java. However, males and females of G. javanicus sp. n. (both from the same island) differ from each other in a number of characters (see above), which either reflect a pronounced sexual dimorphism or indicate that they belong to different species. Until further evidence for the contrary becomes available, these specimens are regarded as conspecific.
As far as we know “Pelitnus javanus Roewer” has never been described and was never mentioned in the literature. It thus is an unpublished name without nomenclatural relevance.
Distribution: Known only from one or two localities on Java [Fig. 1 (21)]. The exact locality of the presumably conspecific female paratypes is unknown. Gnomulus thorelli was found at Cibodas (Schwendinger & Martens, 1999b: 969) and possibly occurs syntopically with G. javanicus sp. n.
Gnomulus lomani sp. n. Figs 125-140
Pelitnus segnipes Loman. - Loman (1902: 182, partim). - Roewer (1923: 63, partim).
Material: BORNEO, Telang (locality not identified), 4 holotype, 8 paratype (ZMB 4247), with one label saying “Pelitnus segnipes Loman, 1892, Fundort (= find locality): Telang, Broneo (Burma)” and another in Gothic handwriting (probably by Roewer) “Pelitnus segnipes Loman, von Loman falsch bestimmt (= misidentified by Loman)’; both specimens leg. F. Grabonsky. - SUMATRA, BORNEO, without exact locality, 2 d, 1 2, “leg. ?, leg. Schwaner?” (ZMA; no types).
Etymology: The species is dedicated to Jan Cornelis Christiaan Loman, a Dutch zoologist of outstanding merit, who published on opilionids from 1879 to 1910.
Diagnosis: Very similar to G. javanicus sp. n., distinguished by the presence of small tubercles on the dorsal side of leg coxa IV and by details of penis morphology, i.e. apex of truncus wider; glans wider than truncus at that point; knee between pro- ximal and distal part of lateral sclerites more bulged towards the truncus (as seen in lateral view); apices of lateral sclerites more widely apart; membraneous tubes enti- rely covered by longer, more V-shaped median plate with less distinct pair of lateral teeth.
Description: 3 (holotype). Coloration: Body dark amber, limbs light amber, with tarsi and distal portions of leg articles slightly lighter; pattern on dorsal and ventral scutum faded.
Carapace with quite low, conical eye tubercle and without lateral tubercles posteriorly below wide, indistinctly divided carapace-abdomen bridge (Figs 136, 137). Dorsal scutal areas distinctly elevated, anterior ones rounded, posteriors keeled; ventral scutal areas swollen, without modified hairs. Palpal coxa with strong ventral process; leg coxa I with small anterolateral one; ventral side of leg coxae II and II with distinct anteroproximal processes, no posteroproximal one on coxa II; dorsal side of coxa IV with a small anterior tubercle. Genital operculum about as long as wide; posterior margin of stigmatic pit with low tubercle.
Chelicerae (Fig. 134) weak, proximal article with distinct dorsodistal to dorsomedian boss; no ventral process.
78 P. J. SCHWENDINGER & J. MARTENS
CH 5
WR
a
n)
tA GE 125 126 A DI è = N
Jl sl Am FT]
Fics 125-140
Gnomulus lomani sp. n., & holotype (125-128, 134, 136, 137), 2 paratypes (129-130, 139), presumably conspecific & (138) and © (133, 140). - Penis, dorsal (125) and lateral view (126); apex of penis, dorsal (127, 129, 131) and lateral view (128, 130); glans penis, lateral view (132). Left chelicera, retrolateral view (133, 134); distal part of left leg II, retrolateral view (135); anterior body, dorsal view (136): anterior body and proximal palp, lateral view (137- 140). - Scale lines 0.1 mm (127-132), 1.0 mm (others).
FURTHER NEW SPECIES OF GNOMULUS 79
Palps (Fig. 137): Femur with strong ventroproximal process and extended dorsodistal to dorsoproximal boss; trochanter with long, digitiform ventral process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus II 2.9 times longer than wide (Fig. 135).
Penis (Figs 125-132; holotype: 125-128): Truncus slender, continuously widening towards apex; distal margin broadly arched. Glans wider than truncus at that point; lateral sclerites convex and with a slightly elevated dorsal ledge in proximal half, distal part narrowly paddle-shaped, pointing away from the truncus (Fig. 128), both sides parallel to each other; knee between proximal and distal half distinctly bulged towards the truncus; apices of lateral sclerites widely apart; median plate quite short, more or less distinctly V-shaped, distally rounded, with lateral teeth reduced to low bulges; membraneous tubes completely covered by median plate; stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
2 (uncertain identification). As the maie but with a more slender palpal femur, distinctly smaller ventral processes on palpal trochanter and femur and with a smaller dorsomedian boss on the proximal cheliceral article.
Measurements: 3 holotype (d paratype in parentheses; no measuremets of available © given because of uncertain identification): Body 5.70 (5.67) long, 4.05 (3.99) wide; carapace region 1.19 (1.19) long, 2.22 (2.20) wide. - Palp and legs:
Tr Fe Pa Ti Mt Ta Total Palp 0.78 1.06 0.90 0.65 - 112 4.51 Leg I 0.56 2.06 1.06 125 1.87 0.94 1.14 Leg II 0.72 2.99 1.56 2.28 2.99 1.31 11.85 Leg III 0.69 2.24 1.22 1.50 2,31 1.00 8.96 Leg IV 0.97 2.81 1.50 212 3.40 1.18 11.98
Variation: The 4 paratype has indistinct posteroproximal processes on coxa II, more strongly elevated dorsal scutal areas (Fig. 139) and a longer median plate on glans penis (Fig. 129).
Relationships: Gnomulus lomani sp. n. is closest to G. javanicus sp. n.; both are in the same phyletic lineage with G. exsudans sp. n., G. hutan sp. n., G. laevis, G. obscurus sp. n. and G. sundaicus.
Remarks: The indication of the type locality [“Telang, Broneo (Burma)”], as given on the label with the types, is misleading. According to information from Jason Dunlop, the ZMB houses extensive material collected by Fritz Grabonsky in Borneo, but none from Burma. Broneo was obviously misspelled for Borneo, but it is unclear why Burma was added in parentheses.
2 d and 1 © (dried and pinned specimens transferred to alcohol) with label reading “Sumatra, leg. ?, Borneo, leg. Schwaner (?)”, lodged in the ZMA and re- corded by Loman (1902: 182), are very similar to the types of G. lomani sp. n. but cannot be assigned to this species with certainty. In one d the distal part of the penis is missing, in the other the penis is bent and its glans partly collapsed (caused by the pin which passed beside the tip of the penis; Figs 131, 132).
80 P. J. SCHWENDINGER & J. MARTENS
The weaker palpal femur, smaller ventral processes on palpal trochanter and femur (Fig. 140) and smaller dorsomedian boss on the proximal cheliceral article (Fig. 133) of the female (conspecific?) are probably due to sexual dimorphism. The same is seen in G. armillatus (see Schwendinger & Martens 1999b, figs 116-123) and - less distinctly so - also in G. exsudans sp. n. (Figs 181-184); it presumably also holds true for G. javanicus sp. n. (Figs 122-124).
Distribution: The type locality cannot be identified, but there are three localities which may correspond: 1) Pulau Talang (1°55’N, 109°46’E), an island off the coast at the western end of Sarawak, 2) Telang (2°07’S, 115°00’E) in southern Kalimantan and 3) Pulau Telang (0°43’N, 104°37’E), an island in the Riau Archipelago, south of Singapore.
Gnomulus obscurus sp. n. Figs 141-149
Material: MALAYSIA (east), Sarawak, Kuching, d holotype (SMF 7373/12, with label: “Pelitnus segnipes, Roewer det. 1939”).
Etymology: Latin: obscurus = hidden, unknown. The specific epithet refers to the previous misidentification of the type specimen.
Diagnosis: Similar to G. annulipes, distinguished by: Body smaller; eye tu- bercle lower, more rounded; penis more slender, with basally narrower, distally wider lateral sclerites and with a W-shaped median plate.
Description: 3 (holotype). Coloration (bleached): Body and cheliceral fingers light amber, no colour pattern on scuta discernible; limbs mostly light orange.
Carapace with widely conical, distally rounded eye tubercle and with a pair of small lateral tubercles below wide, indistinctly divided carapace-abdomen bridge. Anterior dorsal scutal areas little elevated, indistinctly keeled, posterior ones rounded (Fig. 145); a low longitudinal furrow separating anterior areas medially. Ventral scutal areas swollen, without modified hairs. Palpal coxa with large ventral process; leg coxa I with distinct anterolateral one; coxa II with strong, coxa III with small anteroproximal process, coxa II with posteroproximal one. Genital operculum almost semicircular (Fig. 146); posterior margin of stigmatic pit with small tubercle.
Chelicerae (Fig. 147) weak, proximal article with distinct dorsodistal to dorsomedian boss; no ventral process.
Palps (Fig. 148): Ventral side of femur with narrow, slightly anteriad-inclined proximal process and moderately developed dorsodistal to dorsoproximal boss; trochanter with slightly distad-inclined ventral process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus II about 3.5 times longer than wide (Fig. 149).
Penis (Figs 141-144) slender, widest at height of membraneous socket of glans; apex narrow, carrying plenty of setae; distal margin of truncus almost straight. Glans narrower than truncus at that point; lateral sclerites convex and with a moder- ately elevated dorsal ledge in basal half, in distal half laterally compressed, tapering and pointing away from the truncus; median plate short, with a narrow, protruding, somewhat W-shaped median portion; membraneous tubes completely covered by
FURTHER NEW SPECIES OF GNOMULUS 81
ASTA AES
DZ he
CS ‘N I
S
> za Sn i l VASI N
141
Fics 141-149
Gnomulus obscurus sp. n., 3 holotype. - Penis, dorsal (141) and lateral view (142); apex of penis, dorsal (143) and lateral view (144). Body, lateral view (145); genital operculum, ventral view (146); left chelicera, retrolateral view (147); left palp, retrolateral view (148); distal part of left leg II, retrolateral view (149). - Scale lines 0.1 mm (143, 144), 1.0 mm (others).
median plate; stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
2. Unknown.
Measurements: (4): Body 5.64 long, 3.99 wide; carapace region 1.33 long, 2.12 wide. - Palp and legs:
ihe Fe Pa Ti Mt Ta Total Palp 0.84 sini 0.89 0.62 - 1923 4.69 eect 0.64 249 13 1.36 2.12 113 8.78 Leg II 0.74 3.30 153 25] 3.30 1.58 12.96 Leg III 0.67 2.47 1225 1.63 2.56 1.18 9.74 Leg IV 0.94 3.30 1.48 2.34 SA] 155 13.16
Relationships: Penis morphology shows that G. obscurus Sp. n. is close to G. exsudans sp. n., G. javanicus sp. n., G. hutan sp. n., G. laevis, G. lomani sp. n. and G. sundaicus.
Distribution: Known only from Kuching in Sarawak, northern Borneo, where G. obscurus sp. n. apparently occurs sympatrically with G. sundaicus [Fig. 1 (24)].
Gnomulus hutan sp. n. Figs 150-163
Material: MALAYSIA (east), Sarawak, confluent of Sun Oyan and Mujong rivers, E of Kapit, 50 m, d holotype, d paratype, 18.V.1994, leg. I. Löbl & D. Burckhardt (MHNG). Etymology: Malay and Indonesian: hutan = forest; noun in apposition.
82 P. J. SCHWENDINGER & J. MARTENS
Diagnosis: Similar to G. lomani sp. n., distinguished by: Body larger; dorsal scutal areas less elevated; tubercles behind coxa IV present; ventral processes on palpal femur and trochanter smaller; penis stouter, with a narrower apex, a smaller membraneous base of the glans and a longer median plate.
Description 3 (holotype). Coloration mostly dark amber, with dark reticu- lation on carapace region, chelicerae and pedipalps; pattern on abdominal part of dorsal scutum indistinct; leg tarsi light amber, distitarsus I dorsally darkened.
Carapace with conical, terminally rounded eye tubercle; no lateral tubercles below wide, indistinctly divided carapace-abdomen bridge (Fig. 157). Dorsal scutal areas indistinctly elevated. Ventral scutum with a pair of anterolateral tubercles be- hind coxa IV; ventral scutal areas swollen, pallid, covered with very small short hairs (Fig. 156). Palpal coxa with pronounced ventral process; leg coxa I with short, wide anterolateral one; coxae II and III with distinct anteroproximal processes, coxa II with small posteroproximal one; dorsal side of coxa IV with distinct anterior tubercle (Fig. 156). Genital operculum rounded, as long as wide (Fig. 158); posterior margin of stigmatic pit with quite large tubercle.
Chelicerae (Fig. 159) weak, proximal article with distinct dorsodistal to dorsomedian boss; no ventral process.
Palps (Figs 160, 161): Femur with small ventroproximal process, with distinct dorsodistal to dorsoproximal boss and with wide, low prodorsal boss (Fig. 161); trochanter with digitiform, slightly distad-inclined ventral process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus II about 2.8 times longer than wide (Fig. 162).
Penis (Figs 150-155; holotype: 152, 153) fairly stout, widening in distal half, distal margin slightly invaginated; apex narrow, carrying plenty of setae. Glans slightly narrower than truncus at that point; lateral sclerites strongly convex, basal half with moderately elevated dorsal ledge, distal half narrowly paddle-shaped, pointing away from the truncus; median plate long, V-shaped, with a small pair of lateral teeth; membraneous tubes completely covered by median plate; stylus slender, base bulbous, apex with a small pair of ventral subterminal teeth.
2. Unknown.
Measurements: 3 holotype (d paratype in parentheses): Body 7.44 (7.47) long, 4.96 (5.08) wide; carapace region 1.49 (1.46) long, 2.70 (2.67) wide. - Palp and legs:
Ibe Fe Pa Ti Mt Ta Total Palp 0.96 1.30 0.99 0.81 - 155 5.61 leo 0.81 2.42 24 1.43 DI 1.24 9.34 Leg II 0.99 332 11:99 2.39 3.47 1.64 13.36 Leg III 0.87 2.45 1.30 1.61 2.79 1.12 10.14 Leg IV 1.24 3.41 1.67 2.42 4.43 1.30 14.47
Variation: The paratype has smaller ventral processes on its palpal femur and trochanter (Fig. 163) than the holotype (Fig. 160).
FURTHER NEW SPECIES OF GNOMULUS 83
a AB
7
CS [==
Gir WO YI ©
SÒ)
A 151 LEA cee
159
Fics 150-163
Gnomulus hutan sp. n., d holotype (152, 153, 156-162), 4 paratype (150, 151, 154, 155, 163). - Penis, dorsal (150) and lateral view (151); apex of penis, dorsal (152, 154) and lateral view (153); glans penis, lateral view (155). Body, lateral view (156); anterior body, dorsal view (157): genital operculum, ventral view (158); left chelicera, retrolateral view (159); left palp, retrolateral view (160); trochanter and femur of left palp, dorsal view (161); distal part of left leg II, retrolateral view (162); anterior body and proximal palp, lateral view (163). - Scale lines 0.1 mm (152-155), 1.0 mm (others).
Relationships: This species appears most closely related to G. lomani sp. n., followed by G. javanicus sp. n. and G. exsudans sp. n.
Distribution: Known only from the environs of Kapit, in the central part of Sarawak [Fig. 1 (25)].
Gnomulus exsudans sp. n. Figs 164-185
Material: MALAYSIA (east), Sarawak, Gunung Mulu National Park, Base Camp (= Paku Camp, 150 m?), d holotype, 1 d, 1 ® paratypes, 1 juv., VI.1978, leg. F. Wanless (NHML). - Sabah, Sandakan Bay (southwest), Sapagaya Lumber Camp, 2-20 m, 1 d paratype (BMH; penis dissected and retained by W. A. Shear), 7 2 paratypes, 2 juv., 4.-7.X1.1957, leg. J. L. Gressitt (BMH); Sandakan Bay (northwest), Sepilok Forest Reserve, 1-10 m, 7 6,5 @
84 P. J. SCHWENDINGER & J. MARTENS
paratypes, 1 juv., 30.X.1957, leg. J. L. Gressitt (1 dg, 1 ® paratypes in MAR, 1 6, 1 9 paratypes in MHNG, others in BMH).
Etymology: Latin: exsudans (present participle of exsudare) = sweating out. The specific epithet refers to the paired spots of secretion found on the dorsal and ventral scutum of this species.
Diagnosis: Similar to G. hutan sp. n., distinguished by: Body smaller; dorsal scutal areas more elevated; no tubercle on dorsal side of coxa IV; ventral process on palpal femur stronger; penis with wider apex; glans wider than truncus at that point, lateral sclerites distally wider and proximally more strongly elevated above the median plate.
Description: & (holotype). Coloration: Dorsal scutum amber, with dark reti- culation in carapace region, dark margin and dark scutal elevations (broken by light median longitudinal stripe in areas I-IV); ventral scutal elevations swollen and pallid (Figs 185a-c). Legs grey-brown, lighter near the joints; chelicerae, pedipalps and trochanters and tarsi of legs light brown, with dark reticulations on proximal article of chelicerae and on palpal trochanter and femur; tarsalia of leg I ventrally cream.
Carapace with pronounced, distally rounded eye tubercle slightly set back from front margin of scutum and with indistinct lateral tubercles posteriorly below wide, divided carapace-abdomen bridge. Dorsal scutal areas distinctly elevated and narrowly rounded (Fig. 185a, c); ventral scutal areas swollen, without modified hairs. Palpal coxa with pronounced ventral process; leg coxa I with indistinct anterolateral one; ventral side of leg coxae II and III with distinct anteroproximal processes, a small posteroproximal one on coxa H. Genital operculum about as long as wide; strongly pronounced tubercle on posterior margin of stigmatic pit (Fig. 185b).
Chelicerae (Fig. 177) weak, proximal article with dorsodistal to dorsomedian boss; no ventral process.
Palps (Fig. 179): Ventral side of femur with slightly distad-inclined proximal process, about as long as ventrad-directed process on ventral side of trochanter.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus of leg II 3.2 times longer than wide (Fig. 180).
Penis (Figs 164-176; holotype: 164, 165, 167, 168): Truncus slightly constric- ted at height of glans, distal margin broadly arched. Glans wider than truncus at that point; lateral sclerites convex, proximal half with dorsal ledge strongly elevated above median plate, distal half widely paddle-shaped, pointing away from the truncus; knee between proximal and distal half of lateral sclerites rounded, somewhat bulged towards the truncus; median plate V-shaped, with an indistinct pair of lateral teeth; membraneous tubes completely covered by median plate; stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
2. As the male but: Carapace region relatively smaller; ventral processes on palpal femur and trochanter slightly weaker (Figs 183, 184); proximal article of chelicera without dorsomedian boss (Fig. 178); ventral scutal areas not swollen; distitarsus II only 3.0 times longer than wide.
Measurements: 3 holotype (9 from type locality in parentheses): Body 5.95 (6.12) long, 4.18 (4.38) wide; carapace region 1.33 (1.16) long, 2.21 (2.12) wide. - Palp and legs:
FURTHER NEW SPECIES OF GNOMULUS 85
\ ER > = x
PERE BENZ
Fics 164-184
Gnomulus exsudans sp. n., 4 holotype (164, 165, 167, 168, 177, 179, 180), paratypes: d from the type locality (181), dd from Sabah (166, 169-176, 182), 2 from the type locality (178, 183), 2 from Sabah (184). - Penis, dorsal (164, 166) and lateral view (165); apex of penis, dorsal (167, 169) and lateral view (168); glans penis, dorsal (171, 173, 175) and lateral view (170, 172, 174, 176). Left chelicera, retrolateral view (177, 178); left palp, retrolateral view (179): distal part of left leg II, retrolateral view (180); anterior body and proximal palp, lateral view (181-184). - Scale lines 0.1 mm (167-176), 1.0 mm (others).
86 P. J. SCHWENDINGER & J. MARTENS
Fic. 185
Gnomulus exsudans sp. n., d holotype. - Body, dorsal (a), ventral (b) and lateral view (c). - Scale line 1.0 mm.
Tr Fe Pa Ti Mt Ta Total Palp 0.79 (0.79) 1.18 (1.18) 0.93 (0.89) 0.64 (0.59) -- 1.18 (1.23) 4.72 (4.68) Leg I 0.79 (0.64) 2.51 (2.31) 1.08(1.03) 1.40(1.28) 2.16(2.07) 1.16(1.13) 9.10 (8.46) Leg II 0.84(0.84) 3.64 (3.35) 1.57(1.50) 2.68 (2.56) 3.39 (3.25) 1.50(1.40) 13.62 (12.90) Leg III 0.69 (0.69) 2.66 (2.46) 1.28(1.21) 1.67(1.67) 2.68 (2.56) 1.38(1.35) 10.36 (9.94) Leg IV 1.03,0:93) 3.44(3.35) 1.53 (1.48) 2:41 0.39) 3.91 (3:76) 1.43.40) 13:75 (13:31)
FURTHER NEW SPECIES OF GNOMULUS 87
Variation: Range of measurements in dd (n= 10) [2 9 (n= 13) in paren- theses]: Body 5.26-5.95 (5.53-6.18) long, 3.68-4.45 (3.89-4.45) wide, carapace region 1.11-1.33 (1.02-1.17) long, 1.95-2.21 (1.88-2.12) wide. Specimens of the Sarawak and Sabah populations show the same penis morphology but differ in a few external characters. The animals from the type locality in eastern Sarawak have a more bulged anterior carapace margin, an eye tubercle slightly set back from the scutal front margin (Figs 181, 183, 185c) and a distincly less elevated anterolateral process on leg coxa I than specimens from Sabah. Eye tubercles vary from pointed to rounded (Figs 181-184, 185c). The tubercles behind coxa IV are more or less pronounced in different specimens and on either side of the body. The 4 paratype from Sarawak has distinct tubercles below its carapace-abdomen bridge (Fig. 181); these are indistinct or absent in other specimens (Figs 182-184, 185c). All specimens from Sabah are clearly darker in colour, which was probably caused by humid acids (from soil particles) in the preservative.
Remarks: Paired spots of denaturated secretion, arranged in a regular pattern near the lateral margins of the dorsal and ventral scutum, occur in all specimens exa- mined (Figs 181-185). These spots are very distinct in the specimens from Sarawak, less so (missing on ventral scutum) in those from Sabah. Such peculiar secretions have never been found in other oncopodid species and are probably specific. However, as these secretions are quite loosely attached to the cuticle and are easily brushed off, we don’t use them as a diagnostic character. They deserve further exa- mination.
Relationships: According to penis morphology, Gnomulus exsudans sp. n. is closest to G. hutan sp. n.; both are closely related to G. laevis, G. javanicus sp. n., G. lomani sp. n., G. obscurus sp. n. and G. sundaicus. Gnomulus conigerus, which also occurs in the Sepilok Forest Reserve, is clearly more distant and belongs to a different phyletic lineage within the armillatus-group (cf. Schwendinger, 1992: 183, 184, figs 27-41).
Distribution and bionomics: Gnomulus exsudans sp. n. is known from three localities, one in eastern Sarawak, the other two in eastern Sabah. The Sarawak [Fig. 1 (26)] and Sabah [Fig. 1 (27, 28)] populations are separated by about 400 km. According to a note on the label, the specimens from the type locality were swept from shrubs. In the Oncopodidae occurrence off the forest floor is most unusual and has otherwise only been observed in G. sundaicus from western Sarawak (Schwendinger 1992: 187).
Gnomulus carinatus sp. n. Figs 186-190 Pelitnus segnipes Loman. - Loman (1902: 182, partim). - Roewer (1923: 63, partim, fig. 66). - Schwendinger (1992: 187, figs 57-61).
Material: INDONESIA, South Kalimantan, Bandjermasin (= Banjarmasin), é holo- type, leg. Suck (SMF 1259).
Etymology: Latin: carinatus = keeled. The specific epithet refers to the keeled dorsal scutal elevations of this species.
Diagnosis: Externally similar to G. thorelli (male unknown), distinguished by keeled elevations on posterior part of dorsal scutum and by a longer subbasal process
88 P. J. SCHWENDINGER & J. MARTENS
on ventral side of palpal femur. Penis similar to that of G. armillatus but: Truncus with wider apex; glans shorter, wider; lateral sclerites more convex; median plate more rounded and carrying distinct lateral teeth.
Description: 8 (holotype). Coloration: Body reddish brown, colour pattern faded; legs yellow-brown, tarsi I, Il cream.
Carapace with conical, pointed eye tubercle and with a pair of small lateral tubercles posteriorly below wide, indistinctly divided carapace-abdomen bridge. Dorsal scutal elevations rounded in areas I-II, distinctly keeled in posterior areas; ventral scutal areas swollen, covered by few very small hairs without incrustations (Fig. 187). Palpal coxa with large ventral process; leg coxa I with distinct antero- lateral one; ventral side of leg coxae II and III with small anteroproximal processes, coxa II without posteroproximal one. Genital operculum somewhat triangular, slightly wider than long; posterior margin of stigmatic pit with tubercle.
Chelicerae (Fig. 188) weak, proximal article with distinct dorsodistal and indistinct dorsomedian boss; no ventral process.
Palps (Fig. 189): Ventral side of femur with strong subbasal, slightly distad- directed process; trochanter with short, ventrad-directed process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus of leg II about 2.5 times longer than wide (Fig. 190).
Penis (Fig. 186; Schwendinger, 1992: figs 58-61): Truncus in its distal half wider than in proximal half; distal margin broadly arched, median part almost straight. Glans slightly wider than truncus at that point; lateral sclerites strongly convex in proximal portion, with dorsal ledge moderately elevated above median plate, distal half cylindrical, weakly sigmoid, tapering and pointing away from the truncus; median plate short, widely rounded, with a distinct pair of lateral teeth; stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
9. Unknown.
Measurements: (8): Body 6.00 long, 3.98 wide; carapace region 1.23 long, 2.16 wide. - Palp and legs:
ihe Fe Pa u Mt Ta Total Palp 0.79 113 0.89 0.59 - I) 1558 Leg I 0.64 2.02 1.08 1.18 82 0.84 USS Leg II 0.74 2.66 1.38 1.82 2.61 1.28 10.49 Leg II 0.69 2.02 1.08 1.28 DA 0.84 8.12 eal) 0.79 2:50 1.38 1.87. 3:39 0.93 10.88
Relationships: Gnomulus carinatus sp. n. appears most closely related to G. thorelli ($ unknown). These are the only two Gnomulus species known at present, which possess a ventral process distinctly remote from the base of the palpal femur. Regarding penis morphology, most congruence is seen between G. carinatus sp. n. and G. armillatus.
Remark: Gnomulus carinatus sp. n. could be the conspecific male to the female syntypes of G. thorelli, since most differences between them correspond with sexual dimorphism found in other Gnomulus species. However, a wide geographic
FURTHER NEW SPECIES OF GNOMULUS 89
Fics 186-190
Gnomulus carinatus sp. n., 3 holotype. - Glans penis, dorsal view (186). Body, lateral view (187); left chelicera, retrolateral view (188); right palp, retrolateral view (189); distal part of left leg II, retrolateral view (190). - Scale lines 0.1 mm (186), 1.0 mm (others).
separation and the presence of keeled dorsal scutal elevations in G. carinatus sp. n. (rarely seen in Gnomulus) indicate distinctiveness.
Distribution: Known only from the type locality in southern Borneo [Fig. 1 (22)].
THE HAMATUS-GROUP (new)
Diagnosis: Medium-sized (5.1-5.3 mm) species with rounded eye tubercle and without carapace-abdomen bridge; posterior margin of stigmatic pit without tubercle; ventral side of palpal trochanter with slightly distad-directed, spatulate process. Penis subdistally widened; glans with an extensive membraneous socket; lateral sclerites distally truncate; median plate long, very narrow, turned upwards; stylus with a bul- bous base and with a pair of subterminal teeth.
Species account and distribution: Only a single species, G. hamatus sp. n., from Luzon, the Philippines.
Gnomulus hamatus sp. n. Figs 191-203
Material: PHILIPPINES, Luzon, Lagunas, Mt. Banahaw, above Kinabuhayan, trail to Crystalino, 600-700 m, d holotype, 24.XI.1995. - Mt. Makiling, south of Los Bafios, near Mad Springs, 400-450 m, 2 paratype, 19.X1.1995; leg. I. Löbl (both in MHNG).
Etymology: Latin: hamatus = furnished with a hook. The specific epithet refers to the upturned, hook-like median plate of the glans penis.
Diagnosis: Externally similar to G. minor (3 unknown), distinguished by: Body larger; interocular area elevated; dorsal and ventral scutal areas higher; ventral processes on palpal femur and trochanter longer. Distinguished from all other Gno- mulus species by a stout, subdistally very wide penis; its glans with a large membra- neous base and an upturned, hook-like median plate.
90 P. J. SCHWENDINGER & J. MARTENS
TR
Fics 191-203
Gnomulus hamatus sp. n., 3 holotype (191-197, 199-202), © paratype (198, 203). - Penis, dorsal (191) and lateral view (192); apex of penis, dorsal (193) and lateral view (194). Body, lateral (195) and dorsal view (196); anterior body, ventral view (197); left chelicera, retrolateral view (198, 199); left palp, retrolateral view (200); proximal part of right palp, retrolateral view (201); distal part of left leg II, retrolateral view (202); anterior body and proximal palp, lateral view (203). - Scale lines 0.1 mm (193, 194), 0.5 mm (191, 192), 1.0 mm (others).
Description: 3 (holotype). Coloration: Dorsal scutum amber, with dark reti- culation in carapace region, dark margin and dark, medially interconnected, trans- versal bands (Figs 195, 196); central portion of each ventral scutal elevation some- what pallid. Legs dark amber, except for light amber tarsalia (with darkened dorsal sides on tarsi I, III and IV).
Carapace with widely conical, distally rounded eye tubercle; no lateral tu- bercles in posterior portion; carapace-abdomen bridge absent. Dorsal scutal areas moderately elevated, the first one rising steeply behind the carapace region; ventral
FURTHER NEW SPECIES OF GNOMULUS 9]
scutal areas indistinctly elevated, densely covered with short white truncate hairs (Fig. 195). Palpal coxa with small ventral process; leg coxa I with distinct antero- lateral process; ventral side of leg coxa II with distinct anteroproximal and postero- proximal processes, the latter overlapping anteroproximal one on coxa II. Genital operculum wider than long; no tubercle on posterior margin of stigmatic pit (Fig. 197).
Chelicerae (Fig. 199) weak, proximal article with low, slightly forward- inclined dorsodistal to dorsomedian boss and indistinct retroventral tubercle.
Palps (Figs 200, 201): Femur with pronounced ventral process and dorso- proximal boss; ventral side of trochanter with long, spade-shaped, slightly distad- inclined process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus of leg II 2.1 times longer than wide (Fig. 202).
Penis (Figs 191-194): Truncus stout; apex with wide subterminal lateral lobes carrying plenty of setae; distal margin broadly arched. Glans narrower than truncus at that point; membraneous socket very large, shaped like a plate; lateral sclerites short, with broadly truncate tips close to each other; median plate narrow, hook-like, poin- ting away from the truncus and slightly distad; membraneous tubes mostly covered by basal part of median plate; stylus slender, base bulbous, apex with a small pair of subterminal ventral teeth.
2. As the male but palp with shorter ventral processes on femur and trochanter, proximodorsal boss on palpal femur less distinct (Fig. 203); ventral scutal areas darker and less elevated, covered with fewer hairs.
Measurements: 3 (2 in parentheses): Body 5.09 (5.29) long, 3.46 (3.66) wide; carapace region 1.28 (1.24) long, 1.95 (1.90) wide. - Palp and legs:
Tr Fe Pa Ti Mt Ta Total Palp 0.67 (0.59) 0.94 (0.89) 0.74 (0.69) 0.54 (0.49) -- 1.24 (1.24) 4.13 (3.90) LegI 0.52 (0.49) 1.48 (1.53) 0.79 (0.74) 0.86 (0.84) 1.38 (1.33) 0.84 (0.77) 5.87 (5.70) LesII 0.67 (0.69) 1.95 (1.98) 1.03(1.01) 1.33 (1.33) 2.07 (2.07) 0.99 (0.94) 8.04 (8.02)
Leg II 0.59 (0.54) 1.46 (1.46) 0.84 (0.79) 0.91 (0.89) 1.70 (1.68) 0.62 (0.62) 6.12 (5.98) LegIV 0.74 (0.74) 2.15 (2.10) 1.09 (1.09) 1.48 (1.46) 2.67 (2.52) 0.74. (0.74) 8.87 (8.65)
Relationships: In the shape of the stylus, in the slightly distad-inclined ventral process on palpal trochanter and in medium body size, G. hamatus sp. n. corresponds with species of the armillatus-group. The lack of a carapace-abdomen bridge and the presence of aberrant modifications of the penis, however, indicate that this species is closer to the goodnighti-group.
Distribution: Known from two mountains near San Pablo City in central Luzon [Fig. 1 (34, 35)].
THE TUMIDIFRONS-GROUP (new)
Diagnosis: Small (3.4-4.0 mm) species with relatively large eyes, without a carapace-abdomen bridge and with a distad-directed proximal process on ventral side of palpal trochanter; dorsal scutal elevations lighter in colour than areas in between them; posterior margin of stigmatic pit without tubercle. Truncus penis with circular
92 P. J. SCHWENDINGER & J. MARTENS
wrinkles in basal portion; glans with long, golfclub-shaped membraneous tubes protruding from under a narrow, pointed median plate; stylus strong, with a seemingly bulbous base and without subterminal ventral teeth.
Species account and distribution: This group is close to the goodnighti-group and it comprises two species, G. tumidifrons sp. n. and G. matabesar sp. n., from the Moluccas.
Gnomulus tumidifrons sp. n. Figs 204-216
Material: INDONESIA, Moluccas, Halmahera, Buli, Maba, 20 m, & holotype, 3 2 paratypes, 6.-7.X1.1999; leg. A. Riedel (1 2 paratype in MAR, others in MHNG).
Etymology: Latin: tumidus = swollen, frons = forehead; noun in apposition. The specific epithet refers to the characteristic frontal hump on the eye tubercle of this species.
Diagnosis: Similar to G. coniceps, distinguished by: Eyes larger; interocular tubercle smaller, with a distinct hump on frontal side; carapace-abdomen bridge absent; dorsal scutal areas light, less elevated; penis with narrower apex; lateral scle- rites of glans wider in distal portion, not covering long, golfclub-shaped membra- neous tubes.
Description: 3 (holotype). Coloration: Body light amber, with dark reti- culation in carapace region and dark pattern on dorsal and ventral scuta; dorsal scutal elevations light, separated by dark transversal bands and by a dark longitudinal median stripe; ventral scutal elevations I-V pallid (Fig. 216a-c). Chelicerae, pedipalps and leg coxae and trochanters light yellow-brown; other leg segments mostly darkened, except for light amber tarsi and distal portions of tibiae and metatarsi; distitarsus I ventrally cream.
Carapace with small conical eye tubercle bearing a hump on its front side; eyes large; no lateral tubercles on posterior carapace; carapace-abdomen bridge absent. Dorsal scutal areas moderately elevated, VI and VII most strongly so; ventral scutal areas indistinctly swollen (Fig. 216a, c) and covered with short hairs. Palpal coxa with strong ventral process; leg coxa I with indistinct anterolateral one; ventral side of leg coxae II and IH with small anteroproximal processes, indistinct posteroproximal one on coxa II. Genital operculum about as long as wide; no tubercle on posterior margin of stigmatic pit (Fig. 216b).
Chelicerae (Fig. 210) weak, proximal article with distinct dorsodistal and indistinct dorsomedian boss; no ventral process.
Palps (Fig. 211): Femur stout, ventral side with small proximal process; trochanter with distad-directed ventral process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus of leg II 1.6 times longer than wide (Fig. 212).
Penis (Figs 204-208): Truncus fairly stout, widest below glans, slightly constricted at height of glans, with circular wrinkles in proximal part; distal margin broadly arched. Glans slightly wider than truncus at that point; membraneous socket distally pointed; lateral sclerites massiv and convex, proximal portion wide, with a strong dorsal tooth on each side; tips of lateral sclerites fang-like, pointing towards each other; median plate long, narrowly triangular; membraneous tubes long, golf-
FURTHER NEW SPECIES OF GNOMULUS 93
pie)
Fics 204-215
Gnomulus tumidifrons sp. n., 3 holotype (204-208, 210-212), 9 paratypes (209, 213-215). - Penis, dorsal (204) and lateral view (205); apex of penis, dorsal (206) and lateral view (207); glans penis, ventral view (208). Left chelicera, retrolateral view (209, 210); left palp, retrolateral view (211): distal part of left leg II, retrolateral view (212); anterior body and proximal palp, lateral view (213-215). - Scale lines 0.1 mm (206, 207), 1.0 mm (others).
club-shaped, clearly visible in between median plate and lateral sclerites; stylus strong, fairly wide at bulbous base, tip pointed, without subterminal ventral teeth, completely covered by the median plate.
2. As the male, apart from very slightly less pallid ventral scutal elevations.
Measurements: 3 (2 in parentheses): Body 3.42 (3.34) long, 2.28 (2.25) wide; carapace region 0.90 (0.83) long, 1.20 (1.18) wide. - Palp and legs:
Tr Fe Pa Ti Mt Ta Total
Palp 0.45 (0.40) 0.49 (0.47) 0.41 (0.38) 0.28 (0.27) -- 0.57 (0.57) 2.20 (2.09) Leg I 0.35 (0.32) 0.82 (0.81) 0.47 (0.45) 0.47 (0.45) 0.69 (0.68) 0.46 (0.47) 3.26 (3.18) Leg II 0.41 (0.40) 1.04(1.04) 0.63 (0.60) 0.72 (0.69) 1.07 (1.07) 0.54 (0.56) 4.41 (4.36)
Leg III 0.35 (0.33) 0.72 (0.72) 0.50 (0.50) 0.49 (0.48) 0.87 (0.84) 0.39 (0.40) 3.32 (3.27) Leg IV. 0.47 (0.45) 1.01 (0.99) 0.66 (0.66) 0.79 (0.77) 1.28 (1.26) 0.44 (0.42) 4.65 (4.55)
Variation: Range of measurements in 9 9 (n= 3): Body 3.34-3.70 long, 2.25- 2.33 wide, carapace region 0.83-0.90 long, 1.18-1.23 wide.
Relationships: This species 1s most closely related to G. matabesar sp. n.
Distribution and bionomics: Known only from the type locality in the eastern region of Halmahera Island [Fig. 1 (39)]. The animals were collected by sifting leaf litter in a disturbed primary rain forest.
94 P. J. SCHWENDINGER & J. MARTENS
E
Fic. 216
Gnomulus tumidifrons sp. n., d holotype. - Body, dorsal (a), ventral (b) and lateral view (c). - Scale line 1.0 mm.
Gnomulus matabesar sp. n. Figs 217-224
Material: INDONESIA, Moluccas, Halmahera, mountains SW of Tobelo, 850 m, d holotype, 2 juv., 1.X1.1999; leg. A. Riedel (MHNG).
Etymology: Malay and Indonesian: mata = eye, besar = big; noun in apposition. The specific epithet refers to the unusually large eyes of this species.
Diagnosis: Closely related to G. tumidifrons sp. n., distinguished by: Eye tu-
bercle without frontal hump; colour pattern on dorsal scutum less marked; antero-
FURTHER NEW SPECIES OF GNOMULUS 95
proximal processes on ventral side of leg coxae II larger; distitarsus II relatively longer; truncus penis more slender; membraneous socket of glans with broadly rounded distal margin; lateral sclerites basally narrower, without dorsal teeth and with truncate tips; median plate shorter, wider at base, with lateral teeth.
Description: 3 (holotype). Coloration as in G. tumidifrons sp. n., but dark markings generally less pronounced (bleached?).
Carapace with stout, conical, slightly foreward-inclined eye tubercle; eyes large; no lateral tubercles in posterior part; carapace-abdomen bridge absent. Dorsal scutal areas moderately elevated, VI and VII highest (Fig. 221); ventral scutal areas covered with short hairs. Palpal coxa with pronounced ventral process; leg coxa I with indistinct anterolateral one; ventral side of leg coxa II with quite large, III with small and narrow anteroproximal processes, a short, rounded posteroproximal one on coxa II. Genital operculum about as long as wide; no tubercle on posterior margin of stigmatic pit.
Chelicerae (Fig. 222) weak, proximal article with dorsodistal boss; no ventral process.
Palps (Fig. 223): Femur stout, with small ventroproximal process; trochanter with distad-directed ventral process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus of leg H 2 times longer than wide (Fig. 224).
Penis (Figs 217-220): Truncus constricted at height of glans, with circular wrinkles in proximal half; apex with broadly arched distal margin. Glans about as wide as truncus at that point, its membraneous socket widely rounded distally; lateral sclerites short, their broadly rounded tips covering lateral parts of long, golfclub- shaped membraneous tubes; median plate short, its distal part narrowly triangular, its base wide, with distinct lateral teeth; stylus fairly strong, base seemingly bulbous, tip pointed, without subterminal teeth, completely covered by the median plate.
2. Unknown.
Measurements: d : Body 3.59 long, 2.35 wide; carapace region 0.93 long, 1.29 wide. - Palp and legs:
ibe Fe Pa Ti Mt Ta Total Palp 055 0.58 0.46 0.32 - 0.64 2.35 Leg I 0.37 0.97 0.55 0.55 0.86 0.33 3.85 Leg II 0.47 124 0.72 0.89 1.39 0.67 5158 Leg III 0.42 0.84 0.56 0.58 1.08 0.47 3.95 Leg IV 0.52 1.19 072 0.95 JESS 052 5.45
Remarks: The juveniles examined possess a high, conical, but not forward- inclined eye tubercle.
Relationships: Gnomulus matabesar sp. n. and G. tumidifrons sp. n. are closest relatives. Small body size, a distad-directed ventral process on palpal trochanter and modifications of membraneous tubes and stylus indicate phylogenetic proximity to the goodnighti-group from the Philippines. The shape of the membraneous tubes also points to a fairly close relationship with G. latoperculum sp. n. from Sulawesi.
96 P. J. SCHWENDINGER & J. MARTENS
220
BU) DIO =
D
>)
=" = Wi. © N hs
yw
TT MP.
Fics 217-224
Gnomulus matabesar sp. n., & holotype. - Penis, dorsal (217) and lateral view (218); apex of penis, dorsal (219) and lateral view (220). Body, lateral view (221); left chelicera, retrolateral view (222); left palp, retrolateral view (223); distal part of left leg H, retrolateral view (224). - Scale lines 0.1 mm (219, 220), 1.0 mm (others).
Distribution: Known only from the type locality in the northern part of Halmahera Island [Fig. I (38)].
THE LATOPERCULUM-GROUP (new)
Diagnosis: Large (5.7-6.7 mm) species with a wide, undivided carapace- abdomen bridge and with a slightly distad-inclined process on ventral side of palpal trochanter; posterior margin of stigmatic pit with tubercle; genital operculum very wide; penis scoop-shaped, carrying an enlarged pointed stylus with an invaginated base and without subterminal ventral teeth.
Species account and distribution: At present, this group is represented only by G. latoperculum sp. n. from northern Sulawesi.
Gnomulus latoperculum sp. n. Figs 225-240
Material: INDONESIA, Sulawesi, Northern Sulawesi Province, Dumoga - Bone National Park, 9 holotype, ¢ paratype (penis not examined), 2 juv., XI.1985, leg. P. D. Hillyard (Project Wallace Expedition: NHML). - Same province, Kotamobagu, Matalibaru, Gunung Tongara, 800-900 m, 2 2 paratypes, 5.-9.XII.1999, leg. A. Riedel (MHNG).
Etymology: Latin: latus = wide, extensive, operculum = cover, lid; noun in apposition. The specific epithet refers to the unusually wide genital operculum (especially in males) of this species.
Diagnosis: Similar to G. claviger sp. n., distinguished by: Body much larger; eye tubercle lower; carapace-abdomen bridge wider, undivided; genital operculum
FURTHER NEW SPECIES OF GNOMULUS 97
Fics 225-239
Gnomulus latoperculum sp. n., $ holotype (225-230, 237), & paratype (234-236), 2 paratypes (231-233, 238, 239). - Penis, dorsal (225), lateral (226) and ventral view (227); apex of penis, dorsal (228), lateral (229) and distal view (230). Anterior body, ventral view (231); genital operculum, ventral view (232). Left chelicera, retrolateral view (233, 234); left palp, retrolateral view (235); distal part of left leg II, retrolateral view (236); anterior body and proximal palp, lateral view (237-239). - Scale lines 0.1 mm (228, 229), 1.0 mm (others).
wider; truncus penis scoop-shaped; glans carrying lobate lateral sclerites and a basally thick, distally tapering stylus; median plate absent.
Description: è (paratype). Coloration quite pale, light yellow-brown (newly moulted specimen). Dark reticulation in carapace region and dark pattern on dorsal and ventral scuta; dark transversal bands on dorsal scutal elevations broken by light, narrow, longitudinal stripe in areas I-VI; dorsal and ventral scutal elevations each with a pallid transversal band in its centre (Fig. 240a-c). Chelicerae and pedipalps
98 P. J. SCHWENDINGER & J. MARTENS
Fic. 240
Gnomulus latoperculum sp. n., d paratype. - Body, dorsal (a), ventral (b) and lateral view (c). - Scale line 1.0 mm.
with dark reticulation. Legs mostly dark, except for light distal portion on tibiae and light median zone on metatarsi of posterior legs and light tarsi on all legs.
Carapace with small, low, rounded eye tubercle; no lateral tubercles below wide, undivided carapace-abdomen bridge. Dorsal scutal areas moderately elevated;
FURTHER NEW SPECIES OF GNOMULUS 99
ventral scutal areas slightly swollen (Fig. 240a, c), covered with short hairs. Palpal coxa with pronounced ventral process; leg coxa I with small anterolateral one; ventral side of leg coxae II and HI with small conical anteroproximal processes, an indistinct rounded posteroproximal one on coxa II. Genital operculum very large, much wider than long; posterior margin of stigmatic pit with pronounced tubercle (Fig. 240b).
Chelicerae (Fig. 234) weak, proximal article with dorsodistal boss; no ventral process.
Palps (Fig. 235): Femur and trochanter each with a slightly distad-inclined ventral process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus of leg II 2.3 times longer than wide (Fig. 236).
Penis (Figs 225-230): Truncus stout, apex very wide, scoop-shaped (see Fig. 230 for distal view), with almost straight distal margin; setae restricted to two pallid, cushion-shaped oval areas in a lateral subterminal position. Glans large, about as wide as truncus at that point, situated close to anterior margin of truncus; membraneous socket large, distally widely rounded; lateral sclerites lobate, pointing down the truncus; median plate absent; membraneous tubes long, distally flat, wide and roun- ded, clearly visible in between lateral sclerites and stylus; stylus strongly enlarged, very wide at invaginated base, distally tapering, without ventral subterminal teeth.
2. As the male, apart from less elevated, entirely dark ventral scutal elevations and a more or less distinctly narrower genital operculum (Figs 231, 232).
Measurements: 3 holotype (? in parentheses): Body 6.21 (6.36) long, 4.12 (4.09) wide; carapace region 1.38 (1.23) long, 2.44 (2.29) wide. - Palp and legs:
(SHIT
Tr Fe Pa Ti Mt Ta Total Palp 0.89 (0.71) 1.26(1.13) 0.99 (0.84) 0.67 (0.59 1.33 (1.31) 5.14 (4.58)
) > LegI 0.67 (0.62) 2.02(1.92) 1.04(0.94) 1.13(1.11) 1.75(1.73) 0.99 (0.91) 7.60 (7.23) Leg IT 0.79 (0.74) 2.61 (2.54) 1.33 (1.23) 1.70(1.75) 2.56(2.47) 1.16(1.11) 10.15 (9.84) Leg II 0.69 (0.59) 1.87(1.85) 1.06 (0.99) 1.26 (1.23) 2.12 (2.17) 0.79 (0.79) 7.79 (7.62) Leg IV 0.94 (0.84) 2.51 (2.54) 1.33(1.26) 1.85 (1.87) 3.20 (3.20) 0.89 (0.89) 10.72 (10.60)
Variation: Range of measurements in dd (n= 2) and 99 (n=2; in parentheses): Body 5.74-6.21 (6.36-6.68) long, 3.99-4.12 (4.09-4.51) wide, carapace region 1.26-1.38 (1.23-1.48) long, 2.29-2.44 (2.29-2.56) wide. The d paratype has distinct pale transversal bands in the central zones of its dark dorsal scutal elevations (Fig. 240a). This is not observable in the holotype, where the underlying pigmentation is partly detached from the cuticle (due to preservation?). One @ has a distinctly higher eye tubercle and smaller ventral processes on palpal femur and trochanter (Fig. 239). Variation in the size of the genital operculum, see Figs 231, 232, 240b.
Relationships: The relationships of G. latoperculum sp. n. are unclear. Judging from penis morphology it appears most closely related to the goodnighti-species group.
Distribution: Known only from two localities (close to each other) in northern Sulawesi [Fig. 1 (41)]. The 2 paratypes were collected by sifting leaf litter in a selectively logged primary rain forest.
100 P. J. SCHWENDINGER & J. MARTENS
THE GOODNIGHTI-GROUP (see Schwendinger & Martens, 1999b: 974)
Diagnosis: Small (1.9-3.8 mm) species, additionally characterized by: Ventral process on palpal trochanter distinctly distad-directed; posterior margin of stigmatic pit without tubercle; carapace-abdomen bridge absent (G. crucifer, G. minor, G. crassipes sp. n.) or present (all others); glans penis usually with quite long mem- braneous tubes; stylus enlarged (often strongly modified), if slender, then without ventral pair of subterminal teeth (G. coniceps, G. imadatei?); base of stylus bulbous (G. coniceps, G. goodnighti and G. crassipes sp. n.) or invaginated (all others).
Species account and distribution: Nine species: One from Brunei [G. imadatei (Suzuki)] and eight from the Philippines [G. claviger sp. n., G. coniceps Martens & Schwendinger, G. crassipes sp. n., G. crucifer Martens & Schwendinger, G. good- nighti (Suzuki), G. leyteensis Martens & Schwendinger, G. maculatus Martens & Schwendinger, G. minor Tsurusaki (male unknown - assignment uncertain)].
Gnomulus claviger sp. n. Figs 241-255
Material: PHILIPPINES, Luzon, Mt. Banahaw, above Kinabuhayan, trail to Cristalino, 600-700 m, d holotype, 24.X1.1995, leg. I. Löbl (MHNG). - Los Baños, Mt. Makiling (= Mt. Maquiling), 2 4 paratypes, X1.1968 and 5.V.1968, leg. R. A. Morse (AMNH).
Etymology: Latin: clava = club, cudgel; ger (suffix derived from gerere) = furnished with. The specific epithet refers to the club-shaped penis of this species.
Diagnosis: Close to G. maculatus, distinguished by: Colour pattern different; ventroproximal process on palpal femur larger; penis stouter, distally wider, with membraneous socket extending beyond apex of truncus; sclerites of glans, especially stylus, different in shape.
Description: 3 (paratype). Coloration: Body amber, with dark reticulation in carapace region and dark margin and transversal bands on dorsal scutum (cf. Fig. 255a, c); dark transversal bands on ventral scutum less distinct (median portion most clearly pronounced). Genital operculum light throughout (cf. Fig. 255b). Chelicerae and pedipalps with dark reticulation, except for light yellow-brown hand and tarsus, respectively. Legs mostly darkened, except for light tarsi and light distal portion on tibiae and metatarsi.
Carapace with low, widely conical, terminally rounded eye tubercle bearing a small hump on its front side; no lateral tubercles in posterior part; carapace-abdomen bridge formed by two opposing pairs of tubercles. Dorsal scutal areas only indistinctly elevated; ventral scutal areas with few fine hairs, not recognizably modified (cf. Fig. 255a, c). Palpal coxa with strong ventral process; leg coxa I with indistinct antero- lateral one; ventral side of leg coxae II and III with small conical anteroproximal processes, a knob-shaped posteroproximal one on coxa II. Genital operculum clearly wider than long; posterior margin of stigmatic pit without tubercle (Fig. 255b).
Chelicerae (Fig. 249) weak, proximal article with distinct dorsodistal and indistinct dorsomedian boss; no ventral process.
Palps (Fig. 250): Femur short, ventral side with small, ventrad-directed pro- ximal process; trochanter with distad-directed ventral process.
Legs 1324, tarsal formula 2-2-3-3. Distitarsus II 2.1 times longer than wide
FURTHER NEW SPECIES OF GNOMULUS 101
Fics 241-254
Gnomulus claviger sp. n., 3 holotype (245, 246, 254), d paratypes (141-244, 247-253). - Penis, dorsal (241) and lateral view (242); apex of penis, dorsal (243, 245, 247) and lateral view (244, 246, 248). Left chelicera, retrolateral view (249); left palp, retrolateral view (250); distal part of left leg II, retrolateral view (251); distal part of left leg I, dorsal view (252). Anterior body and proximal palp, lateral view (253, 254). - Scale lines 0.1 mm (243-248), 0.5 mm (others).
(Fig. 251); distitarsus I somewhat egg-shaped, clearly wider than preceding leg seg- ments (Fig. 252).
Penis (Figs 241-248; holotype: 245, 246): Truncus stout, strongly widening in distal half; apex very wide, with broadly arched distal margin. Glans very close to tip of truncus, slightly narrower than truncus at that point; membraneous socket distally
P. J. SCHWENDINGER & J. MARTENS
102
Fic. 255 Gnomulus claviger Sp. n., 3 paratype. - Body, dorsal (a), ventral (b) and lateral view (c). -
Scale line 1.0 mm.
FURTHER NEW SPECIES OF GNOMULUS 103
inflatable; lateral sclerites each with a lateral spur at about mid-length, tips tapering, inclined towards each other, pointing down the truncus; median plate long, narrowing towards the truncate tip; membraneous tubes short, mostly covered by median plate; stylus distinctly enlarged, base invaginated, tip broadly truncate, with distolateral edges drawn into recurved hooks.
2. Unknown.
Measurements: 3 paratype: Body 2.45 long, 1.56 wide; carapace region 0.62 long, 0.94 wide. - Palp and legs:
tor: Fe Pa Ti Mt Ta Total Palp 0.31 0.40 0.33 0.20 - 0.50 1.74 Eee I 027 0.68 0.40 0.40 0.57 0.42 2.74 Leg II 0.35 0.93 Onl 0.60 0.88 0.51 3.78 Leg III 0.29 0.64 0.41 0.40 0.73 0.33 2.80 Bes IV 0.35 0.93 0.56 0.68 1.01 0.37 3.90
Variation: Measurements, external and genital characters of all three d differ only to a minor extent: Body 2.43-2.56 long, 1.56-1.72 wide, carapace region 0.62- 0.64 long, 0.93-0.98 wide; eye tubercles, see Figs 253, 254, 255c.
Remarks: In two males the membraneous socket of the glans penis is distally extended into a flat cap (Figs 245-248); in the third male the socket appears to be deflated and its distal portion retracted (Figs 241-244). An inflatable socket has not been observed in other oncopodids and it is not clear whether it has any function during copulation and whether inflation also occurs on the expanded penis (when the glans is folded upwards).
Relationships: The strongly modified stylus of G. claviger sp. n. clearly places this species in the goodnighti-group, closest to G. maculatus. Strong resemblance in the shape of the truncus penis between G. claviger sp. n. and G. latoperculum sp. n. 1s considered to be due to convergence.
Distribution: Known only from two mountains on Luzon Island [Fig. 1 (34, 35)], where this species occurs together with G. hamatus sp. n. On one of these mountains, Mt. Makiling, a third congeneric species, G. minor (generic placement has yet to be confirmed by males), was found.
Gnomulus crassipes sp. n. Figs 256-273
Material: PHILIPPINES, Luzon, Mt. Banahaw, near school, 500 m, about 1 km from Kinabuhayan, 3 holotype, 2 6, 1 @ paratypes, 1 juv., 26.X1.1995; summit trail, 800 m, 1 d paratype, 25.X1.1995; above Kinabuhayan, trail to Cristalino, 600-700 m, 3 ® paratypes, 24.X1.1995. All specimens leg. I. Löbl (1 3, 1 9 paratypes in MAR, others in MHNG).
Etymology: Latin: crassus = thick, stout, pes = leg; noun in apposition. The specific epithet refers to the incrassate metatarsus III of males in this species.
Diagnosis: Close to G. crucifer (also possessing the unusual tarsal formula 2-2-2-2), distinguished by: Body smaller; colour pattern different; ventroproximal process on palpal femur larger; truncus penis subdistally constricted; glans with longer lateral sclerites; stylus different in shape.
104 P. J. SCHWENDINGER & J. MARTENS
Description: 3 (holotype). Coloration: Body light brown, with dark reticu- lation on carapace region, chelicerae and pedipalps; abdominal region of dorsal scutum amber, with dark margin and dark transversal bands (laterally more or less distinctly touching each other) on all areas (Fig. 273a, c); transversal bands on ventral scutum weakly pronounced (Fig. 273). Femora to metatarsi of legs darkened, tarsi light brown.
Carapace with very low, widely rounded eye tubercle; posterior portion of carapace elevated, slightly higher than eye tubercle; no lateral tubercles in posterior part; carapace-abdomen bridge absent. Dorsal scutal areas only indistinctly elevated, ventral scutal areas slightly more so, without hairs (Fig. 273a, c). Palpal coxa with pronounced ventral process; leg coxa I with indistinct anterolateral one; ventral side of leg coxae II and HI with conical anteroproximal processes, a knob-shaped postero- proximal one on coxa II. Genital operculum wider than long; no tubercle on posterior margin of stigmatic pit (Fig. 273b).
Chelicerae (Fig. 266) weak, proximal article with slightly forward-inclined dorsodistal and indistinct dorsomedian boss; no ventral process.
Palps (Fig. 267): Femur short, with widely rounded ventroproximal process; trochanter with distad-directed ventral process.
Legs 3142, tarsal formula 2-2-2-2. Distitarsus II 1.2 times longer than wide (Fig. 268); metatarsus HI recognizably inflated (Figs 269, 270).
Penis (Figs 256-263; holotype: 260, 261): Truncus slender, strongly constric- ted below axe-shaped (in dorsal view) apex; distal margin almost straight; a pair of lateral setae situated above constriction, few other setae below it. Glans slightly wider than truncus at that point; membraneous socket distally wide, rounded; lateral scle- rites long, pointing down the truncus, their tips unevenly rounded; median plate short, pointed, mostly covered by lateral sclerites; membraneous tubes long, covered by lateral sclerites; stylus enlarged, base bulbous, distal portion compressed at the sides, with a dorsal boss at some distance from the blade-shaped tip, sperm duct opening on the tip of a narrow medioventral spine pointing towards the truncus.
9. As the male but metatarsus II not incrassate (Figs 271, 272).
Measurements: 3 holotype ( in parentheses): Body 1.94 (2.09) long, 1.17 (1.26) wide; carapace region 0.56 (0.60) long, 0.70 (0.72) wide. - Palp and legs:
Tr Fe Pa Ti Mt Ta Total Palp 0.20 (0.21) 0.25 (0.25) 0.24(0.24) 0.16 (0.16) = - 0.32. (033) 2,7719) Leg I 0.21 (0.21) 0.42 (0.44) 0.29(0.30) 0.21 (0.22) 0.25(0.36) 0.22(0.23) 1.70(1.76) Leg II 0.24 (0.27) 0.54(0.57) 0.36(0.38) 0.31 (0.32) 0.54 (0.55) 0.27 (0.28) 2.26 (2.37) ( (
Leg IMI 0.21(0.21) 0.33 (0.35) 0.28(0.29) 0.23(0.23) 0.41(0.41) 0.15(0.15) 1.61 (1.64) Leg IV 0.25 (0.27) 0.50(0.52) 0.37(0.39) 0.37 (0.39) 0.56(0.58) 0.17 (0:17) 2.22 2.31)
Variation: Range of measurements in dd (n = 4) and 2 9 (n= 4; in paren- theses): Body 1.90-1.98 (1.99-2.09) long, 1.13-1.20 (1.22-1.26) wide, carapace region 0.55-0.56 (0.56-0.60) long, 0.68-0.70 (0.72) wide. In some specimens the dark transversal bands on the dorsal scutum are broken by light, narrow longitudinal stripes in areas II and III: the ventral scutal bands are partly broken in some specimens. In one male the median plate of the glans penis