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Full text of "Revue suisse de zoologie"

aininALES 



de la 

SOCIÉTÉ SUISSE DE ZOOLOGIE 

et du 

MUSÉUM D'HISTOIRE NATURELLE 

de la Ville de Genève 



tome 1 09 
fascicule 1 
2002 



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GENEVE MARS 2002 ISSN 0035 - 418 X 




REVUE SUISSE DE ZOOLOGIE 



TOME 109 — FASCICULE 1 

Publication subventionnée par: 

Académie suisse des Sciences naturelles ASSN 

. Ville de Genève 

Société suisse de Zoologie 



VOLKER MAHNERT 

Directeur du Muséum d'histoire naturelle de Genève 

CHARLES LIENHARD 

Chargé de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

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 à des experts d'institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie 
comparée, physiologie. 

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 

Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 



Prix de l'abonnement: 

SUISSE Fr. 225.— UNION POSTALE Fr. 230.- 

(en francs suisses) 

Les demandes d'abonnement doivent être adressées 

à la rédaction de la Revue suisse de Zoologie, 

Muséum d'histoire naturelle, C.P. 6434, CH-121 1 Genève 6, Suisse 



ANNALES 

de la 

SOCIÉTÉ SUISSE DE ZOOLOGIE 

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REVUE SUISSE DE ZOOLOGIE 



TOME 109 — FASCICULE 1 

Publication subventionnée par: 

Académie suisse des Sciences naturelles ASSN 

Ville de Genève 

Société suisse de Zoologie 



VOLKER MAHNERT 
Directeur du Muséum d'histoire naturelle de Genève 

CHARLES LIENHARD 

Chargé de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

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 à des experts d'institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie 
comparée, physiologie. 

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 

Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 



Prix de l'abonnement: 

SUISSE Fr. 225.— UNION POSTALE Fr. 230, 

(en francs suisses) 

Les demandes d'abonnement doivent être adressées 

à la rédaction de la Revue suisse de Zoologie, 

Muséum d'histoire naturelle, C.P. 6434, CH-121 1 Genève 6, Suisse 



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 "Muséum 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 6: 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. 



SMonicixonis schwenmngeri sp. n. 




-*do 



~+ pe 



Figs 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 histoty: 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. 



SM0DIC1N0DES SCHWENDINGERI SP. N. 7 

The differentiation of the "sister" genera Smodicinus Simon, 1895, Paras- 
modi.x 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. 

Jézéquel, J. F. 1966. Araignées de la savane de Singrobo (Côte d'Ivoire). V. Note complé- 
mentaire sur les Thomisidae. Bulletin du Muséum National d'Histoire Naturelle, Paris 
37:613-630. 



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 revisionai 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 1 & Peter VOGEL 

Institute of Ecology, University of Lausanne, BB, IE-ZEA, 

CH-1015 Lausanne-Dorigny, Switzerland. 

E-mail: brigitte.reutter@ie-zea.unil.ch 
1 Ethology - Evolution - Ecology, University of Rennes I - CNRS (UMR 6552), 

Campus de Beaulieu, Bât. 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. flavieollis 
(Melchior, 1834). A. alpicola was originally considered as a high-altitude subspecies 
of A. flavieollis (Heinrich, 1951, 1952) and later described as a new species by Storch 
and Lütt (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 morphometries 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 1 1.09.2001 



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 et al, 2001). However, it requires to use fresh or frozen 
blood samples. 

Although morphologic (Storch & Lütt, 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 
(Tegelström & 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 ALP1COLA 



11 



France 




Fig. 1 
Geographic distribution of the sampled localities. 



Table 1 
A survey of material examined (sampling localities and number of individuals). All specimens 
were previously identified using morphometric (m) and/or allozymic (a) methods. 



Locality (map symbol) 


A. sylvaticus 


A. flavicollìs 


A. alpi cola 


Switzerland 










Gordevio TI (S 1 ) 




4 (a) 






Prosito TI (S2) 


3 (a) 








Martigny VS (S3) 




4(a) 






Sanetsch VS (S4) 


1 (a) 


1 (a) 


6 (a) 




Bourg St. Bernard VS (S5) 


1 (m) 


2 < m ' 


2 ( m ) 




ChurGR(S6) 


10 (m) 


10 (m) 






Haslital BE (S7) 


3 (a) 


1 (a) 


2 (m) 


Germany 










Garmisch (G) 




6 (a) 




Austria 












Silbertal (Al) 


1 (a,m) 


2 (a.m) 


9 (m) 




Hohe Tauern (A2) 




5 (m) 


7 (m) 


Italy 












Valle d'Aosta (II) 


7 (a,m) 


1 (a) 


3 (a.m) 




Gran Paradiso NP (12) 


7 (m) 


7 (m) 


2 (m ) 




Vinschgau (13) 


g(m) 


1 (m) 


5 < m > 


France 


Morzine (FI) 




1 (m) 






Mt. Cenis (F2) 


5 Cm) 




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 et 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 pi 
reaction mixture contained the two primers (1 pM each), 2.5 mM MgCL, 0.25 mM 
each dNTP, 2.5 pi reaction buffer, 5 pi 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, i.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 pi volume 
consisting of 0.5 pM primer, 4 pi Dye mix, and 5.5 pi 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 Spel according to the following protocol: 
10 pi of the amplified DNA were digested in a 25 pi reaction mixture comprising 2.5 
units Spel (A//CTAGT) (Life Technologies) and 2.5 pi 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 \ 3 

found to meet these criteria. Spel cleaved the amplified part of the cytochrome b gene 
at the position 723 into two fragments of 3 1 1 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-ge\ 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. 



S ■§ .8 ** 



* * W u 



& ^ « 




s P eI W4— 500 bp 



Fie. 2 



Representative examples of fragment patterns after Spel endonuclease digestion of an 866 bp 
fragment of the cytochrome b gene. Apodemus alpicola showed two fragments after Spei 
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 syfnpatry 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 et 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 Jtirg Paul Müller from the Bündner 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 ALP1COLA \ 5 



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84-89. 
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15 B. A. REUTTER, E. PETIT & P. VOGEL 



Vogel, P., Maddalena, T., Mabille, A. & Paquet, G. 1991. Confirmation biochimique du 
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Revue suisse de Zoologie 109 (1): 17-22; mars 2002 



Proportionsänderungen beim Mj im Gebiss des mitteleuropäischen 
Dachses Mêles mêles (Mammalia, Carnivora) 

Beatrice BLÖCHLINGER & Peter LÜPS 

Naturhistorisches Museum der Burgergemeinde Bern, Bernastrasse 15, 

CH-3005 Bern, Schweiz. 

Changing proportions in Mj in the European badger, Mêles mêles 
(Mammalia, Carnivora). - We measured M, trigonid- and talonid-length 
in 62 recent and 110 neolithic badger Mêles 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 Mi 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 Degerb0l (1933) bei 
Dachsen aus Dänemark zeigen konnte. 

Analog zu Degerb0l bei den dänischen Dachsen, fanden Grundbacher et 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 Mj auf. Da der P 4 mit dem Mj- 
Trigonid ein gut harmonierendes Kauwerkzeug bildet, stellt sich die Frage nach 
allfälligen Veränderungen der prozentualen Anteile von Trigonid und Talonid unter 
der Voraussetzung des gleich lang gebliebenen Zahnes. 



Manuskript angenommen 28.08.2001 



lg B. BLÖCHLINGER & P. LÜPS 

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: 

1. Intakte Kiefer rezenter, als Unfallopfer und Hegeabschüsse in den Jahren 1967-1982 

gesammelter Dachse (Sammlung Naturhist. Museum Bern, Lüps, 1984): 

- je 12 S und 9 im Alter von 6-12 Monaten, d.h. Tiere mit völlig entwickeltem, aber 
noch wenig abgenutztem Gebiss (Lüps & Roper, 1988a) 

- 21 6 und 17 9, mindestens 13 Monate alt (vgl. Lüps et al., 1987, Grundbacher et ai, 
1990) 

2. Unterkiefer aus zwei neolithischen Stationen: 

- Seeberg Burgäschisee-Süd (ca. 3750 - 3700 v. Chr., Schibier & Suter, 1990): 16 
Unterkieferäste (s. Jéquier, 1963) 

- Twann (ca. 3840 - 3530 v. Chr., Suter & Schifferdecker, 1986): 94 Unterkieferäste 
(vgl. Grundbacher et 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. 



M j - PROPORTIONSANDERUNGEN BEI MELES MELES 



19 




^fc 



Abb. 1 
Linke Vorbacken- und Backenzahnreihe (Ansicht buccal) eines rezenten männlichen Dachses, 
mit eingezeichneten Messstrecken am Mj. 



RESULTATE 

Weder bei den Zähnen der adulten, noch bei denjenigen der 6 bis 12 Monate 
alten rezenten Dachse Hess 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 t (Gesamtlänge, grösste 
Breite und Höhe: Lüps & Roper, 1988a; Gesamtlänge und Breite: Grundbacher et ah, 



20 



B. BLÖCHLINGER & P. LÜPS 



Tab. 1 

Messwerte am Mj für neolithische und rezente Dachse in mm. 

(Student T-Test für unabhängige Stichproben) 







Dachse 


Dachse 


P 


Zunahme/Abnahme 






neolithisch 


rezent 




% 


Gesamtlänge 


n 


94 


62 


0.136 


- 




X 


16.45 


16.27 








s 


0.70 


0.78 






Trigonid Länge 


n 


92 


62 


0.025 


+2.7 




X 


8.26 


8.49 








s 


0.69 


0.45 






Trigonid Breite 


n 


93 


62 


0.165 


- 




X 


5.23 


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 








s 


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: 
Degerböl, 1930, für das Schweizerische Mittelland: Clutton-Brock, 1990, Grund- 
bacher et al., 1990). Für den Mj Hess 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 
Neuchâtel, 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 4 zu einem grossflächigen M 1 geführt hat (Petter, 1971, Barysh- 
nikov & Potapova, 1990). Der Metaconus des reduzierten P 4 steht dem Mj -Trigonid 



M j - PROPORTIONSÄNDERUNGEN BEI MELES MELES 21 

gegenüber, der Metaconus des M 1 opponiert zum Talonid des Mj. 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 Mj: 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 et 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 

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Clutton-Brock, J. 1990. Animal remains from the neolithic lake village site of Yvonand IV, 
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22 B. BLÖCHLINGER & P. LÜPS 



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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 für Hans R. Stampfli. Helbing & Lichten- 
hahn, Basel, XVI + 325 pp. 

Stocker, G. & Lüps, 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. In: A Study of Trophic 
Diversity within Guilds (pp. 410-436). In: 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-B ALS 

Museum of Natural History, CP 6434, CH-121 1 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 Trägardh, 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. JUV ARA-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 25 



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 (idjl, 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 (ipl, 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 all simple 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, Vili. 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: A3 14, 19, lo*, 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; XI 551, 19, litter from Ericetum near northern 
slope. Manhartstal near Grossau, 15.9.1960; - Steiermark: T256, 2 9, fir woods, sifting of moss 
under Vaccinium, 1400-1500m, Seckhauer Zinken above Jörgerhütte, 4.6.1960; X1576, le? 
forest litter, Lassnitzklamm near Deuschlandsberg, 14.7.1964 ; X1585, 19, \S , 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; XI 575. 1 9, 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: 7 9, 6o\ 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, la, 
deciduous forest litter, Berzeasca, Almaj Mountain, 15.8.1970, leg. I. Juvara-Bals. Two slides 
deposited in MHNG. - SLOVENIA: Carniola: la, 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. IB). Podonotal region with 19 pairs of setae ( jl-j6; 
zl, z2, z4-z6, si, 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 (idJl, idJ2-double, idJ3, 
idJ4; idZ4, idS4, idZ5, idRl, idS7). Peritrematal region as in figure 1A. 

Length of setae: jl =48-54um; other j setae about 48um, setae of s series about 
42pm, seta r5 =27-30pm, zl = 17um; setae on opisthonotum uniform, their lengths 
about 36-42pm. 

Idiosomal venter. Sternal shield subrectangular, sternal setae moderately long, 
stl =48um, st2 =36pm, st3 =44pm; gland pore gvl 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 



27 




Fig. 1 
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. 
2 A). 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 =48 u, JV3 =30pm, 
others about 36pm. 

Legs. Coxa II with palmate ridge situated anterolaterally; trochanter IV with 
setae pdl and pd2 short, simple. Measurements: Tarsus I =150-172. 5pm; tarsus IV = 
184-195.5mm. Epigynium: height (h) =179-198pm, length of base (b) =172.5-207pm, 
st5-st5 '=1 15-147pm, h/b = about 1. Sternal shield: stl-stl'= 69-75pm, st2-st2' =87- 
96pm, st3-st3' =97- 103pm. 



28 



I. JUVARA-BALS 




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 




Fig. 3 
Male: Heteroparasitus tirolensis (Sellnick): A, B. H. coronarius (Karg): C-F; H. quadratus 
("Witalinskij: 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- JUV ARA-BALS 

Male 

Idiosomal dorsum. Idionotal systems as in female, setal lengths: jl =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 =24pm, stl = 40|jm; 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, 2 9,1c? sifting of alder litter, near 
the Mühlauer river close to a waterfall. Mühlau by Admont, 20.4.1940; A278, 19, lo\ sifting 
of decayed tree stumps, Leichenberg by Admont, southern slope, 1.11.1942; X582, 19, \S, 
sifting litter, beech woods, Damberg near Steyer, northern slope, 27.4.1946; - Burgenland: 
X1690a, 28, 19, beech litter, Mandelstein by Weitra, 1.10.1966; - Niederösterreich: X1649, 
1 9, 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 36-42pm; 
opisthonotal setae 24-36pm. 

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 92pm; opisthogastric setae about 30pm. 

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- 16 lmm, 
tarsus IV = 177-184pm. Epigynium: h =180.5pm, st5-st5' =122pm, b = 149.5pm, h/b 
= 1,21. Sternal shield: stl-stl' = 61pm, st2-st2' = 94.3pm, st3-st3' = 106pm. 

Male 

Idiosomal dorsum. Length of setae: jl = 48-54pm, seta si =12pm, other s 
setae 24-36pm; setae on opisthonotum about 24pm. 

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: stl =42pm, st2 = 36pm, st3 =30pm; length of opisthogastric setae about 24pm. 

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. 
6Kj. 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 =145-152pm; tarsus IV =165.6-177pm. 



32 i- juv ara-bals 

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 H. coronarius. Holo- 
parasitus rotulifer is another, valid species related to the Holoparasitus s.str. species, 
whose males are provided with an excipulum. Males of the related H. 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 H. tirolensis, distinguishes females; other differen- 
tiating characteristics are the shape of the endogynium and of the metagynial sclerites. 
Karg (1971) thought that H. 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, 2c? Myslenice near Krakow, Southern Poland, type 
locality, litter in a mixed forest, 10.1 1.1969, 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- 136pm; length of sternal shield setae 60pm. 

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 zi, si, 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 gvl, 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 pll stout and pilose, tarsus with seta pd2 very long. 

Heteroparasitus (Medioparasitus) athiasae sp. n. 

Type material: SPAIN: Sp408 slide J430 1 6, holotype; paratypes: 39, 3 6, Montes del 
Invernardero near Verin (Provincia Orense), Sierra de San Married, Campobecerros, sifting of 
litter near a stream, 24.7.1955 . 

Other material: SPAIN: Sp42, 23, Ceiro de Mirador, Sierra de Luna near Algesiras 
(Andalusia), sifting of litter and humus under Erica sp., 28.2.1951; Sp211, 2$, 1 <3\ sur- 
rounding of Pontevedra, Cuesta del Ralo Saluda, sifting of litter under Cyprus, 1.7. 1952; 
Sp483, 13, 1 2 , Isela de One, Prov. Pontevedra, sifting of litter under Ulex europaeus, 
4.8.1956: Sp485, 1 9, Near the road Gondomas-Tuy (Sierra de la Grora), Bayona, sifting of 
litter (Quercus sp. forest), 5.8. 1956; Sp500, 2 ? , surroundings of Ezaro, south of Cabo Fisterra, 
sifting of litter (Quercus robur forest), 15.8.1956; Sp571, 23 , Sierra de Son (Leon) near San 
Feliz de las Lavanderas, Astorga, sifting of litter under Quercus and Erica arborea, 1 1.8.1957; 
Sp567, 3 2, 3<3\ Rio Duerna valley, Molina Ferreda near Astorga, sifting of litter of Quercus 
pyrenica and Erica arborea, 10.8.1957; Sp579 1 o\ 4? Near San Saturino (Lugo), sifting of 
litter under Rubus, 15.8.1957; Sp587 1 6, El Fito, West of Aviles, (Oviedo), sifting of litter 
under Betula sp., Quercus sp., 16.8.1957; Sp496, 66 , 12, 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. 



34 



I. JUVARA-BALS 




Fig. 4 
Medioparasitus athiasae sp. n. Female: A-idiosoma, dorsal view; B-peritrematal region; Ci- 
sternal shield, paragynia and endogynium; D- epigynium; E- endogynium, F-opisthogaster. 



A REVISION OF HETEROPARASITUS 



35 




Fig. 5 
Medioparasitus athiasae sp. n. Male: A-H. Female: I-M. A-sternogenital region; B-genital 
region with subgenital sclerite (sg. sci.); 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 




Fig. 6 
Male. Mediopavasitus athiasae sp. n.: A-I; Heteroparasitus coronarius (Karg): J, K, L; H. 
tirolensis (Sellnick) : M, N; H. quadratus (Witalinski): 0, 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 ridse, 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 jl =42- 
45um, si =6-7um; r2, r3, s2 = 13-14um; other setae about 42-48um; opisthonotal 
setae from 48um (J3, S3, S4) to 54 urn (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 gvl 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. 51). 
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 vl 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: stl- 
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' =108-1 15pm, 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: jl =36pm, zl =12um, s2- 
s3 =18um, s6 =48um, J3-J4 =54um, other setae of series z =36-42pm. Opisthonotal 
setae of series J and S =48pm, setae of series R =42pm. Adenotaxy: gland pores gdj2, 
gdj4, gdz6, gdZl 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. 5 A). 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. JUV ARA-BALS 

(Fig. 5B). Opisthogastric shield with 7 pairs of ventral setae, their length from 24 to 
36pm. Gland pore gvl 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 
=159u. 

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 
II-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 peri tre - 
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 (a), the genital orifice (S) and the epi- 
gynium and endogynium (9). 

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 gv 1 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) 

1 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 urn, 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- 
soma length 450-1400um 2 



40 I- JUV ARA-BALS 

2 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 urn . . . 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 2060pm 3 

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 
chelicera with three or four teeth 5 

4 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 prestemal sclerites, 
epigynium triangular or subpentagonal; femur II of male with triangular 
or different- shaped apophysis; idiosoma oval-shaped, length 880 to 
2060pm 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 

700pm 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 
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. 

5 Female dorsal and peritrematal shields united anteriorly, peritrematal 
shield fused with opisthogaster; extension of peritrematal shield behind 
sdamata discernible in male; male without transverse suture on idiosoma ... 6 



A REVISION OF HETEROPARASITUS 4] 

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 

dorsum of idiosoma 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- 
soma length 800-1200um 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 350-800pm 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- 1200pm 

Mixogamasus Juvara-Bals, 1972 

Type species: M. intermedins Juvara-Bals, 1972 
Palpgenual setae al land 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 787-896pm, of female 1028- 1300pm 

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 350-800pm, of female 

440-900pm Leptogamasus Trägardh, 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 450pm, 

male unknown Zelogamasus Hennessy & Farrier, 1989 

Type species: Z. oligochaetns 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 700-885pm, of female 

1 170-1300pm Amblygamasus Berlese, 1906 

Type species: Gamasus dentipes Koch, 1839 



42 I. JUV ARA-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 iv5 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 420-1 100pm, of female 450- 

1200pm 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 

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 llpairs 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 et ah, 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 is 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, i.e.: Leptogamasus Träg- 
ardh, 1936 (type species: L. suecicus Trägardh, 1936), Emogamasus 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 Emogamasus, 
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 i.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 withl8-20 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 gvl 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 gv 1 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. Lobi, P. Schwen- 
dinger and C. Lienhard (Geneva, Switzerland). I am indebt to Drs Axel Christian 
(Görlitz, 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 

Athias-Henriot, C. 1967. Observations sur les Pergamasus. I. Sous-genre Paragamasus Hull, 
1918 (Acariens anactinotriches, Parasitidae). Mémoires du Muséum d'Histoire Natu- 
relle série A (Zoologie) 49(1): 1-197. 

Athias-Henriot, C. 1969. Les organes cuticulaires sensoriels et glandulaires des gamasides. 
Poroïdotaxie et adénotaxie. Bulletin de la Société Zoologique de Prance 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 françaises du sous-genre Leptogamasus s. s. (Arachnida, 
Gamasida, Parasitidae: genre Leptogamasus). Annales de la Société entomologique de 
France 8(1): 189-204.^ 

Athias-Henriot, C. 1973. Paragamasus (Meriadenogamasus) franzi (nov. subgen), Perga- 
masidae, neu für Nepal (Arachnida, Gamasida, Parasitina). Verhandlungen der Zoolo- 
gisch-Botanischen Gesellschaft in Wien 1 13: 93-102. 

Berlese, A. 1906. Monographia del genere Gainasus Latr. Redia 3: 66-304. 

Evans, G. O & Till, W. M. 1979. Mesostigmata mites of Britain and Ireland (Chelicerata: 
Acari, Parasitiformes). An introduction to their external morphology and classification. 
Transactions of the Zoological Society of London 35: 139-270. 

Holzmann, C. 1969. Die Familie Parasitidae Oudemans, 1901. Schriftenreihe für Ver- 
gleichende Milbenkunde, Fürth/Bayern. Acarologie 13: 3-55. 

Hyatt, K. H. 1987. Mites of the genus Holoparasitus Oudemans, 1936 (Mesostigmata: Para- 
sitidae) in the British Isles. Bulletin of the British Museum (Natural History), Zoology 
series 52 (4): 139-164. 

International Commission on Zoological Nomenclature. 1999. International Code of Zoo- 
logical Nomenclature. The International Trust for Zoological Nomenclature, London, 
I-XXIX, 1-306. 

Johnston, D. E. & Moraza, M. L. 1991. The idiosomal adenotaxy and poroidotaxy of 
Zerconidae (Mesostigmata: Zerconina). ///.: Dusbabk, F. & Bukva, V. (eds). Modern 
Acarology. SPB Academic, The Hague, 2: 349-356. 

Juvara-Bals, I. 1972. Mixogamasus un nouveau genre de Parasitidae (Acariens anactino- 
triches) de Roumanie. Acarologia 14: 3-114. 

Juvara-Bals, I. 1975. Sur le genre Holoparasitus Oudemans, 1936 et sur certains caractères 
morphologiques de la famille Parasitidae Oudemans (Parasitiformes). Acarologia 17: 
384-409. 

Juvara-Bals, I. 1981. Nouvelle definition du genre Leptogamasus Trägardh, 1936 (Acarina, 
Gamasida, Parasitidae) et description de six nouvelles espèces. Revue suisse de Zoo- 
logie 88(1): 77-93. 

Karg. W. 1971. Acari (Acarina), Milben. Unterordnung Anactinochaeta (Parasitiformes). Die 
freilebenden Gamasina (Gamasides). Tierwelt Deutschlands, Jena Verlag 59: 1-475. 

Karg, W. 1993. Acari (Acarina), Milben: Parasitiformes (Anactinotricha). Cohors Gamasina 
Leach. Raubmilben. Tierwelt Deutschlands, Gustav Fischer Verlag, Jena 59: 1-523. 

Klompen, J. S. H., Keirans, J. E., Filippova, N. A. & Oliver JR., J. H. 1996. Idiosomal lyri- 
fissures, setae, and small glands as taxonomic characters and potential indicators of 
ancestral segmentation patterns in larval Ixodidae (Acari: Ixodida). International Jour- 
nal of Acarology 22 (2): 1 13-134. 

Koehler, H. 2000. Gamasina von TNT-belasteten Standorten ("Werk Tanne", Harz). Ab- 
handlungen und Berichten des Naturkundemuseums Görlitz 72: 1 15-120. 

Krantz, G. W. & Redmond, B. L. 1987. Identification of glandular and poroidal idionotal 
systems in Macrocheies perglaber F. & P. (Acari: Macrochelidae). Experimental and 
Applied Acarology 3: 243-253. 

Lindquist, E. E. 1994. Some observations on the chaetotaxy of the caudal body region of 
gamasinae mites (Acari: Mesostigmata), with a modified notation for some ventro- 
lateral body setae. Acarologia 35: 323-326. 

Lindquist, E. E. & Evans, G. O. 1965. Taxonomic concepts in the Ascidae, with a modified 
setal nomenclature for the idiosoma of the Gamasina (Acarina, Mesostigmata). Memoir 
of the entomological Society of Canada 41: 1-64. 

Lindquist, E. E. & Moraza, M. L. 1998. Observation on homologies of idiosomal setae in 
Zerconidae (Acari: Mesostigmata), with modified notation for some posterior body 
setae. Acarologia 39 (3): 203-226. 



46 I. JUVARA-BALS 



Micherdzinski, W. 1969. Die Familie Parasitidae Oudemans 1901 (Acarina, Mesostigmata). 

Panstwowe Wydawnictwo Naukowe, Krakow, 1-660. 
Oudemans, A. C. 1936. Kritisch Historisch Overzicht der Acarologie. Brill, Leiden, IIIA, 1805- 

1850. 

Schmölzer, K. 1995. Catalogus faunae austriae Teil IX f., U-ordn: Anactinochaeta (Parasi - 
tiformes). Verlag der Österreichischen Akademie der Wissenschaften, Wien, 1-179. 

Sellnick, M. 1968. Zwei neue Pergamasus Arten aus Österreich. 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 Höhlenkunde, der Eiszeitforschung 
und den Naturgebieten, Biologische Serie, Brno, 8: 1-79. 

Witalinski, W. 1972. Mites of the genus Holoparasitus Oudemans, 1936 (Acarina, Parasi- 
tidae). Acta Zoologica cracoviensia 17 (9): 217-238. 



Revue suisse de Zoologie 109 (1): 47-1 13; mars 2002 



A taxonomic revision of the family Oncopodidae III. Further new 
species of Gnomulus Thorell (Opiliones, Laniatores) 

Peter J. SCHWENDINGER 1 & Jochen MARTENS 2 

1 Muséum d'histoire naturelle, case postale 6434, CH-1211 Genève 6, Switzerland. 

2 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. exsudons (Sarawak, Sabah), G. hamatus (Phi- 
lippines), G. kutan (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. ryssiè (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 S0rensen, 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, 1 1 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 in here). 

Four of the six groups distinguished by Schwendinger & Martens (1999b: 979) 
are re-evaluated and five additional groups are added. The aboi'ensis-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 SV11 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. Genève; NSMT National Science Museum, Tokyo; SMF Naturmuseum und 
Forschungsinstitut Senckenberg, Frankfurt; ZMB Museum für Naturkunde der 
Humboldt-Universität, Berlin; ZMC Zoologisk Museum, K0benhavn; ZMH Zoolo- 
gisches Institut und Museum, Universität 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 




10° 



£X^«^>^5^ 



8Z? 



Fig. 1 
Records of Gnomulus species treated in this paper. - 1 Omei Shan (G. sinensis sp. n.), 2 Cue 
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. ijssie 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. kutan sp. n.), 26 Gunung Mulu N. P. (G exsudons 
sp. n.j, 27 Sepilok (G exsudons sp. n.), 28 Sapagaya (G. exsudons 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 S holotype 
(MHNG), leg. W. Schawaller, 19.III.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: 6 (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 III, 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 



51 




Figs 2-8 
Gnomulus sinensis sp. n., S 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 II. 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. 

9 . Unknown. 

Measurements: (6): Body 4.32 long, 3.29 wide; carapace region 1.06 long, 
2.00 wide. - Palp and legs: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


lp 


0.67 


0.96 


0.62 


0.44 


- 


0.91 


3.60 


gl 


0.52 


1.58 


0.77 


0.82 


1.43 


0.74 


5.86 


gli 


0.59 


2.03 


1.01 


1.26 


2.03 


1.11 


8.03 


g III 


0.52 


1.48 


0.79 


0.86 


1.63 


0.59 


5.87 


g IV 


0.64 


1.93 


1.03 


1.31 


2.42 


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 
[Fig. 1(1)]. 



Gnomulus spiniceps sp. n. Figs 10-19 

Material: VIETNAM, Ninh Binh Province, Cue Phuong National Park, 450 m, about 
40 km NW of Ninh Binh, Ô holotype (NSMT-Ad 174), leg. S. Nomura, 15.X.1995. 



52 



P. J. SCHWENDINGER & J. MARTENS 






Fig. 9 
Gnomulus sinensis sp. n., 6 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 




Figs 10- 

Gnomulus spiniceps sp. n., 8 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: 8 (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 






Fig. 19 
Gnomuliis spiniceps sp. n., 6 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. 

9 . Unknown. 

Measurements: (cT): Body 5.46 long, 4.06 wide; carapace region 1.48 long, 
2.31 wide. - Palp and legs: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.84 


1.23 


0.84 


0.62 


- 


1.28 


4.81 


Legi 


0.64 


2.02 


0.96 


1.03 


1.77 


0.76 


7.18 


Leg II 


0.79 


2.66 


1.33 


1.67 


2.53 


1.16 


10.14 


Leg III 


0.59 


2.02 


0.96 


1.08 


1.94 


0.69 


7.28 


Les 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 

(2)]. 

The A5L/-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 105-1 13). 

New material: MALAYSIA (peninsula), Perak, Maxwell Hill near Taiping (type 
locality) [Fig. 1 (11)]. at 1 150 m, 2 â, 1 9, 24.- 25.XI.1999, at 1250 m, 1 6, 23.XI.1999; all 
specimens leg. G. Cuccodoro & I. Lobi (1 S,\ 9 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 S has a distinctly 
narrower apex and a lateral glans sclerite with an inwards pointing apex (only on one 
side, probably deformed; Fig. 22). 

The 9 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 




Figs 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 1 1, S holotype (MHNG), 1 o\ 1 2 paratypes, 3 juv., 24.III.1993; 
G. Jasar, 1720 m, 1 6, 2 9 paratypes, 25.III.1993; trails 4 and 13 (E of Tanah Rata), 1500 m, 1 
6 paratype, 23.III.1993; trail 9 (between Tanah Rata and Robinson Fall), 1400 m, 2 9 
paratypes, 27. III. 1993, all leg. I. Lobi & F. Calarne. - Ringlet, 960 m, 1 juv., leg. T. Jaccoud. 1 
o\ 1 9 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: â (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 III 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 




Figs 23-34 
Gnomulus monticola sp. n., â holotype (27, 28, 30, 32, 33), 6 paratypes (23-26, 29), 9 
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 






Fig. 35 
Gnomulus monticola sp. n., 6 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. 

9 . As the male, no external sexual dimorphism discernible. 



FURTHER NEW SPECIES OF GNOMULUS 59 

Measurements: 6 holotype ( 9 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: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.41 (0.41) 


0.47 (0.47) 


0.38 (0.39) 


0.27 (0.27) 


-- 


0.56 (0.56) 


2.09(2.10) 


Legi 


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) 


Legïï 


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 III 


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) 



LeglV 0.44(0.47) 1.22(1.27) 0.69(0.72) 0.88(0.89) 1.44(1.44) 0.50(0.50) 5.17(5.29) 

Variation: Range of measurements in S â (n=4) and 9 ? (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 â 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 S (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, 6 holotype, 10./1 3.III. 1993, leg. I. Lobi & F. Calarne (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: ê (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 III 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 




Figs 36-45 
Gnomulus pilosus sp. n., 6 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. 

$ . Unknown. 

Measurements: 6: Body 3.45 long, 2.32 wide; carapace region 0.74 long, 1.28 
wide. - Palp and legs: 



FURTHER NEW SPECIES OF GNOMULUS 6 1 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.38 


0.50 


0.39 


0.27 


- 


0.57 


2.11 


Legi 


0.38 


0.98 


0.55 


0.57 


0.82 


0.66 


3.96 


Leg II 


0.49 


1.32 


0.71 


0.88 


1.34 


0.71 


5.45 


Les III 


0.39 


0.98 


0.58 


0.61 


1.07 


0.46 


4.09 



Leg IV 0.49 1.39 0.77 0.96 1.61 0.53 5.75 

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-grovp (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, 6 holotype, 5. II. 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 5 (not a type). 9.VI.2001, leg. P. 
Schwendinger. All specimens in MHNG. 

Etxmologx: The latinized Greek suffix -oidetis 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: 6 (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 




Figs 46-55 
Gnomulus rostratoideus sp. n., S holotype (46-54), 9 (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 III 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. 

9 (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: â 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: 





Tr 


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) 


Legi 


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) 


Leg IV 


0.77 (0.67) 


2.38(1.98) 


1.21 (1.07) 


1.62(1.39) 


2.65 (2.28) 


0.75 (0.63) 


9.38 (8.02) 



Variation: The second 2 measures: Body length 5.26, width 3.22, carapace 
length 1.49, width 1.78. Both 9 from Singapore are smaller than the â 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 6 from Singapore, we 
tentatively place the 2 2 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 6), 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. 



54 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, S holotype (MHNG), 1 6, 3 9 paratypes, 3 juv., 23.- 
30.XI.1989, leg. I. Lobi, D. Agosti & D. Burckhardt (1 9 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 â, with a distinct eye tubercle in both sexes; ventral side of leg coxa II 
with an anteroproximal process, process on coxa III larger; trochanter III 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: S (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 III, 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 




Figs 56-72 
Gnomulus tuberculatus sp. n., 6 holotype (60, 61, 62, 63, 66-68), S 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). 



9 . 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: S holotype (9 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) 

Legi 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) 

Legll 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 III 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 m S S (n=2) and 9 9 (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 9 it is almost pointed 
(Fig. 71). 

Relationships: Genital morphology and certain characteristics of chelicerae, 
palps and leg coxae I indicate that Gnomulus tubereulatus 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 (Thoreil), G. drescoi (Silhavy)], two from Java 
[G. javanicus sp. n., G. thorelli (S0rensen) (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. kutan 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, 6 holotype, 2 S, 1 9 paratypes, 12.XI.1998. 
- Trat Province, Laem Ngop District, Ko Chang (= Elephant Island) National Park, forest above 
White Sand Beach, 20-50 m, 3 6 paratypes, 6.-8.IX.1993, 1 juv., 24.VIII.1992 (all specimens 



FURTHER NEW SPECIES OF GNOMULUS 



67 






Fig. 73 
Gnomulus tuberculatus sp. n., 6 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 6 paratype, 
3.-23.XII.1999 (leg. A. Schulz). 1 S 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 




Figs 74-90 
Gnomulus marginatus sp. n., 6 holotype (74-77. 85-88), 6 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). 



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 






Fig. 91 
Gnomulus marginatus sp. n., S 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: â (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. 

Legsl324, 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. 

9 . 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: â holotype (9 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 


Ti 


Mt 


Ta 


Total 


Palp 


0.89 (0.89) 


1.48(1.40) 


1.08(1.08) 


0.81 (0.79) 




1.55(1.53) 


5.81 (5.69) 


Legi 


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) 


Leg II 


0.89 (0.86) 


3.17(3.15) 


1.48(1.48) 


2.21 (2.12) 


3.25 (3.20) 


1.33(1.28) 


12.33 (12.09) 


Leg III 


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 ô â (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 â paratype with a 
bifid ventral tooth on proximal cheliceral article (Fig. 84) and with a truncate ventral 
process on palpal trochanter (Fig. 90). 

Relationships: Gnonudus 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 7 ] 

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, S holotype, 17.XI.1985, leg. I. Lobi & 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: 6 (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 II 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. 

$ . Unknown. 



72 



P. J. SCHWENDINGER & J. MARTENS 




Figs 92-101 
Gnomulus ryssie sp. n., S 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: (<?): Body 6.22 long, 4.38 wide; carapace region 1.13 long, 
2.21 wide. - Palp and legs: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.69 


1.03 


0.83 


0.59 


- 


1.15 


4.29 


Legi 


0.54 


1.92 


0.89 


0.98 


1.62 


0.79 


6.74 


Leg II 


0.64 


2.51 


1.13 


1.77 


2.51 


0.98 


9.54 


Leg III 


0.57 


1.92 


0.98 


1.18 


2.02 


0.59 


7.26 


Leg IV 


0.74 


2.56 


1.21 


1.80 


3.10 


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: MYANMAR, Tenasserim, Malewoon (= Maliwun), 6 holotype (MSNG, Nr. 
10203), lee. 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: S (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 III 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. 1 10): 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. 

9 . Unknown. 

Measurements: (<?): Body 5.74 long, 4.21 wide; carapace region 1.44 long, 
2.13 wide. - Palp and legs: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.87 


1.24 


0.99 


0.64 


- 


1.29 


5.03 


Legi 


0.69 


2.45 


1.16 


1.34 


2.08 


1.06 


8.78 


Leg II 


0.82 


3.37 


1.58 


2.33 


3.22 


1.34 


12.66 


Leg III 


0.72 


2.50 


1.29 


1.56 


2.57 


1.14 


9.78 


Leg IV 


0.89 


3.32 


1.56 


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 




Figs 102-111 
Gnomulus leofeae sp. n., 6 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 S specimen was collected on Gunung (= Mount) Kerinci 
(2160 m, 17.-1 8. 11.2000, leg. P. J. Schwendinger, MHNG), Jambi Province, Sumatra 
[Fig. 1 (20)], which largely accords with the S from the same locality mentioned by 
Schwendinger & Martens (1999b: 963, 965, figs 1 14, 122). The new S 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 

Pelitmis segnipes Loman. - Loman (1902: 182, partim). - Roewer (1923: 63, partim). 

Material: INDONESIA, Java, Mt. Gede, SE of Bogor, 3 holotype (ZMH, with labels: 
"Pelitmis segnipes, Loman det. 1901/02, H. Fruhstorfer vend. 18.11.1897" and "Pelitnus 
segnipes, Roewer det. 1914, No. 1258"). - Java, without exact locality, 2 5 paratypes (SMF 
1602, with label: "Pelitmis javanus n. sp. Roewer, typus, 1 â , 1 9 , 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: 6 (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. 

9 . 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 



116 . i 117 ( \ 




Figs 112-124 
Gnomulus javanicus sp. n., 6 holotype (112-117, 119, 124), 9 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 (1 14, 1 15), 1.0 mm (others). 



ventral scutal areas not swollen. Legs 1324; distitarsus II about 2.1 times longer than 
wide (Fig. 121). 

Measurements: â holotype (9 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: 





Tr 


Fe 


Pa 


Ti 


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) 


Legi 


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 


0.69 (0.64) 


3.24(2.32) 


1.53(1.16) 


2.37(1.63) 


3.24(2.32) 


? (1.13) 


? (9.20) 


Leg III 


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) 



Leg IV 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 ? 9 (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 77 

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), 3 holotype, S 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 o\ 1 9, "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: 6 (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 III 
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 




Figs 125-140 
Gnomulus lomani sp. n.. S holotype (125-128. 134. 136, 137), 9 paratypes (129-130, 139), 
presumably conspecific S (138) and 9 (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. 

9 (uncertain identification). As the male 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: 6 holotype (a paratype in parentheses; no measuremets of 
available 9 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 


- 


1.12 


4.51 


Legi 


0.56 


2.06 


1.06 


1.25 


1.87 


0.94 


7.74 


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 2.12 3.40 1.18 11.98 

Variation: The 6 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 kutan 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 ô and 1 9 (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 â 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, 6 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: 6 (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 




Figs 141-149 
Gnomulus obscurus sp. n., 6 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: {8): Body 5.64 long, 3.99 wide; carapace region 1.33 long, 
2.12 wide. - Palp and legs: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.84 


1.11 


0.89 


0.62 


- 


1.23 


4.69 


Legi 


0.64 


2.42 


1.11 


1.36 


2.12 


1.13 


8.78 


Leg II 


0.74 


3.30 


1.53 


2.51 


3.30 


1.58 


12.96 


Leg III 


0.67 


2.47 


1.23 


1.63 


2.56 


1.18 


9.74 


Leg IV 


0.94 


3.30 


1.48 


2.34 


3.77 


1.33 


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, 6 holotype, Ô paratype, 18.V.1994, leg. I. Lobi & D. Burckhardt (MHNG). 
Etymology: Malay and Indonesian: hutan = forest; noun in apposition. 



82 P. J- SCHWENDINGER & J. MARTENS 

Diagnosis: Similar to G. lontani 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 S (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. 

9 . Unknown. 

Measurements: 6 holotype (o* 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 
lees: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


lp 


0.96 


1.30 


0.99 


0.81 


- 


1.55 


5.61 


g I 


0.81 


2.42 


1.21 


1.43 


2.23 


1.24 


9.34 


gli 


0.99 


3.32 


1.55 


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 




Figs 150-163 
Gnomulushutansp.n., S holotype (152, 153, 156-162), 6 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?), 6 holotype, 1 <J, 1 9 paratypes, 1 juv., VI. 1978, leg. F. Wanless 
(NHML). - Sabah, Sandakan Bay (southwest), Sapagaya Lumber Camp, 2-20 m, 1 S paratype 
(BMH; penis dissected and retained by W. A. Shear), 7 9 paratypes, 2 juv., 4.-7.XI.1957, leg. 
J. L. Gressitt (BMH); Sandakan Bay (northwest), Sepilok Forest Reserve, 1-10 m, 7 o\ 5 9 



84 P- J- SCHWENDINGER & J. MARTENS 

paratypes, 1 juv., 30.X.1957, leg. J. L. Gressitt (1 <?, 1 9 paratypes in MAR, 1 o\ 1 2 
paratypes in MHNG, others in BMH). 

Etymology: Latin: exsudons (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: S (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 II. 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. 

9 . 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: 6 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 




Figs 164-184 
Gnomulus exsudans sp. n., 6 holotype (164, 165, 167, 168, 177. 179, 180), paratypes: S from 
the type locality (181), 6 6 from Sabah (166, 169-176, 182), 9 from the type locality (178, 
183), 9 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 






Fig. 185 
Gnomulus exsudans sp. n., <S 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) 


Legi 


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 (2.39) 


3.91 (3.76) 


1.43(1.40) 


13.75(13.31) 



FURTHER NEW SPECIES OF GNOMULUS 87 

Variation: Range of measurements in S â (n= 10) [9 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 â 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), 6 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 



gg 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: â (holotype). Coloration: Body reddish brown, colour pattern 
faded; legs yellow-brown, tarsi I, II 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. 

? . Unknown. 

Measurements: (â): Body 6.00 long, 3.98 wide; carapace region 1.23 long, 
2.16 wide. - Palp and legs: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


lp 


0.79 


1.13 


0.89 


0.59 


- 


1.13 


4.53 


g I 


0.64 


2.02 


1.08 


1.18 


1.82 


0.84 


7.58 


gli 


0.74 


2.66 


1.38 


1.82 


2.61 


1.28 


10.49 


gin 


0.69 


2.02 


1.08 


1.28 


2.21 


0.84 


8.12 



Leg IV 0.79 2.56 1.38 1.87 3.35 0.93 10.88 

Relationships: Gnomulus carinatus sp. n. appears most closely related to G. 
thorelli (6 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 




Figs 186-190 
Gnomulus carinatus sp. n., S 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, 6 holotype, 24.XI.1995. - Mt. Makiling, south of Los Banos, near 
Mad Springs, 400-450 m, ? paratype, 19.XI.1995; leg. I. Lobi (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 (6 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 




Figs 191-203 
Gnomulus hamatus sp. n., S holotype (191-197, 199-202), 9 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: S (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 91 

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. 

9 . 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: â (9 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) 


Lei II 


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 III 


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) 


Leg IV 


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, S holotype, 3 9 
paratypes, 6.-7. XI. 1999; leg. A. Riedel (1 9 paratype in MAR, others in MHNG). 

Etymology: Latin: tumidiis = 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: â (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 III 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 




Figs 204-215 
Gnomulus tumidifrons sp. n., 6 holotype (204-208, 210-212), 2 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. 

9 . As the male, apart from very slightly less pallid ventral scutal elevations. 

Measurements: â (9 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) 


Legi 


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 $ ? (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 is 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 






Fig. 216 
Gnomulus tumidifrons sp. n., 6 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, S 
holotype, 2 juv., 1. XI. 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: â (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 II 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. 

9 . Unknown. 

Measurements: â: Body 3.59 long, 2.35 wide; carapace region 0.93 long, 1.29 
wide. - Palp and legs: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.55 


0.58 


0.46 


0.32 


- 


0.64 


2.55 


Legi 


0.37 


0.97 


0.55 


0.55 


0.86 


0.55 


3.85 


Leg II 


0.47 


1.24 


0.72 


0.89 


1.39 


0.67 


5.38 


Leg III 


0.42 


0.84 


0.56 


0.58 


1.08 


0.47 


3.95 


Leg IV 


0.52 


1.19 


0.72 


0.95 


1.55 


0.52 


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 




Figs 217-224 
Gnómulus matabesar sp. n., 6 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 II, retrolateral view (224). - 
Scale lines 0. 1 mm (2 1 9, 220), 1 .0 mm (others). 

Distribution: Known only from the type locality in the northern part of 
Halmahera Island [Fig. 1 (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, S holotype, 6 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 9 paratypes, 5.-9.XII.1999, leg. A. Riedel (MHNG). 

Etymology: Latin: latus = wide, extensive, operculum = cover, lid; noun in apposition. 
Lhe 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 




Figs 225-239 
Gnomulus latoperculum sp. n., 6 holotype (225-230, 237), 6 paratype (234-236), 5 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: 6 (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 






Fig. 240 
Gnomulus latoperculwn sp. n., 6 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 III 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. 

9 . 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: 6 holotype (9 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: 





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 II 


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 III 


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 â <5 (n= 2) and 9 9 (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 S 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 9 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 9 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. imadateil); 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, 6 holotype, 24.XI.1995, leg. I. Lobi (MHNG). - Los Banos, Mt. Makiling (= Mt. 
Maquiling), 2 6 paratypes, XI. 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: <S (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 




Figs 241-254 
Gnomulus claviger sp. n., 6 holotype (245, 246, 254), S 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 



102 



P. J. SCHWENDINGER & J. MARTENS 






Fig. 255 
Gnomulus claviger sp. n., S paratype. - Body, dorsal (a), ventral (b) and lateral view (c). 
Scale line 1 .0 mm. 



FURTHER NEW SPECIES OF GNOMULUS \ 03 

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. 

$ . Unknown. 

Measurements: S paratype: Body 2.45 long, 1.56 wide; carapace region 0.62 
long. 0.94 wide. - Palp and legs: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.31 


0.40 


0.33 


0.20 


- 


0.50 


1.74 


Legi 


0.27 


0.68 


0.40 


0.40 


0.57 


0.42 


2.74 


Leg II 


0.35 


0.93 


0.51 


0.60 


0.88 


0.51 


3.78 


Leg III 


0.29 


0.64 


0.41 


0.40 


0.73 


0.33 


2.80 


Leg IV 


0.35 


0.93 


0.56 


0.68 


1.01 


0.37 


3.90 



Variation: Measurements, external and genital characters of all three 6 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. is 
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, 6 holotype, 2- S, I- 5 paratypes, 1 juv., 26.XI.1995; summit trail, 800 m. 1 o* 
paratype, 25.XI.1995; above Kinabuhayan, trail to Cristalino, 600-700 m, 3 9 paratypes, 
24.XI.1995. All specimens leg. I. Lobi (1 o\ 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: â (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 III 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 III 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 III not incrassate (Figs 27 1 , 272). 

Measurements: â holotype (9 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 (0.33) 


1.17(1.19) 


Legi 


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 III 


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 6 S (n = 4) and 9 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 is narrowly truncate instead of pointed 
(Fig. 262). 



FURTHER NEW SPECIES OF GNOMULUS 



105 




Figs 256-272 
Gnomulus crassipes sp. n., 6 holotype (260, 261, 266, 267-270), 6 paratypes (256-259, 262, 
263), ? paratype (264, 271, 272). - Penis, dorsal (256) and lateral view (257); apex of penis, 
dorsal (258, 260, 262) and lateral view (259, 261); glans penis, lateral view (263). Left 
chelicera, retrolateral view (265, 266); left palp, retrolateral view (267); distal part of left leg II, 
retrolateral view (268); distal part of left leg III, dorsal (269, 271) and lateral view (270, 272). - 
Scale lines 0.1 mm (258-263), 0.5 mm (others). 



Relationships: Modifications of the glans (especially of its stylus) and a distad- 
directed ventral process on the palpal trochanter place G. crassipes sp. n. in the 
goodnighti-group. Congruence in external morphology (tarsal formula 2-2-2-2, no 
carapace-abdomen bridge) and penis morphology (especially the reduction of the 
median plate) suggest closest relationship with G. crucifer. 

Distribution: Known only from a mountain on Luzon Island [Fig. 1 (34)J, 
where the new species occurs together with G. claviger sp. n. and G. hamatus sp. n. 



106 



P. J. SCHWENDINGER & J. MARTENS 






Fig. 273 
Gnomulus crassipes sp. n., S paratype. - Body, dorsal (a), ventral (b) and lateral view (c). 
Scale line 1.0 mm. 



FURTHER NEW SPECIES OF GNOMULUS ] QJ 

UNIDENTIFIED MATERIAL 

THAILAND: 1 juv. (leg. L. Deharveng; MHNG) was collected in front of 
Tham (= Cave) Pu Lub, ca. 300 m, near Phu Kradung village, Khon Kaen Province, 
northeastern Thailand [Fig. 1 (3)]. 

2 9 (leg. I. Lobi & D. Burckhardt and P. J. Schwendinger; MHNG), similar to 
G. ryssie sp. n. and G. marginatus sp. n., were found on Khao (= Mount) Khieo, 1 150 
m, in the Khao Yai National Park, Nakhon Ratchasima Province, northeastern 
Thailand [Fig. 1 (4)]. 

4 9 (leg. M. Andersen & A. R. Rasmussen; ZMC) from Ko (= Island) Siray 
off Phuket Island [Fig. 1 (9)] appear to be close to G. marginatus sp. n. and G. ryssie 
sp. n. 

1 9 (leg. A. Schulz; MHNG), very similar to the 4 9 from Ko Siray, was 
found in the Khao Sok National Park (Phang Nga Province; not indicated on Fig. 1). 

MALAYSIA (peninsula): 1 9 (leg. P. J. Schwendinger; MHNG), belonging to 
the <75//-group, was sifted from leaf litter near Jeram Pasu Waterfall, ca. 30 km south 
of Kota Baharu, Kelantan [Fig. 1 (10)]. 

1 9 (leg. A. Riedel; MAR) from Maxwell Hill, 1200 m, Taiping, Perak (see 
Martens & Schwendinger, 1998: 549) [Fig. 1 (11)] occurs syntopically with G. laru- 
ticus but clearly does not belong to any of the known <7.s7/-group species. It possesses 
an unusually large genital operculum. 

1 juvenile (leg. P. J. Schwendinger; MHNG) from a forest near Chenderiang 
[Fig. 1 (12)] possibly belongs to G. pulvillatus (see Schwendinger & Martens, 1999b: 
957). 

MALAYSIA (Sarawak): 2 9 (leg. I. Lobi & D. Burckardt; MHNG) from 
Santubong Peninsula (0-100 m), N of Kuching [Fig. 1 (23)], appear to be close to G. 
laevis, but are distinguished by the lack of ventral processes on palpal femur and 
trochanter. 

1 9 (leg. I. Lobi & D. Burckhardt; MHNG) from near Kapit (the type locality 
of G. hutan sp. n.) [Fig. 1 (25)] is distinct by the lack of a ventral process on palpal 
trochanter and by the rare tarsal formula 2-2-2-2. 

MALAYSIA (Sabah): 1 9 (leg. R. Leakey?; NHML) from Mount Kinabalu 
National Park, 1540 m [Fig. 1 (29)] and a large juvenile (leg. W. Schawaller; MAR) 
from the same mountain at 1500-1600 m externally largely correspond with G. 
exsudans sp. n., but presumably belong to a different species. 

The same also applies to 4 juveniles (leg. P. Lehtinen; Zoological Museum 
Turku) from Tiger Hill near Tawau, southeastern Sabah [Fig. 1 (30)]. 

INDONESIA (Sumatra): 1 9 (collector unknown; MHNG) from Deli near 
Medan, Northern Sumatra Province [Fig. 1 (19)] seemingly belongs to the siana- 
tranus-group. A juvenile (leg. A. Riedel; MHNG) from Bukit Lawang (west of 
Medan) [Fig. 1 (18)] may also be conspecific. 

INDONESIA (Kalimantan): 1 9 (leg. P. Beron & T. Ivanova; National 
Museum of Natural History, Sofia) from Nunukan Island (near the border to Sabah) 
[Fig. 1 (31)] is externally similar to G. exsudans sp. n. but possesses a distinctly larger 
body and a lower eye tubercle. 



108 p - J - SCHWENDINGER & J. MARTENS 

INDONESIA (Moluccas): 1 juv. (leg. A. Riedel; MHNG) from Morotai Island 
(north of Halmahara) [Fig. 1 (37)] is similar to the species of the tumidifrons-gvoup 
but has a different colour pattern than the juveniles from Halmahera. 

INDONESIA (Irian Jaya): 19,1 juv. (leg. A. Riedel; MHNG) from Gunung 
Susu on Waigeo Island is externally similar to G. iumidifrons. 

PHILIPPINES (Luzon): 1 9 (leg. L. Deharveng; MAR) from near Sagada 
[Fig. 1 (32)] resembles G. crucifer but differs by larger size and the common tarsal 
formula 2-2-3-3. 

1 juv. (leg. L. Deharveng; MAR) from the Quezon National Park [Fig. 1 (33)] 
clearly differs from the species found at the nearby mountains, Mt. Makiling and Mt. 
Banahaw. 

PHILIPPINES (Leyte): 1 9 (fairly large; mentioned in Martens & Schwen- 
dinger 1998: 549) and 1 juv. (leg. J. Martens & W. Schawaller; MAR), from north of 
Baybay [Fig. 1 (36)] differ from G. leyteensis, which occurs at the same locality. 

DISCUSSION 

The species groups 

At present 48 Gnomulus species are known, which we arrange in 1 1 preli- 
minary species groups. Two of them {aborensis-gxoxxp, rostratus-group) were treated 
earlier (Schwendinger & Martens, 1999b) and are not further considered here. 

1. Gnomulus sinensis sp. n. and G. spiniceps sp. n., here placed in the sinensis- 
group, are both close to the aborensis-group. Discoveries of further new species in the 
northern part of the distribution area of Gnomulus will most likely close the narrow 
gap between these two groups. 

2. In the most species-rich assembly, the armillatus-group with 20 spp., details 
of penis morphology indicate four different phyletic lineages: 

A) Gnomulus piliger from southern Thailand, G. leofeae sp. n. from southern 
Myanmar, G. pulvillatus from peninsular Malaysia, G. armillaius from Sumatra and 
G. carinatus sp. n. from southern Kalimantan all possess narrow, cylindrical, pointed, 
slightly sigmoid lateral glans sclerites pointing away from the truncus, in addition to a 
short, broadly rounded median plate. 

B) Gnomulus lyssie sp. n. and G. marginatus sp. n. from central and south- 
eastern Thailand are similar but have a long, more or less pointed median plate. 

C) Gnomulus baham from Brunei and G. conigerus from Sabah also have 
cylindrical, pointed lateral sclerites, but they are U-shaped instead of sigmoid and 
their median plate is quite long and pointed. Genitalic characters of all these species 
appear to be plesiomorphic, as cylindrical, pointed lateral sclerites are also present in 
the primitive Palaeoncopus. Therefore these three subgroups are possibly not mono- 
phyletic. 

D) Gnomulus javanicus sp. n. (from Java), G. exsudans sp. n., G. kutan sp. n., 
G. laevis, G. lontani sp. n., G. obscurus sp. n. and G. sundaicus (from northern 
Borneo), on the other hand, all possess paddle-shaped or blade-shaped lateral sclerites 
with laterally compressed apices, which are probably apomorphic. These 7 Gnomulus 



FURTHER NEW SPECIES OF GNOMULUS 109 

species, presumably together with G. annulipes (the apices of its small and hook-like 
lateral sclerites are not clearly compressed), seem to form a distinctly monophyletic 
group. 

In G. thorelli from Java and G. drescoi from Sumatra the males are still 
unknown and therefore their relationships remain unclear. We assume that the former 
species is close to G. carinatus sp. n. and the latter close to (or even identical with) 
G. armillatus. 

3. The <7s//-group is a distinct, seemingly quite primitive lineage, restricted in 
its distribution to peninsular Malaysia. The fortunate find of further G. laruticus 
specimens confirms that the unusual tarsal formula 2-2-2-2 is in fact a specific 
character and not just a deformity of the holotype. The same is presumably also the 
case in the unrelated G. crucifer (with the same tarsal formula) from the Philippines. 

4. Gnomulus tuberculatus sp. n., the second species of the s umatranus- group, 
makes an external distinction from the armillatus-group and from the aborensis-group 
less clear-cut than before. With regard to penis morphology, however, the two species 
of the sumatranus-group show more congruence with the externally much different 
asli-group species. The synapomorphic presence of a subdistal process on the ventral 
side of their palpal femur, in addition to their large size, show that G. sumatranus and 
G. tuberculatus sp. n. belong to a distinct lineage and do not just present links 
between the armillatus- (or aborensis-) group and the asli-group. 

5. Gnomulus rostratoideus sp. n. is externally almost indistinguishable (except 
for an unbroken dark margin around the dorsal scutum) from the species of the 
rostratus-group, which are characterised by a strong, beak-like, forward-inclined eye 
tubercle. The penis of the new species, however, does not show the same derived 
morphology (i.e. stylus shaped like a flattened bell). Instead it resembles the penes of 
the goodnighti-group species in that its cylindrical stylus lacks ventral subterminal 
teeth. This congruence is considered to be due to convergent reduction. Therefore G. 
rostratoideus sp. n. is probably a primitive relative of the species in the rostratus- 
group. We place it here in a species group of its own, the rostratoideus-group, and 
regard it as sister to the distinct and clearly monophyletic rostratus-group. 

6. Gnomulus hamatus sp. n., the only representative of the hamatus-group, 
possesses several characters which appear to be primitive for the genus Gnomulus 
(i.e. medium body size, an only slightly distad-inclined ventral process on palpal 
trochanter and stylus penis slender, with bulbous base and with a pair of subterminal 
ventral teeth). The absence of a carapace-abdomen bridge and the presence of pro- 
nounced modifications on the truncus penis and certain parts of the glans (i.e. sub- 
distally wide truncus, enlarged membraneous socket, truncate lateral sclerites, hook- 
shaped median plate), however, show that this species is much closer to the good- 
nighti-group than to the armillatus-group. Therefore we place G. hamatus sp. n. in a 
species group of its own and regard it as the sister of the goodnighti-group. 

7. With 1 1 species known at present, the goodnighti-group is second to the 
armillatus-group in species-richness but it is more diverse with regard to penis 
morphology. Moreover, the group has a fairly restricted distribution in the Philippines 
and northern Borneo, which indicates that this is a relatively young phylogenetic 
lineage undergoing rapid speciation. This group is probably paraphyletic. 



HO P. J. SCHWENDINGER & J. MARTENS 

Gnomulus goodnighti and G. leyteensis show the primitive situation of 
retaining a ventral pair of subdistal teeth on their stylus penis. Otherwise both species 
perfectly correspond with others from this group, mostly because their styli are dis- 
tinctly enlarged. Gnomulus coniceps and seemingly also G. imadatei (not examined; 
see Suzuki, 1969: fig. 4d, e), on the other hand, lack subdistal teeth on their styli (and 
are otherwise quite typical for this group), but these styli appear to be slender, not 
enlarged. Hence, the presence of an enlarged stylus or the absence of subterminal 
teeth on the stylus alone are not exclusive characteristics of the goodnighti-group (see 
also the rostratoideus-, tumidifrons- and latoperculum-growp). 

Gnomulus crassipes sp. n. and G. crucifer are related by the unusual tarsal for- 
mula 2-2-2-2, by the reduction of their carapace-abdomen bridge and by similar 
modifications of the glans penis. The lack of a carapace-abdomen bridge appears to 
link them with G. minor (male unknown) and with the tumidifrons-group, but this is 
not reflected in a similar penis morphology. The carapace-abdomen bridge probably 
has been reduced parallel in different lineages, i.e. in the goodnighti-group, in the 
tumidifrons-group and in the hamatus-group. 

8. Gnomulus tumidifrons sp. n. and G. maiabesar sp. n. from the Moluccas, 
here placed in the tumidifrons-group, clearly represent a distinct phyletic lineage close 
to the goodnighti-group. Small body size, a distad-directed ventral process on palpal 
trochanter, fairly long membraneous tubes in the glans and an enlarged stylus are 
shared by both groups. The two species of the tumidifrons-group are characterized by 
large eyes, a glans penis with fairly large, golfclub-shaped membraneous tubes, which 
are not covered by the narrow, spike-like median plate. Similar membraneous tubes 
are also found in G. latoperculum sp. n. The tumidifrons-group is possibly an offshoot 
of the goodnighti-group, which in this case would be paraphyletic. 

9. Gnomulus latoperculum sp. n. (latoperculum-group) appears intermediate 
between three species groups. In external morphology it resembles species of the 
armillatus-group (by its fairly large size, its wide, undivided carapace-abdomen 
bridge and by a slightly distad-inclined ventral process on palpal trochanter), but its 
penis shows modifications otherwise mostly found in the goodnighti-group (i.e. 
enlarged cylindrical stylus with invaginated base and without subdistal pair of teeth, 
long membraneous tubes, reduced median plate, lobate lateral sclerites). As in the 
case of the tumidifrons-group, G. latoperculum sp. n. is presumably also a highly 
derived descendant of the (then paraphyletic) goodnighti-group. Most similarities in 
penis morphology are found between G. latoperculum sp. n. and G. claviger sp. n., 
but we consider these as convergent. A close relationship possibly also exists between 
Gnomulus latoperculum sp. n. and both species of the tumidifrons-group from the 
Moluccas, as indicated by geographical proximity and by the long, distally flattened 
and widened (golfclub-shaped) membraneous tubes of the glans penis. The latter may, 
however, be yet another case of convergence. 

Zoogeography 

Gnomulus has clearly the largest distribution of all oncopodid genera, covering 
the entire range of the family (Fig. 1 and Schwendinger & Martens, 1999b: fig. 1). 
The newly discovered species considerably expand the previously known distribution 



FURTHER NEW SPECIES OF GNOMULUS \ \ \ 

of the Oncopodidae towards the northeast and the southeast. Gnomulus sinensis sp. n. 
marks the northeasternmost occurrence in this family and G spiniceps sp. n. stands at 
the eastern periphery of the known distribution on mainland Asia. Looking at the 
sparse and widely separated oncopodid records in this region (with the exception of a 
relatively well-investigated peninsular Malaysia), one can see how little we still 
known about the diversity of these enigmatic opilionids, let alone about their biology. 
These patchy records are not least due to restricted accessability of certain areas, 
either because of the political situation in Myanmar and northeastern India, or because 
of dangers from land mines and unexploded bombshells in Indochina. Unfortunately 
this restricts zoological investigations in situ, but not large-scale habitat destruction in 
these areas. 

The most surprising discovery from the new material is the occurrence of 
oncopodids beyond Wallace's line, i.e. Gnomulus latoperculum sp. n. in northern 
Sulawesi, G matabesar sp. n. and G tumidifrons sp. n. in the northern Moluccas and 
an unidentified species (only 9 and juvenile available) on an island off the north- 
western tip of New Guinea. Considering that oncopodids require constantly high 
humidity and looking at the relationships between these three nominal species and at 
their taxonomic distinctiveness from congeners west of Wallace's line, we can 
exclude that they have been introduced by man. The closest relatives of these three 
species are found in the Philippines and not on the Greater Sunda Islands. Therefore it 
appears that the ancestors of these autochthonous taxa have arrived from the north, 
probably in fairly recent geological times. It will be exciting to learn from further 
collections on the Lesser Sunda Islands and on New Guinea, how far the Oncopo- 
didae have advanced into the Australian region. 

The different species groups occupy quite distinctly outlined geographical 
ranges with only few disjunctions. The aborensis-gvoxxp species occur in and around 
the Himalayas and its foothills extending into northern Thailand, whereas the 
sinensis-group occupies the northeastern part of the generic range on mainland Asia, 
i. e. southern China and northern Vietnam. The asli-group and the rostratus-group are 
known only from peninsular Malaysia (new finds for the latter also in Thailand), the 
sumatranus-group only from Sumatra and the tumidifrons-group so far only from the 
Moluccas (probably also from Waigeo Island off New Guinea). The monotypic 
rostratoideus-group, latoperculum-group and hamatus-gvowp occur in southern penin- 
sular Malaysia, on Sulawesi and on Luzon Island (Philippines), respectively. The 
goodnighti- group is found on the Philippine islands and one of its species also on 
Borneo. Only the armillatus-group has a wide distribution, ranging from northeastern 
Thailand to Sumatra, Java and probably the whole of Borneo. Most species appear to 
have small distribution areas, only two members of the armillatus-group are note- 
worthy exceptions: The known range of G armillatus stretches over 170 km, that of 
G. exsudons sp. n. over about 400 km. 

Remarkable is the seemingly syntopical occurrence of three congeneric species 
(G hamatus sp. n., G claviger sp. n. and G crassipes sp. n.) on Mt. Banahaw, Luzon 
Island. The latter two species even belong to the same derived species-group. From 
the nearby Mt. Makiling also three species are known, i.e. G hamatus sp. n., G 
claviger sp. n. and G minor. In this case, however, it is not clear whether the latter 



112 P- J- SCHWENDINGER & J. MARTENS 

species (male unknown) is really congeneric with the other two. At the same moun- 
tain there is additionally an undescribed Biantoncopus and an oncopodid species, 
which at present we cannot attribute to any genus. The presence of presumably yet 
another Gnomulus species (1 juvenile available) in the nearby Quezon National Park 
shows that the Oncopodidae have experienced an exceptionally vivid process of 
speciation in the Philippines and on Luzon Island in particular. 

ACKNOWLEDGEMENTS 

We gratefully acknowledge loans and donations of specimens from the 
following colleagues: Petar Beron (National Museum of Natural History, Sofia), 
Giulio Cuccodoro and Ivan Lobi (MHNG), Hieronymus Dastych (ZMH), Louis 
Deharveng (Laboratoire d'Ecologie Terrestre, Toulouse), Giuliano Doria (MSNG), 
Jason Dunlop (ZMB), Manfred Grasshoff (SMF), Janet Beccaloni and Paul Hillyard 
(NHML), Pekka Lehtinen (Zoological Museum Turku), Hirotsugu Ono (NSMT), 
Norman Platnick (AMNH). Nikolaj Scharff (ZMC), Wolfgang Schawaller and 
Alexander Riedel (Staatliches Museum für Naturkunde, Stuttgart), Andreas Schulz 
(Leichlingen). J. Dunlop also provided information on the collection of Fritz Gra- 
bonsky. William Shear (Hampden-Sydney College) kindly sent us material from 
different collections, which he intended to study himself. Käthe Rehbinder (Univer- 
sität Mainz) most skillfully and patiently produced the whole-animal figures. Jürgen 
Gruber (Naturhistorisches Museum Wien) commented on the manuscript. The Ger- 
man Academic Exchange Service (Deutscher Akademischer Austauschdienst, Bonn) 
supported P.J. S. with a 1 -months research grant at the beginning of this project. The 
Feldbausch and Wagner Foundations of the University of Mainz provided travel funds 
toJ.M. 

REFERENCES 

International Commission on Zoological Nomenclature 2001. Opinion 1966. Gnomulus 

Thorell, 1890 (Arachnida, Opiliones): Gnomulus sumatramis Thorell, 1891 designated 

as the type species. Bulletin of Zoological Nomenclature 58 (1): 66-67 '. 
Loman, J. C. C. 1902. Neue aussereuropäische Opilioniden. Zoologische Jahrbücher, Abteilung 

für Systematik 16: 163-216. 
Martens, J. & Schwendinger, P. 1998. A taxonomic revision of the family Oncopodidae I. 

New genera and new species of Gnomulus Thorell (Opiliones, Laniatores). Revue 

Suisse' de Zoologie 105 (3): 499-555. 
Roewer, C. F. 1923. Die Weberknechte der Erde. Fischer, Jena. 

Roewer, C. F. 1935. Südostasiatische Opiliones der Sammlung Fea und Modigliani des Natur- 
historischen Museum in Genua. Annali des Museo civico di Storia Naturale di Genova 

59: 12-25. 
Schwendinger, P. J. 1992. New Oncopodidae (Opiliones, Laniatores) from Southeast Asia. 

Revue suisse de Zoologie 99 (1): 177-199. 
Schwendinger, P. J. & Martens, J. 1999a. Case 3116. Gnomulus Thorell, 1890 (Arachnida, 

Opiliones): proposed designation of G. sumatramis Thorell, 1891 as the type species. 

Bulletin of Zoological Nomenclature 56 (3): 171-173. 
Schwendinger, P. J. & Martens, J. 1999b. A taxonomic revision of the family Oncopodidae 

II. The aenus Gnomulus Thorell (Opiliones, Laniatores). Revue suisse de Zoologie 106 

(4): 945-982. 
Suzuki, S. 1969. On a collection of opilionids from Southeast-Asia. Journal of Science of the 

Hiroshima University; Series B, Div. 1 (Zoology) 22: 1 1-77. 



FURTHER NEW SPECIES OF GNOMULUS 



113 



INDEX [Names of Gnomulus species treated in this paper] 



armillatus 74 
carinatus 87, 89 
claviger 100, 101,102 
crass ipes 103, 705, 106 
exsudons 83, 85. 86 
hamatus 89.90 
kutan 81, Si 
javanicus 75. 76 
laruticus 55. 56 
latoperculum 96, 97, 98 
leofeae 72, 74 
lomani 77, 78 



marginatus 66, 68, 69 
matabesar 94, 96 
monticala 56, 57, 58 
obscurus 80, 87 
pilosus 59, 60 
rostradoideus 61, 62 
ryssie 71,72 
sinensis 50, 57, 52 
spiniceps 5 1 , 55, 54 
tuberculatus 64, 65, 67 
tumidifrons 92, 95, 94 



Added in proof. We have only recently received further Gnomulus specimens from 
the Philippines, Malaysia and Singapore (all in MHNG; leg. L. Deharveng, S. Huber, 
L. Monod, A. Schulz, P. J. Schwendinger), which will be treated in a later paper. 



Revue suisse de Zoologie 109 (1): 1 15-125; mars 2002 



Taxonomic consideration of the genus Odontobuthus Vachon 
(Scorpiones, Buthidae), with description of a new species l 

Wilson R. LOURENÇO & Adeline PÉZIER 

Laboratoire de Zoologie (Arthropodes), Muséum National d'Histoire Naturelle, 

61 rue de Buffon, F-75005 Paris, France. E-mail: arachne@mnhn.fr 



Taxonomic consideration of the genus Odontobuthus Vachon (Scor- 
piones, Buthidae), with description of a new species. - The two species 
of the genus Odontobuthus Vachon known at present are reassessed and 
considered to be valid. A new species, Odontobuthus bidentatus sp. n. is 
described from Iraq and from the region of Borazdjan, southern Iran. Some 
comments on the geographical distribution of the genus are added and a 
key to its species is provided. 

Key-words: Scorpions - Odontobuthus - Iraq - new species - geographical 
distribution. 

INTRODUCTION 

The genus Odontobuthus was introduced by Vachon in 1950. Its type species, 
Bitthus doriae Thorell, 1876, was originally described from Teheran. Since its 
establishment, Odontobuthus has been thought to consist of only two species, O. 
doriae, distributed in Iran and Iraq, and Odontobuthus odonturus (Pocock, 1897) from 
Kelat Frontier, Sind (before in India, now in Pakistan). The genus is presumably 
distributed throughout India, Pakistan, Iran and Oman (Fet & Lowe, 2000). 

For some time, the validity of these two species has been questioned. After 
describing O. odonturus (under Buthus odonturus) in 1897, Pocock changed his mind 
and, in his Fauna of India (Pocock, 1900), considered it to be only a sub-species of 
B. doriae, B. doriae odonturus {^Odontobuthus doriae odonturus). This opinion was 
retained by others authors, including Takashima (1945). More recently, in their new 
version of the Fauna of India, Tikader and Bastawade (1983) redescribed Pocock's 
holotype under O. doriae odonturus and suggested that the genus might be mono- 
typic. Finally Kovarik (1997) listed both species and justified their validity on the 
basis of the different number of lateral lobes on the metasomal segment V. 

Because of various imprécisions in the diagnosis both of the genus and of the 
species, and also doubts about their precise geographical distribution (see Fet & 
Lowe, 2000), we decided to investigate the large collection of Odontobuthus available 
in the Paris Museum, together with several specimens now deposited in the Geneva 



The study was supported by the Department of cultural affairs, City of Geneva, Switzerland. 
Manuscript accepted 23.08.2001 



116 W. R. LOURENÇO & A. PÉZIER 

Museum. We concluded that the two species in this genus are valid and can be 
justified by more than the single character given by Kovarik (1997). Furthermore, 
some specimens from Iraq and from the south of Iran were found to represent a new 
congeneric species, which is described below. The geographical distribution of O. 
dorioe is probably limited to Iran, whereas the presence of O. odonturus in Iran still 
requires confirmation. The records by Vachon (1966), Habibi (1971), and Farzanpay 
(1988) are most certainly based on misidentifications. Revised diagnoses are given for 
the genus and for the two known species, followed by a description of the new 
species. A key is also provided for the three species. 

Abbreviations: MHNG. Muséum d'histoire naturelle, Genève. MNHN. Mu- 
séum National d'Histoire Naturelle, Paris. NHMW. Naturhistorisches Museum, Wien. 

TAXONOMIC TREATMENT 

Odontobuthus Vachon, 1950 

Type species by original designation: Buthus doriae Thorell. 

Diagnosis: Scorpions of small to medium size, ranging from 40 to 70 mm in 
total length. General coloration yellow to pale yellow. Dentate margins of fixed and 
movable fingers with 10 to 14 oblique rows of granules separated by stronger 
accessory granules. Carapace carinae strong, showing a lyre-shaped configuration. 
Tergites tricarinated. Ventral carinae of metasomal segments II-III and ventral trans- 
verse carina of segment IV armed with very strong teeth. Ventrolateral carinae of 
metasomal segment V with several strong lobated granules. Anal arch composed of 
strong lateral lobes and more reduced ventral lobes. Trichobothriotaxy type A-ß. 

Odontobuthus doriae (Thorell) Figs 1-8 

Diagnosis: Scorpions of medium size, ranging from 65 to 70 mm in total 
length. General coloration yellow to pale yellow. Dentate margins of fixed and 
movable fingers with 13-14 oblique rows of granules. Ventral carinae of metasomal 
segments II-III armed with 3 pairs of very strong conical teeth. Ventrolateral carinae 
of metasomal segment V with 4-5 strong lobated granules. Anal arch composed of 2 
strong lateral lobes and 6 more reduced ventral lobes. 

Variation in pectinal teeth counts. Males: 27-35. Females: 18-24. 

Material examined: IRAN, Borudjerd.l male (T. Habibi) MNHN RS 4667; Cazvin, 1 
male. 1 female. 1971 (T. Habibi) MHNG; Karady, 1 male, 1 female, 1971 (T. Habibi) MHNG; 
Kashan, 1 female (T. Habibi) MNHN RS 4669, 2 males, 4 females, MNHN RS 1824, 1 female, 
MNHN RS 1823, 4 males, 13 females (A. Chabaud) MNHN RS 1828, 2 males, 22 females (A. 
Chabaud) MNHN RS 1827. 3 males. 2 females (A. Chabaud) MNHN RS 1826, 5 males, 12 
females (A. Chabaud) MNHN RS 4645, 6 males. 16 females (A. Chabaud) MNHN RS 4646; 
Region of Teheran, 4 males, 5 females, 1974 (Farzanpay) MHNG; Rezayeh, 1 female (T. 
Fatemi) MNHN RS 4341; Teheran, 2 males, 2 females (T. Habibi) MNHG; 1 female (T. 
Fatemi) MNHN RS 4403: 20 km NW of Kashan toward Quom, 1 female. 1974 (J. Delacour) 
MNHN RS 7884; 132 km NW of Rafssanjam, 1 14 km SE of Yazd, 1 female (J. Garzoni, A. de 
Chambrier); 100 km SE of Esfahan, after Shiraz, 1 male, 1974 (J. Delacour) MNHN RS 7883; 
70 km W of Sirjan toward Chiraz. 1 female, 1974 (J. Delacour) MNHN RS 7881; 100 km NW 
of Sirjan toward Kerman, 1 male. 3 females, 1974 (J. Delacour) RS 7880; 130 km W of Sirjan 



THE GENUS ODONTOBUTHUS 



17 




Figs 1-8 
Odontobuthus doriae (female). 1-4. Trichobothrial pattern. 1. Chela, external aspect. 2. Femur, 
dorsal aspect. 3-4. Tibia, external and dorsal aspects. 5-6. Metasomal segments II and III, 
lateral aspect. 7. Chelicera, dorsal aspect. 8. Metasomal segment V and telson, lateral aspect, 
showing anal arch. 



118 W. R. LOURENÇO & A. PÉZIER 



toward Chiraz, 1 female, 1974 (J. Delacour) RS 7882; 1 male (Loeffler) NHMW; 1 male 
MNHN RS 1821; 3 males, 3 females MNHN RS 1819; 1 female MNHN RS 1818; 1 male, 
1 female (Doria) MNHN RS 1825. 

Odontobuthus odonturus (Pocock) Figs 9-12 

Diagnosis: Scorpions of small to medium size, ranging from 40 to 50 mm in 
total length. General coloration yellow; carapace carinae covered with dark pigment. 
Dentate margins of fixed and movable fingers with 10-11 oblique rows of granules. 
Ventral carinae of metasomal segments II-III armed with 3 pairs of very strong 
flattened spinoid teeth. Ventrolateral carinae of metasomal segment V with 3 strong 
lobated granules. Anal arch composed of 3 moderate lateral lobes and 4 more reduced 
ventral lobes. 

Variation in pectinal teeth counts. Males: 26-29. Females: 16-21. 

Material examined: PAKISTAN, Indus Delta, 2 males, 7 females, MHNG; Karachi 
1 male, 1 female, MNHN RS 1822; Karachi Rehri village, 7 females, 1965 (J. A. Anderson), 
MHNG; Korangi-Colony, 1 male MHNG. 

Odontobuthus bidentatus sp. n. Figs 13-26 

Type material. Holotype (male): IRAQ, 180 km north of Bagdad, Khanagin-Dyala, 
11/1964 (G. Pringle). Paratypes (1 female): IRAQ. 40 km SW of Mossoul, 10/XI/1981 (B. 
Moutis). (1 male, 1 female): IRAN, region of Borazdjan, HI/1971 (T. Habibi). All specimens 
deposited in the Natural History Museum, Geneva. 

Etymology: The specific name refers to the presence of 2 pairs of teeth on the 
ventral carinae of metasomal segments II-III. 

Diagnosis: Scorpions of medium size, ranging from 60 to 65 mm in total 
length. General coloration yellow; carapace with a dark triangular spot on the anterior 
edge. Dentate margins of fixed and movable fingers with 13-14 oblique rows of 
granules. Ventral carinae of metasomal segments II-III armed with 2 pairs of very 
strong conical teeth. Ventrolateral carinae of metasomal segment V with 2-3 strong 
lobated granules. Anal arch composed of 3 strong lateral lobes and 4 reduced ventral 
lobes. 

Description (based on holotype and paratypes): 

Coloration. Generally yellow with only a few dark spots or pigmented zones 
on the body. Prosoma: carapace with a triangular dark spot in the anterior region; 
zone between anterior median carinae not pigmented; eyes surrounded by black 
pigment. Mesosoma: tergites I- VII with slightly pigmented confluent zones. Meta- 
soma: all segments including vesicle yellowish; aculeus yellowish at the base and 
reddish at its extremity. Venter; pectines pale-yellow. Chelicerae yellowish; teeth 
dark reddish. Pedipalps: some carinae and rows of granules on the dentate margins of 
the fingers reddish. 

Morphology. Prosoma: Anterior margin of carapace only slightly emarginate. 
Carapace carinae moderately to strongly developed; anterior median carinae strong; 
central lateral and posterior median carinae showing lyre configuration; central 
median carinae moderately developed. All furrows moderately developed to strong. 
Intercarinal spaces moderately to weakly granular. Median ocular tubercle anterior to 



THE GENUS ODONTOBUTHUS 



119 






10 



12 



Figs 9-12 
Odontobuihus odonturus (male). 9. Metasomal segment V and telson, lateral aspect, showing 
anal arch. 10. Chelicera, dorsal aspect. 11-12. Metasomal segments II and III, lateral aspect. 



the center of the carapace; median eyes separated by almost one and half ocular 
diameters. Five pairs of lateral eyes; the first four arranged in one line, the fifth pair 
situated anteriorly, just next to the second. Mesosoma: Tergites I- VI tricarinate; all 
carinae strong. Tergite VII pentacarinate, with all carinae strong. Intercarinal spaces 
moderately granular. Sternites: two carinae present on sternites III- VI, moderately 
developed to weak; VII with four carinae, strong. Pectines long; pectinal teeth count 
32-31 (32-32 in male paratype; 25-26, 27-27 in female paratypes). Metasoma: 
Segments I-II with 10 carinae; ventral carinae of segment I with a few stronger gra- 
nules; on segment II, two pairs of very strong conical granules. Segments III-IV with 
8 carinae; ventral carinae of segment III with 2 pairs of very strong conical granules. 
Segment V with 7 carinae; the ventrolateral one armed with 2-3 strong lobated gra- 
nules. Dorsal furrows of all segments weakly developed, smooth; intercarinal spaces 
very weakly granular, almost smooth. Telson smooth. Aculeus moderately long; 
subaculear tubercle absent. Chelicerae with 2 very much reduced basal denticles on 
the movable finger (see Vachon, 1963). Pedipalps: Trichobothrial pattern ortho- 
bothriotaxic, type A (according to Vachon, 1974); dorsal trichobothria of femur in 
beta configuration (see Vachon, 1975). Femur pentacarinate; all carinae moderately 
crenulate. Tibia with 7 carinae, moderately crenulate; anterior margin with 4 spinoid 
granules. Chelae with vestigial carinae only. Dentate margins on fixed and movable 
fingers composed of 13-14 oblique rows of granules, separated by stronger accessory 
granules. Legs: Ventral aspect of tarsi with numerous thin long setae. Strong tibial 
spurs present on legs III-IV. Pedal spurs present, moderately developed to strong on 
all legs. 



120 



W. R. LOURENÇO & A. PÉZIER 





Figs 13-14 
Odontobuthus bidentatus sp. n., male paratype, dorsal and ventral aspects (photos by CI. 
Ratton, MHNG). 



THE GENUS ODONTOBUTHUS 



121 





Figs 15-16 
Odontobuthus bidentatus sp. n., female paratype, dorsal and ventral aspects (photos by CI. 
Ratton, MHNG). 



122 w - R - LOURENÇO & A. PÉZIER 

Morphometric values (in mm): male holotype (in parentheses female para- 
type). Total length 65.7 (61.2); carapace length 9.0 (8.0), anterior width 5.8 (5.2), 
posterior width, 10.4 (9.4); metasomal segment I length 5.2 (4.6), width 6.0 (5.2); 
metasomal segment V length 8.6 (7.1), width 4.2 (3.7), depth 3.6 (3.0); vesicle width 
4.0 (3.6), depth 4.1 (3.6); pedipalp: femur length 7.8 (6.2), femur width 2.2 (2.0), tibia 
length 8.8 (7.2), tibia width 3.2 (2.7), chela length 16.4 (13.2), chela width 4.3 (2.5), 
chela depth 4.8 (3.2), movable finger length 12.0 (9.6). 

Key to the species of Odontobuthus 

1 Anal arch with 2 strong lateral lobes and 6 reduced ventral lobes . . . O. doriae 

(1) Anal arch with 3 strong or moderately developed lateral lobes and 4 
reduced ventral lobes 2 

2 Dentate margins of fixed and movable fingers with 10-11 rows of 
granules; ventral carinae of metasomal segments II-III with 3 pairs of 
spinoid teeth O. odonturus 

(2) Dentate margins of fixed and movable fingers with 13-14 rows of gra- 
nules; ventral carinae of metasomal segments II-III with 2 pairs of spi- 
noid teeth O. bidentatus sp. n. 

Some comments on the geographical distribution of the species 

From our study we have concluded that Odontobuthus doriae has most pro- 
bably only a limited distribution in the central and northwestern regions of Iran (Fig. 
27). Its presence in Iraq still requires confirmation. It may be that the records of O. 
doriae from Iraq is due to misidentification of specimens which actually belong to the 
new species described in here. The distribution of Odontobuthus odonturus has only 
been confirmed for Pakistan and India. Its presence in the southern regions of Iran 
also requires confirmation. We assume that the records by Habibi (1971) and 
Farzanpay (1988) also result from misidentification of the new species, which is 
present in the southwest of Iran. This apparently disrupted distribution pattern of the 
new species does not stand alone. The localities recorded for O. bidentatus sp. n., 
show a similar pattern of distribution as observed for the buthid Compsobuthus 
matthiesseni Binila by Sissom and Fet (1998). This species is widely distributed 
within the drainage systems of the Tigris and Euphrates Rivers. Such a pattern of 
distribution applies to sites where the altitude ranges from 150 to 200 m. In contrast, 
at the sites where O. doriae was collected the altitude is much greater, ranging from 
1000 to 3000 m. 

The record of O. odonturus for Oman (Fet & Lowe, 2000) requires further 
investigation. We have examined a few poorly preserved specimens, labeled O. 
doriae, from the MNHN collections, which are presumably collected in Oman by M. 
Maindron. These specimens originate from outside the known distribution of either 
O. doriae or O. odonturus. We therefore prefer to postpone a decision until more 
material from Oman becomes available. 



THE GENUS ODONTOBUTHUS 



123 




Figs 17-26 
Odontobuthus bidentatus sp. n. (male holotype). 17. Movable finger, cutting edge. 18. Meta- 
somal segment V, lateral aspect, showing anal arch. 19-20. Metasomal segments II and III. 
lateral aspect. 21. Telson, lateral aspect. 22. Chelicera, dorsal aspect. 23-26. Trichobothrial 
pattern. 23. Chela, external aspect. 24. Femur, dorsal aspect. 25-26. Tibia, external and dorsal 
aspects. 



124 



W. R. LOURENÇO & A. PÉZIER 




# Odontobuthus doriae 

▼ Odontobuthus odonturus 

O Odontobuthus bidentatus sp. n. 




OMAN 



OMAN SEA 



Fig. 27 
Map showing the localities of the Odontobuthus material examined. 



ACKNOWLEDGEMENTS 

We are grateful to Didier Geffard (Laboratoire de Zoologie, Arthropodes, 
MNHN) for technical assistance and to Prof. John L. Cloudsley-Thompson (London) 
and Peter Schwendinger (MHNG) for reviewing the manuscript. 



REFERENCES 

Farzanpay, R. 1988. A catalogue of the scorpions occuring in Iran, up to January 1986. Revue 
Arachnologiqiie 8 (2): 33-44. 

Fet, V. & Lowe, G. 2000. Family Buthidae C. L. Koch, 1837 (pp. 54-286). In: Fet, V., Sissom, 
W. D., Lowe, G. & Braunwalder, M. E. (eds). Catalog of the scorpions of the world 
(1758-1998). The New York Entomological Society. 

Habibi, T. 1971. Liste de scorpions de l'Iran. Bulletin of the Faculty of Science Teheran Uni- 
versity 2 (4): 42-47. 

Kovarik, F. 1997. Results of the Czech Biological Expedition to Iran. Part 2. Arachnida: 
Scorpiones, with description of Iranohuthus krali gen. n. et sp. n. and Hottentotta 
zagrosensis sp. n. (Buthidae). Acta Societatis Zoologicae Bohemoslovenicae 61: 39-52. 

Pocock, R. I. 1897. Descriptions of some new species of scorpions from India. Journal of the 
Bombay Natural History Society 1 1 : 102- 1 1 7. 

Pocock, R. I. 1900. Arachnida. In: Blanford, W. T. (ed.). The Fauna of British India, 
including Ceylon and Burma. Taylor and Francis, London, xii, 279 pp. 

Sissom, W. D. & Fet, V. 1998. Redescription of Compsobuthus matthiesseni (Scorpiones, 
Buthidae) from Southwestern Asia. Journal of Arachnology 26: 1-8. 

Takashima, H. 1945. Scorpions of Eastern Asia. Acta Arachnologica 9 (3-4): 68-106. 



THE GENUS ODONTOBUTHUS 125 

Tikader, B. K. & Bastawade. D. B. 1983. The Fauna of India. Vol. 3. Scorpions (Scor- 
pionida: Arachnida). Zoological Survey of India, Calcutta, 671 pp. 

Vachon. M. 1950. Etude sur les scorpions. Ill (suite). Description des scorpions du Nord de 
l'Afrique. Archives de l'Institut Pasteur d'Algérie 28 (2): 152-216. 

Vachon, M. 1963. De l'utilité, en systématique, d'une nomenclature des dents des chélicères 
chez les Scorpions. Bulletin du Muséum National d'Histoire Naturelle, Paris 2è sér., 35 
(2): 161-166. 

Vachon, M. 1966. Liste des Scorpions connus en Egypte, Arabie, Israël, Liban, Syrie, Jor- 
danie, Turquie, Irak, Iran. Toxicon 4: 209-218. 

Vachon. M. 1974. Etude des caractères utilisés pour classer les familles et les genres de 
Scorpions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles trichobo- 
thriaux et types de trichobothriotaxie chez les Scorpions. Bulletin du Muséum National 
d'Histoire Naturelle, Paris, 3è sér., n° 140, Zool. 104: 857-958. 

Vachon, M. 1975. Sur l'utilisation de la trichobothriotaxie du bras des pédipalpes des Scor- 
pions (Arachnides) dans le classement des genres de la famille des Buthidae Simon. 
Comptes Rendus des Séances de l'Académie de Sciences, Paris, sér. D, 281: 1597-1599. 



Revue suisse de Zoologie 109 (1): 127-141; mars 2002 



Nouvelles additions à la faune de scorpions néotropicaux 
(Arachnida) 1 

Wilson R. LOURENÇO 

Laboratoire de Zoologie (Arthropodes), Muséum National d'Histoire Naturelle, 

61 rue de Buffon. F-75005 Paris, France. E-mail: arachne@mnhn.fr 



New records of scorpions for the Neotropics (Arachnida). - In this paper 
are presented the results of the study of an interesting collection of 
neotropical scorpions now deposited in the Geneva Museum. The 
collection is composed of 7 families, 14 genera and 34 species. Two new 
species, Tityus apiacas sp. n. and Tityus vaissadei sp. n. (Buthidae) are 
described. 

Key-words: Scorpion - Neotropics - new species - Tityus. 

INTRODUCTION 

Dans une publication récente (Lourenço, 1997a), j'ai pu apporter une première 
contribution concernant des scorpions de la région néotropicale déposés au Muséum 
d'histoire naturelle de Genève. 

La faune des Scorpions de la région néotropicale peut être considérée comme 
une des plus étudiées au monde, avec de nombreuses contributions depuis le début du 
19ème siècle, jusqu'à la synthèse globale de Mello-Leitào (1945). 

Dans une perspective plus moderne, plusieurs travaux d'ensemble concernant 
la systématique et la biogéographie ont été réalisés depuis une vingtaine d'années. 
Parmi les plus importants peuvent être cités, Maury (1979), Lourenço (1982a,b, 1983, 
1988; 1991, 1994a, 1995, 1997a,b), Francke & Stockwell (1987). Cependant, compte- 
tenu de la très grande diversité de la faune scorpionique de la région néotropicale 
(Lourenço, 1994b), la découverte de nouvelles espèces, mais aussi la confirmation de 
nouvelles stations pour des espèces déjà connues est chose courante. 

Le présent travail est le résultat de l'étude d'une collection hétérogène de Scor- 
pions néotropicaux déposés désormais au Muséum d'histoire naturelle de Genève. En 
plus de la description de deux espèces nouvelles, quelques considérations sont faites 
sur des espèces rares. Pour le restant des espèces, seule une liste est proposée. 

La présentation du matériel étudié est faite dans l'ordre alphabétique des 
familles. Les pays concernés sont: Grandes Antilles: Hispaniola, Haiti, République 
Dominicaine; Petites Antilles: Martinique, Montserrat, Sainte Lucie, Saint Martin; 



1 Etude subventionnée par le Département municipal des affaires culturelles de la Ville de 
Genève. 
Manuscrit accepté le 21 .06.2001 



128 W. R. LOURENÇO 

Amérique du Sud: Argentine, Bolivie, Brésil, Colombie, Equateur, Guyane française, 
Mexique, Pérou, Trinidad, Venezuela. 

A l'exception de quelques spécimens de Tityus apiacas, le matériel cité dans le 
présent travail est déposé au Muséum d'histoire naturelle de Genève (MHNG). 

BOTHRIURIDAE Simon, 1880 
Bothriurus Peters, 1861 

Bothriurus patagonicus Maury, 1968 

Matériel: Argentine, Prov. Mendoza, Los Lenas (litière), 2/III/1985 (N. Fernandez), 
1 femelle. Répartition: sud de l'Argentine. 

Urophonius Pocock, 1893 

Urophonius iheringii Pocock, 1893 

Matériel: Brésil. Porto Alegre, 22/IX/1986 (E.K. Bastos), 1 mâle. Répartition: sud du 
Brésil et Uruguay. 

BUTHIDAE CL. Koch, 1837 
Ananteris Thorell, 1891 

Ananteris balzani Thorell, 1891 

Matériel: Brésil, District Fédéral. Brasilia, 10/111/1976 (leg. W. Lourenço), 1 femelle ; 
Taguatinga, 2/IX/1976 (W. Lourenço), 1 mâle, 1 femelle. Répartition: nord de l'Argentine, 
Paraguay, centre et nord du Brésil; régions savanicoles. 

Ananteris cussinii Borelli, 1910 

Matériel: Venezuela, Estado Aragua, Caguâ, Ansia Miranda, 11/1975 (M. Gonzalez- 
Sponga), 1 mâle, 1 femelle (topotypes). Répartition: nord du Venezuela. 

Ananteris venezuelensis Gonzalez-Sponga, 1972 

Matériel: Venezuela, Estado Bolivar, entre El Dorado y Santa Elena de Uairén, 
12/1/1978 (M. Gonzalez-Sponga), 1 mâle, 1 femelle (topotypes). Répartition: centre du Vene- 
zuela. 

Caribetityus Lourenço, 1999 

Caribetityus elii (Armas & Marcano Fondeur, 1 992) 

Matériel: République Dominicaine. Prov. La Vega, Lorna de Casabito (1410-1440 m), 
tropical moist forest, 12/V/1998 (D. Huber), 9 mâles, 7 femelles, 16 immatures. Obs: Imma- 
tures nés en laboratoire. Répartition: île d'Hispaniola. 

Isometras Ehrenberg, 1828 

Isometrus maculatus (DeGeer, 1778) 

Matériel: Antilles, île de San Martin, Pic du Paradis humide (350 m), 28/VII/1996 
(M. Breuil). 1 mâle. Obs: Nouvelle station pour l'espèce. Répartition: cosmopolite sur des 
régions tropicales et sub-tropicales. 



SCORPIONS NEOTROPICAUX 129 



Microtityus Kjellesvig-Waering, 1966 

Microtityus Consuelo Armas & Marcano Fondeur, 1987 

Matériel: République Dominicaine, Prov. Borahona, 1,2 km d'Ojeda (rocky trail, bush), 
7/V/1998 (D. Huber), 3 mâles, 6 femelles. Répartition : île d'Hispaniola. 

Rhopalurus Thorell, 1 876 

Rhopalurus abudi Armas & Marcano Fondeur, 1987 

Matériel: République Dominicaine, Prov. Pedernales, 3,9 km N de Manuel Goja (sand 
bush), 8/V/1998 (D. Huber), 1 mâle, & femelle; Solar en ciudad Santo Domingo, 18/VII/1988 
(J.A. Ottenwalder), 1 femelle. Répartition: île d'Hispaniola. 

Rhopalurus princeps (Karsch, 1879) 

Matériel: Haiti, Côte ouest, 22/IV/1984 (D. Rigolage), 2 mâles, 1 femelle, 1 juvénile. 
Répartition: île d'Hispaniola. 

Tityus CL. Koch, 1836 

Tityus apiacas sp. n. Figs 1-10 

Holotype mâle: Brésil, Etat du Mato Grosso, Apiacas (9° 34' S - 57° 23' W), 10- 
15/11/1997 (G. Skuk & V. Xavier). Déposé au Musée de Zoologie, Université de Säo Paulo 
(MZUSP). Paratypes: Même localité que pour l'holotype, 2 mâles, 2 femelles (MZUSP). Brésil, 
Etat de Mato Grosso, Aripuanà, Chapada Dardanelos (10° 10' S - 59° 27' W), 2-13/XI/1996 
(G. Skuk), 2 mâles, 1 femelle (MHNG). 

Etymologie. Le nom de la localité typique (Apiacas) est placé en apposition au 
nom générique. 

Diagnose. Espèce de grande taille (cf. Tableau I), de couleur sombre appar- 
tenant au groupe de ""Tityus asthenes\ La nouvelle espèce est voisine de Tityus 
cambridgei Pocock et de Tityus dinizi Lourenço, mais peut être distinguée de ces 
dernières espèces par: (i) absence des carènes ventrales au quatrième anneau du 
metasoma chez les mâles, (ii) présence de plusieurs papilles arrondies sur la lamelle 
médiane proximale des peignes chez les femelles. Tityus apiacas sp. n., peut être 
également distinguée de T. dinizi par l'absence de dilatation de la lame basilaire 
intermédiaire des peignes chez les mâles. Finalement la répartition géographique des 
trois espèces est nettement distincte (Fig. 19). 

Description fondée sur l'holotype mâle. Coloration générale brun noirâtre. 
Prosoma: Plaque prosomienne brun noirâtre; tubercule oculaire et yeux latéraux 
noirâtres, plus foncés que la carapace. Mesosoma: Tergites I à VII de même colo- 
ration que celle de la plaque prosomienne, avec quelques zones jaunâtres estompées. 
Sternites brunâtres avec quelques zones jaunâtres estompées; le Vème avec un 
triangle jaunâtre. Metasoma: Anneaux caudaux I à V brun noirâtre avec des zones 
jaunâtres diffuses; le Vème plus foncé. Telson plus clair que l'anneau V; aiguillon à 
base jaune-rougeâtre et à extrémité rougeâtre. Peignes jaune-clair; opercule génital, 
sternum, hanches et processus maxillaires d'un brun clair avec quelques zones 
jaunâtres. Pédipalpes brun noirâtre; doigts de la main noirâtres. Pattes brun noirâtre 



130 



W. R. LOURENÇO 




Figs 1-5 
Tityus apiacas sp. n. (Holotype mâle). 1. Carapace, 2. Pédipalpe, vue dorsale avec tricho- 
bothries. 3. Peignes et opercule génital. 4. Metasoma et telson, vue latérale. 5. Sternite V. 



SCORPIONS NEOTROPICAUX 



13] 




Figs 6-10 
Tityus apiacas sp. n. (Paratype femelle). 6. Carapace. 7. Pédipalpe, vue dorsale avec tricho- 
bothries. 8. Peignes et opercule génital. 9. Metasoma et telson, vue latérale. 10. Sternite V. 



132 w - R - LOURENÇO 

avec des taches jaunâtres diffuses. Chélicères jaunâtre foncé avec une trame de taches 
noires sur son ensemble; base des doigts très foncée. 

Morphologie. Prosoma: Front de la plaque prosomienne avec une échancrure 
frontale moyennement marquée. Tubercule oculaire antérieur par rapport au centre de 
la plaque prosomienne; yeux médians séparés par un diamètre oculaire; trois paires 
d'yeux latéraux. Plaque prosomienne moyennement granulée; carènes médianes 
oculaires allant du bord antérieur jusqu'en arrière du tubercule oculaire, bien mar- 
quées; carènes médianes postérieures bien marquées; sillon médian postérieur 
profond; d'autres sillons moyennement profonds. Mesosoma: Tergites I-VII moyen- 
nement granulés; carène axiale présente sur tous les tergites, et plus marquée dans les 
derniers; dans le Vlème couvrant la moitié du tergite; tergite VII avec cinq carènes, 
l'axiale limitée au tiers antérieur; les deux médianes et les deux latérales fusionnées 
dans la région proximale. Sternum triangulaire; peignes avec 20-19 dents; lame 
basilaire intermédiaire non dilatée. Sternites faiblement granulés; stigmates linéaires. 
Metasoma: Anneau I avec 10 carènes; anneaux II et III avec 8 carènes; anneau IV 
avec 6 carènes, les ventrales absentes; anneau V avec 5 carènes; espaces interca- 
rénaux faiblement granulés dans tous les anneaux; anneau V avec la face dorsale lisse; 
vésicule avec quelques granules très réduits; aiguillon long; épine sous-aiguillonaire 
développée et aiguë avec deux granules ventraux. Pédipalpes: Fémur à 5 carènes, tibia 
à 7 carènes, la carène interne-dorsale à granules spiniformes. Pince allongée avec 8 
carènes. Tranchant des doigts mobiles avec 16-16 séries obliques de granules. 
Chélicères avec la dentition caractéristique des Buthidae (Vachon, 1963); les dents 
ventrales du doigt mobile réduites. Trichobothriotaxie du type A-a, orthobothrio- 
taxique (Vachon, 1974, 1975). Pattes: tarses armés de nombreuses soies ventrales. 

Femelle paratype. Coloration générale plus claire que chez le mâle; pattes et 
pédipalpes brun-rougeâtre; triangle jaunâtre sur le sternite V petit et estompé. Corps 
plus trapu que chez le mâle; anneaux du metasoma plus courts avec 10-8-8-8-5 
carènes. Peignes avec la lame intermédiaire basilaire fortement dilatée; lamelle 
médiane proximale avec des nombreuses papilles arrondies. Pédipalpes plus courts et 
plus robustes que chez le mâle; tranchant des doigts mobiles avec 16-16 séries 
obliques de granules. 

Variation du nombre de dents des peignes. Mâles : 18-19, 19-19, 19-20(x2). 
Femelles: 19-19(x3). 

Tityus asthenes Pocock, 1893 

Matériel: Equateur, Napo, Sacha, 10/VI/1994 (W. Lourenço), 1 mâle. Répartition: 
Equateur et côte pacifiques de Colombie et Panama. 

Tityus bahiensis (Perty, 1834) 

Matériel: Brésil, Minas Gérais, Uberlandia, 30/VI/1986 (W. Lourenço), 2 mâles. 
Répartiton: centre et sud-est du Brésil. 

Tityus bastosi Lourenço, 1984 

Matériel: Equateur. Napo. Sacha, ll/VI/1994 (W. Lourenço), 1 femelle + exuvia. 
Répartition: Amazonie occidentale, Brésil, Equateur, Pérou, Colombie. 



SCORPIONS NEOTROPICAUX 133 

Tityus cambridgei Pocock, 1 897 

Matériel: Brésil, Para, Belém, 15/111/1970 (A. Correa), 6 mâles; Belém/Abaetetuba (in 
Palmae - Raphia taedigera), 27/V/1996 (G. Couturier), 1 mâle. Répartition: Amazonie 
orientale, Brésil et Guyane française. 

Tityus columbianus (Thorell, 1876) 

Matériel: Colombie, Iza, 1 1 /IIP 1993 (W Lourenço), 1 femelle, 8 juvéniles (forme 
sexuée); Mosquera, 26/11/1988 (W. Lourenço), 5 femelles; 8/IIP1993 (W. Lourenço), 3 
femelles, 30 juvéniles (forme parthenogénétique), + exuvia. Répartition: cordillère orientale de 
Colombie. 

Tityus fasciolatus Pessôa, 1935 

Matériel: Brésil. D.F., Braslândia, 7/VP1976 (W. Lourenço), 9 mâles, 21 femelles, 
7 immatures. Répartition: région centrale du Brésil. 

Tityus forcipula (Gervais, 1843) 

Matériel: Colombie, Penas Biancas (rainforest, 2100 m), 15/11/1988 (W. Lourenço), 
1 femelle. Répartition: côte pacifique de Colombie et Equateur. 

Tityus fuehrmanni Kraepelin, 1914 Fig. 11 

Matériel: Colombie, Angelopolis, 11/1998 (W. Lourenço), 1 mâle, 1 femelle, 3 juvéniles 
(topotypes). Répartition: cordillère centrale de Colombie. 

Tityus insignis (Pocock, 1889) 

Matériel: Antilles, Sainte Lucie, 2/IX/1997 (W. Lourenço), 1 mâle + Exuvie (topotype). 
Répartition: endémique de Sainte Lucie. 

Tityus melanostictus Pocock, 1 893 

Matériel: Trinidad, Majuba Road, Petit Valley, Diego Martin, 1 /VU/ 1970 (F.H. 
Aitken), 1 femelle. Répartition: Trinidad, nord du Venezuela. 

Tityus metuendus Pocock, 1897 

Matériel: Brésil, Amazonas, Manaus, Reserva Ducke, V/1972 (A. Correa), 1 femelle; 
Acre, Cruzeiro do Sul (primary rainforest), 7/IV/1981 (W. Lourenço), 1 femelle; Pérou, Iquitos, 
Loreto, Jenaro Herrera, V/1996 (G. Couturier), 1 femelle (parthenogénétique), 20 juvéniles 
(mâles). Répartition: Amazonie centrale et occidentale, Brésil et Pérou. 

Tityus pictus Pocock, 1 893 

Matériel: Antilles, St. Vincent (W.I.), Union, 1/1995 (J. Dandin), 1 femelle. Répartition: 
sud des Petites Antilles. 

Tityus serrulatus Lutz & Mello, 1922 

Matériel: Brésil, Minas Gérais, Belo Horizonte, 25/X/1987 (W. Lourenço), 1 femelle. 
Répartition: régions centre et sud-est du Brésil. 

Tityus silvestris Pocock, 1 897 

Matériel: Brésil, Para, Belém, 15/111/1970 (A. Correa), 1 mâle, 3 femelles. Répartition: 
bassin amazonien, Brésil, Pérou, Equateur. 



134 



W. R. LOURENÇO 




Fig. 1 1 
Tityus fuehrmanni, Femelle d'Angelopolis, Colombie, localité typique. 



Tityus vaissadei sp. n. 



Figs 12-18 



Tityus nematochirus: Lourenço, 1997b : 75 

Holotype mâle; 8 mâles et 15 femelles paratypes (MHNG). Colombie, Dept. Boyaca, 
Otanche. 100 km à W. de Tunja, région de Muzo, VI/1986 (Indiens de la Mine de Muzo). 

Etymologie. Le nom spécifique est attribué en hommage à M. Alain Vaissade, 
Maire de la Ville de Genève. 

Diagnose. Espèce de grande taille (cf. Tableau I), de couleur sombre appar- 
tenant au groupe de "Tityus asthenes". La nouvelle espèce peut être distinguée de 
Tityus nematochirus, par la forme des doigts de la pince des pédipalpes bien plus 
encourbés vers l'antérieur, par la présence de granules spiniformes plus fortement 
marqués sur les carènes dorsales des anneaux II-III du metasoma et par quelques 
différences dans la dentition des chélicères. 

La nouvelle espèce peut également être distinguée de Tityus oteroi Lourenço et 
de Tityus antioquensis Lourenço & Otero Patino par les caractères suivants : 



SCORPIONS NEOTROPICAUX 135 



Tableau I. Mensurations (en mm) des espèces décrites (M = mâle, F = femelle) 

Tityus apiacas Tityus vaissadei 

MF M F 

81,0 73,0 



Longueur totale 


97,0 


76,0 


Prosoma 






- Longueur 


8,8 


8,2 


- Largeur antérieure 


6,8 


6,0 


- Largeur postérieure 


10,1 


9,6 


Anneau caudal I 






- Longueur 


7,2 


6,2 


- Largeur 


4,4 


4,2 


Anneau caudal V 






- Longueur 


11,7 


9,2 


- Largeur 


5.1 


4,2 


- Hauteur 


4,9 


4,1 


Vésicule 






- Largeur 


3,8 


3,0 


- Hauteur 


3.5 


2,9 


Pédipalpe 
- Fémur longueur 


13,6 


9,3 


- Fémur largeur 


2,8 


2,4 


- Tibia longueur 


13,8 


9,7 


- Tibia largeur 


3,0 


3,4 


- Pince longueur 


21,8 


16,6 


- Pince largeur 


2,8 


2,8 


- Pince hauteur 


2,5 


2,7 


Doigt mobile 






- Longueur 


13,9 


11,0 



8.8 


7,9 


7,1 


5,9 


10,2 


9,5 


6,8 


5,2 


4,4 


4,6 


11,3 


8,9 


4,7 


4,0 


4,4 


4,0 


3,7 


3,2 


3,6 


3,0 


18,3 


9,8 


2,2 


2,4 


18,5 


10,4 


2,4 


3,3 


29,5 


17.1 


2,0 


2,9 


2,0 


2,8 



18,0 11,6 



De T. oteroi par des pédipalpes bien plus longs et affilés; en outre, les carènes 
dorsales des anneaux I-IV du metasoma chez T. oteroi présentent des épines 
postérieures très développées. 

De T. antioquensis par la coloration, noirâtre chez la nouvelle espèce est 
rougeâtre chez T. antioquensis; en outre, la lame basilaire intermédiaire chez T. antio- 
quensis n'est pratiquement pas dilatée, les carènes dorsales du metasoma ne pré- 
sentent pas des granules spiniformes et les nombres de dents aux peignes sont plus 
faibles allant de 14 à 16. 

Description fondée sur l'holotype mâle. Coloration générale brun-noirâtre. 
Prosoma: Plaque prosomienne brun-noirâtre sans taches; tubercule oculaire et yeux 
latéraux noirâtres. Mesosoma: Tergites I à VII avec la même coloration que celle de la 
plaque prosomienne, mais légèrement plus foncés. Sternites brun-rougeâtre avec une 
grande tache plus claire dans la région postérieure du Vème. Metasoma: Anneaux 
caudaux I à V brun-noirâtre. Telson brun-rougeâtre; aiguillon à base jaune foncé et à 
extrémité rougeâtre. Peignes jaune-clair; opercule génital, sternum, hanches et pro- 
cessus maxillaires d'un brunâtre tacheté. Pattes et pédipalpes brun-rougeâtre; tarses 



136 



W. R. LOURENÇO 




12 




Figs 12-13 
Tityus vaissadei sp. n. Holotype-mâle (12) et paratype femelle (13), vue dorsale. 



SCORPIONS NEOTROPICAUX 



137 



18 




Figs 14-18 
Figs 14-15. Tranchant des doigts mobiles des pédipalpes avec la courbure caractéristique vers 
l'antérieur. 14. Tityus nematochirus (holotype-mâle). 15. Tityus vaissadei (holotype-mâle). Figs 
16-17. Doigts fixe et mobile des chélicères. 16. Tityus nematochirus (holotype-mâle). 17. Tityus 
vaissadei (holotype-mâle). 18. Pédipalpe de Tityus vaissadei (holotype-mâle), vue dorsale avec 
trichobothries. 



138 



W. R. LOURENÇO 



ì^Tityus cambridgei 
Q Tityus dinizi 
£ Tityus apiacas 
A Tityus nematochirus 
W Tityus vaissadei 




Fig. 19 
Carte avec les localités typiques de Tityus apiacas sp. n. et Tityus vaissadei sp. n. et espèces 



associées. 



des pattes jaunâtres. Chélicères brun-rougeâtre foncé avec une trame de taches noires 
sur son ensemble. 

Morphologie. Prosoma: Front de la plaque prosomienne avec une échancrure 
frontale moyennement marquée. Tubercule oculaire antérieur par rapport au centre de 
la plaque prosomienne; yeux médians séparés par un peu plus d'un diamètre oculaire; 
trois paires d'yeux latéraux. Plaque prosomienne faiblement granulée; carènes mé- 
dianes oculaires allant du bord antérieur jusqu'au début du tubercule oculaire; carènes 
médianes postérieures faiblement marquées; sillon interoculaire bien marqué. Meso- 
soma: Tergites moyennement granulés; carène axiale présente sur tous les tergites; 
tergite VII avec cinq carènes, l'axiale limitée au tiers antérieur; les deux médianes et 
les deux latérales fusionnées dans la région proximale. Sternites faiblement granulés, 
presque lisses; stigmates longs et linéaires. Peignes avec 21-21 dents; lame basilaire 
intermédiaire avec une faible dilatation, arrondie. Metasoma: Anneaux I avec 10 
carènes; anneaux II à IV avec 8 carènes; anneau V avec 5 carènes; espaces intercaré- 
naux faiblement granulés; vésicule peu granulée; épine sous-aiguillonaire courte et 
spinoide avec deux granules dorsaux. Pédipalpes: Fémur à 5 carènes, tibia à 7 
carènes, la carène interne-dorsale sans granules spiniformes. Pince avec 9 carènes très 
estompées. Tranchant des doigts mobiles avec 16-16 séries de granules. Chélicères 
avec la dentition caractéristique des Buthidae; le doigt mobile avec 2x2 petites dents 



SCORPIONS NEOTROPICAUX 139 

en position basale (Vachon, 1963). Trichobothriotaxie du type A-a, orthobothrio- 
taxique (Vachon, 1974, 1975). 

Femelle. Coloration plus claire que chez le mâle, brun-rougeâtre; pattes et 
pédipalpes rougeâtre. Corps plus trapu; pédipalpes beaucoup plus courts que chez le 
mâle. Granulation et carènes plus marquées. Lame basilaire intermédiaire dilatée et 
ronde. 

Variation dans le nombre de dents des peignes. Mâles: 19-19, 19-20(x3), 20- 
19, 20-20(x2), 21-21. Femelles: 17-17, 18-18(x2), 19-18, 19-19(x5), 20-18, 20-19, 
20-20(x4). 

Remarque. Dans une note précédente sur les scorpions de Colombie (Lou- 
renço. 1997b), le présent matériel a été identifié comme étant Tityus nematochirus 
Mello-Leitâo, 1940, espèce décrite de Villavicencio. A ce moment les types étaient 
considérés comme perdus. Récemment, j'ai pu localiser au Musée National de Rio de 
Janeiro, Brésil, un exemplaire mâle, appartenant à la série typique. Malgré le très 
mauvais état de conservation de cet exemplaire (conservé à sec puis mis en alcool), 
j'ai pu constater que le matériel collecté à Muzo appartenait à une espèce distincte. 
Déjà dans deux cas précédents, deux autres espèces assez proches de T. nematochirus 
ont été décrites (Lourenço, 1998; Lourenço & Otero Patino, 1998). 

Tityus zulianus Gonzalez-Sponga, 1981 

Matériel: Venezuela, Estado Zulia, Cupure, 7/III/2000 (A. Borges), 1 mâle. Répartition: 
région nord-ouest du Venezuela. 



CHACTIDAE Pocock, 1893 

Broteochactas Pocock, 1893 

Broteochactas brejo Lourenço, 1988 

Matériel: Brésil, Cearâ, Maranguape (Mts.), 11/1948 (Stanford expedition), 1 femelle 
(holotype). Remarque: Depuis sa description, ce spécimen est resté dans les mains de l'auteur, 
sans affectation d'un musée précis, en raison des doutes sur son appartenance. Arrivant à la 
conclusion que cet exemplaire est libre, je le dépose désormais dans la collection du Muséum 
d'histoire naturelle de Genève. Répartition: seule connue de la localité typique. 

Chactopsis Kraepelin, 1912 

Chactopsis anduzei Gonzalez-Sponga, 1982 

Matériel: Venezuela, Terr. Federal Amazonas, Cano Pava, X/1982 (P. Anduze), 
1 femelle (topotype). Répartition: sud du Venezuela. 

Taurepania Gonzalez-Sponga, 1978 

Taurepania vestigialis Gonzalez-Sponga, 1978 

Matériel: Venezuela, Estado Bolivar, Riberas del curso medio del Rio Yuruani, 
12/VI/1975 (A. Lancini), 2 femelles (topotypes). Répartition: sud du Venezuela. 



140 w - R - LOURENÇO 

DIPLOCENTRIDAE Peters, 1861 
Oieclus Simon, 1880 
Oieclus purvesii (Becker, li 



Matériel: Petites Antilles, Montserrat, Parishof St. Peterw woodlands, 16°45'605- 
62°12'956, secondary forest under stone, 27/11/2001 (M. Stevens), 1 immature. Répartition: 
Petites Antilles. 



EUSCORPIIDAE Laurie, 1896 
Euscorpius Thorell, 1876 

Euscorpius flavicaudis (DeGeer, 1778) 

Matériel: Uruguay, Port de Montevideo, X/1997 (C.A. Toscano-Gadea), 1 mâle, 1 
femelle. Obs: Cette espèce typiquement européenne a pu s'introduire et s'acclimater dans la 
zone portuaire de Montevideo (voir Toscano-Gadea, 1998). Répartition: Afrique du Nord, 
Europe du Sud. 

IURIDAE Thorell, 1876 
Hadruroides Pocock, 1 893 

Hadruroides maculatus (Thorell, 1876) 

Matériel: Pérou, Gran Tumbez. IV/1951 (R. Bauer), 2 mâles, 1 femelle, 1 juvénile. 
Répartition: Equateur, Peru. 

VAEJOVIDAE Thorell, 1876 
Syntropis Kraepelin, 1900 

Syntropis macrura Kraepelin, 1900 

Matériel: Mexique, Baja California Sur, N. Los Aripes, 25/VI/1985 (W. Lourenço & G. 
Polis), 2 mâles, 2 femelles (topotypes). Répartition: Mexique, Baja California. 

REMERCIEMENTS 

Je tiens à remercier le Dr V. Mahnert, directeur du Muséum d'histoire natu- 
relle de Genève, de m' avoir facilité la réalisation de la présente étude et Mlle 
Christine Beau, stagiaire au Laboratoire de Zoologie (Arthropodes), M.N.H.N., pour 
la réalisation de plusieurs illustrations. 

REFERENCES BIBLIOGRAPHIQUES 

Francke, O. F. & Stockwell, S.A. 1987. Scorpions from Costa Rica. Special Publications of 
the Museum, Texas Tech University 25: 1-65. 

Lourenço. W. R. 1982a. Révision du genre Ananteris Thorell, 1891 (Scorpiones, Buthidae) et 
description de six espèces nouvelles. Bulletin du Muséum National d'Histoire Natu- 
relle, Paris, 4e sér. 4 (A 1/2): 119-151. 



SCORPIONS NEOTROPICAUX \4] 



Lourenço, W. R. 1982b. Révision du genre Rhopalurus Thorell, 1876 (Scorpiones, Buthidae). 
Revue Arachnologique 4 : 107-141. 

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Revue suisse de Zoologie 109 (1): 143-153; mars 2002 



A re vision of Oreophoetes Renn, 1904, and description of a new 
genus (Insecta: Phasmatodea: Anareolatae: Diapheromeridae: 
Diapheromerinae: Oreophoetini) 

Oliver ZOMPRO 

Max-Planck-Institut für Limnologie, Arbeitsgruppe Tropenökologie, 

August-Thienemannstraße 2, D-24306 Plön, Deutschland. 

E-mail: zompro@mpil-ploen.mpg.de 

Website: www.sungaya.de 

A revision of Oreophoetes Rehn, 1904, and description of a new genus 
(Insecta: Phasmatodea: Anareolatae: Diapheromeridae: Diapherome- 
rinae: Oreophoetini). - The genus Oreophoetes Rehn, 1904, is revised. 
Diagnoses are given for all species included and synonyms are listed. 
Bacteria nigripes Scudder, 1875, is reduced to a subspecies of O. peruana 
(Saussure, 1868). A lectotype is designated for Bacteria peruana Saussure, 
1868. The egg of Oreophoetes mima (Giglio-Tos, 1898) is described for 
the first time. A new genus and species of this tribe from Northern Peru, 
Oreophoetophasma hennemanni gen. n., sp. n., is described. 

Key-words: Phasmatodea - Oreophoetes Rehn, 1904 - Oreophoetophasma 
hennemanni gen. n., sp. n. - synonymy - eggs. 

INTRODUCTION 

Because of several distinguishing characters the genus Oreophoetes Rehn, 
1904, formerly treated as member of the Heteronemiini auct. (see Zompro, 2001c), 
had to be transferred to Diapheromeridae: Diapheromerinae: Oreophoetini by Zompro 
(2001a: 53). This tribe includes some of the most colourful phasmids which are 
distributed in the Andes of NW South America. The species are specialized fern- 
eaters. 

The Oreophoetini are closer related to the Ocnophilini than to the Diaphero- 
merini. As in Ocnophilini, their abdomen is strikingly short, the third antennomere is 
strongly elongated and the meso- and metafemora are rectangular and not trapezoidal 
in cross-section. The Diapheromerini have a comparably longer abdomen, much 
shorter third antennal segment, and meso- and metafemora which are trapezoidal in 
cross-section. 

MATERIAL AND METHODS 

Material was examined in several public and private collections. Livestock was 
obtained from The Phasmid Study Group, England, and reared in cages described by 
Zompro (1996). Preservations were executed as described by Zompro (1996). 



Manuscript accepted 26.09.2001 



144 °- ZOMPRO 

Measurements were taken using an ocular micrometer of Russian make, 
drawings were made using a Zeiss-Citoval 2 binocular with trawing tube. 

Material was examined from the following collections: Academy of Natural 
Sciences, Philadelphia, USA [ANSP]; Deutsches Entomologisches Institut, Ebers- 
walde, Germany [DEIC]; Museum d'histoire naturelle, Geneva, Switzerland 
[MHNG]; Universita di Torino, Torino, Italy [MIZT]; Museo de Historia Natural, 
Universidad Nacional Mayor de San Marcos, Lima, Peru [MUSM]; Naturhistorisches 
Museum, Vienna, Austria [NHMW]; Staatliches Museum für Naturkunde, Stuttgart, 
Germany [SMNS]; Staatliches Museum für Tierkunde, Dresden, Germany [SMTD]; 
Zoologisches Museum der Humboldt-Universität, Berlin, Germany [ZMHB]; Zoo- 
logisches Museum der Universität Hamburg, Germany [ZMUH]; Zoologische Staats- 
sammlung, München, Germany [ZSMC]; Frank H. Hennemann, Freinsheim, Ger- 
many [FHH] and the author's collection [OZ], affiliated with Zoologisches Museum 
der Christian- Albrechts-Universität, Kiel, Germany [ZMUK]. 

OREOPHOETINI 

Diagnosis. Small, colourful, Anareolatae. Profemora, at least in male, straight. 
Abdominal segments II to X combined shorter than thorax with median segment. 
Third antennal segment more than two times as long as scapus and pedicellus 
combined. Egg rounded rhombic in transverse cross-section. 

The Oreophoetini includes only two genera, Oreophoetes Rehn, 1904 and 
Oreophoetophasma gen. n. 

Key to the males of Oreophoetini 

1 Head globose, as wide as long Oreophoetophasma hennemanni 

Head rounded rectangular, distinctly longer than wide Oreophoetes 2 

2 Head uniformly yellow or orange 3 

Head with four black spots Oreophoetes peruana peruana 

3 Head and pronotum uniformly yellow Oreophoetes peruana nigripes 

Head orange, pronotum black Oreophoetes mima 

Key to the known females of Oreophoetini 

1 Head subglobose, vertex distinctly raised .... Oreophoetophasma hennemanni 
Head rounded rectangular, vertex flat Oreophoetes 2 

2 Head with four black spots Oreophoetes peruana 

Head with at best two small black spots between the eyes . Oreophoetes mima 

Key to the known eggs of Oreophoetini 

1 Capsule brownish red with irregular, darker spots Oreophoetes 2 

Capsule uniformly black Oreophoetophasma hennemanni 

2 Micropylar plate reaching operculum Oreophoetes peruana 

Micropylar plate not reaching operculum Oreophoetes mima 



A REVISION OF OREOPHOETES ] 45 



Oreophoetes Rehn, 1904 

Oreophoetes Rehn, 1904: 56; Kirby, 1904: 350; Giglio-Tos, 1910: 31; Hebard, 1924: 145; 

Bradley & Galil, 1977: 180; Sellick, 1997: 102; 1998: 213; Zompro, 2001a: 53. 

Type-species: Bacteria peruana Saussure, 1868, by original designation. 
AUophyllus Brunner von Wattenwyl, 1907: 317. 

Type-species: Bacteria peruana Saussure, 1868, by subsequent designation of Hebard, 

1924: 145. 

Complementary description. Small to medium sized, colourful phasmids. Head 
rounded rectangular, vertex flat, not raised. Eyes small, projecting hemispherically. 
Antennae in both sexes considerably longer than body. Scapus flattened, rounded 
rectangular, pedicellus half as long and two thirds as wide, subcylindrical. Pronotum 
rectangular with curved lateral margins, shorter and narrower than head. Mesonotum 
twice as long as metanotum, of similar width as pronotum. 

Profemora straight. Protibiae longer than profemora, probasitarsus at least as 
long as combined length of remaining segments. Meso- and metafemora rectangular 
in cross-section. 

Median segment shorter than one third of metanotum. Abdominal segments II 
to X combined shorter than thorax and median segment combined. Abdominal 
segment II twice as long as median segment. II to VII of similar width, VII slightly 
shorter or of equal length. VIII of male as long as X, IX longer. Segment VIII slightly 
widened posteriorly, IX with dorsomedian carina, X with a notch posteriorly, IX with 
dorsomedian carina, X with a notch posteriorly. Subgenital plate swollen, with 
prominent margin posteriorly. Vomer simple, triangular. Cerci simple, slightly 
curved. VIII of female indistinctly wider than VII, longer than IX or X. X as long as 
IX. Cerci strong, straight. Subgenital plate boat-like, pointed posteriorly, with 
ventromedian carina. 

Comments concerning Oreophoetes Rehn, 1904. The strongly elongated third 
antennomere is a striking feature which only occurs in the genus Neophasma 
Redtenbacher, 1906. a member of the suborder Areolatae, further it is one of the main 
distinguishing characters of the recently described fossil Archipseudophasmatidae 
from Baltic amber (Zompro, 2001b), which are also members of Areolatae. 

The species of this genus are able to spray from their prothoracal glands; this 
spray burns on the skin and is possibly harmful for the eyes. 

Oreophoetes mima (Giglio-Tos, 1898) 

Bacunculus mimus Giglio-Tos, 1898: 25; Hebard, 1924: 145. 
Oreophoetes mimus: Giglio-Tos, 1910: 32. 
Heteronemia mimus: Kirby, 1904: 348. 

Material examined. 30 6, 16 9, 1 9n5, 1 9n4, 12 eggs. 

Lectotype (designated by Brock, 1998: 302): S, Gualaquiza [MIZT]. - Paralectotypes: 
2 9, Gualaguiza; 4 6, Ecuador. 28.IX.1905 Dr. Fr. Ohaus leg 1905. id. vend. 30.1.1907. cfr. 
Reisebericht 1907; 18 6, 7 9, 1 9n5, Ecuador. Sabanilla b. Zamora, Prov. Loja 16. / 23. / 27. / 
29.IX. / 1. / 2.X. 1905 Dr. Fr. Ohaus leg. id. vend. 30.1.1907. cfr. Reisebericht 1907; 2 S, 1 9, 
Ecuador. Sabanilla b. Zamora, Prov. Loja 29.IX. / 2.X.1905 Dr. Fr. Ohaus leg. id. vend. 
30.1.1907. cfr. Reisebericht 1907 [OZ no. 66-1 -3]; 6 6, 5 9, 4 eggs, Ecuador, Prov. Zamora, 



146 °- ZOMPRO 



Chinchipe, Rio Sabanilla, 27.XI.1998, leg. D. Berger [OZ no. 66-4 -8, 10-15]; 3 9,1 2n4, 8 
eggs, Ecuador, Prov. Zamora, Chinchipe, Estacion Cientifica San Francisco, 1.1999, leg. D. 
Berger [OZ no. 66-9, 16-18]. 

Diagnosis. This species is extremely variable in size, but usually smaller than 
O. peruana. Body shiny. In males vertex uniformly orange, genae sometimes black; 
abdomen dark, thorax red with irregular black spots. Head of females with two small 
black spots between eyes, vertex uniformly yellow. Body yellow with irregular black 
spots or stripes. Femora irregularly annulated, not only with a long dark area in the 
apical half as in some females of O. peruana. 

Measurements (mm), males. Body: 38.4-52.1; head: 2.7-3.3; pronotum: 2.4- 
3.0; mesonotum: 9.4-12.3; metanotum: 4.7-6.0; mediansegment: 1.3-1.7; profemora: 
13.0-17.8; protibiae: 16.5-22.3; mesofemora: 10.1-13.5; mesotibiae: 11.8-15.9; 
metafemora: 13.0-18.1; metatibiae: 17.2-23.2. 

Measurements (mm), females. Body: 41.1-55.0; head: 3.3-4.8; pronotum: 2.4- 
3.2; mesonotum: 9.8-13.8; metanotum: 4.7-5.6; mediansegment: 1.7-2.2; profemora: 
12.0-16.8; protibiae: 13.7-19.2; mesofemora: 9.0-13.0; mesotibiae: 10.0-15.4; 
metafemora: 12.6-18.8; metatibiae: 14.5-22.6. 

Egg. General colour brownish red with irregular darker spots. Capsule slightly 
punctured, rounded rhombic in transverse cross-section, lateral edges flattened. 
Micropylar plate reaching close to the operculum anteriorly, short median line 
present, with broad, laterally parallel margin. Polar area with deep transversal 
impression. Operculum oval, with a group of short, light bristles in its center. 
Measurements (mm). Length: 2.55; width: 1.15; height: 2.35. 

Oreophoetes peruana peruana (Saussure, 1868) 

Bacteria peruana Saussure, 1868: 65; 1870: 160, pi. 3 fig. 12a, b. 

Oreophoetes peruana: Rehn, 1904: 56; Kirby, 1904: 350; Giglio-Tos, 1910: 31; Hebard, 1924: 

145; Mottaz & Tudor, 1989: 15; Potvin. 1998: 28; Sellick, 1998: 213 fig. 22g, h; 

Zompro, 2001a: 53 fig. 68-69, 122-123. 
Allophyllus peruanus: Brunner von Wattenwyl, 1907: 317, pi. 14: 2a, b. 
Bacunculus festae Giglio-Tos, 1898: 22; Hebard, 1924: 145; synonymized by Giglio-Tos, 

1910: 31. Lectotype designated by Brock. 1998: 302. 
Heteronemia festae: Kirby, 1904: 348. 
Bacunculus festuca Giglio-Tos, 1898: 24; Hebard, 1924: 145; synonymized by Giglio-Tos, 

1910: 31. Lectotype designated by Brock, 1998: 303. 
Allophyllus festuca: Brunner von Wattenwyl, 1907: 318. 
Heteronemia festuca: Kirby, 1904: 348. 
Bacunculus (?) gramen Gislio-Tos 1898: 26; Hebard, 1924: 145; synonymized by Brock, 1998: 

303. 
Oreophoetes gramen: Giglio-Tos 1910: 32. 
Heteronemia gramen: Kirby. 1904: 349. 

Material examined. 25 6 , 16 2, 1 2n5, several eggs. 

Lectotype (by present designation): d, Iquitos, Pérou, 600/81, Ane. coll. [MHNG]. 
Paralectotypes: 1 S, Pérou [MHNG]; 1 2n5, lectotype of Bacunculus festuca Giglio-Tos, 
1898: Valle del Santiago [MIZT]; 1 6, holotype of Bacunculus gramen Giglio-Tos, 1898: 
Valle del Santiago [MIZT]; 1 <J, 1 2: Banos, Ecuador [MHNG]; 1 2, no data [MHNG]; 1 S: 



A REVISION OF OREOPHOETES \ 47 

Peru [ZMHB]: 1 6, 3 9: Santa Inez, [Ecuad.], R. Haensch S. [ZMHB]; 1 6, Ecuador, Bos, 
8.X.99, Haensch S. G. [ZMHB]; 1 Ô, Ecuador, A 2500, Haensch S. G. [ZMHB]; 1 o\ Ecuador, 
Bos, 22.X.99, Haensch S. G. [ZMHB]; 2 o\ Ecuador, M. 7.4.99, Haensch S. G. [ZMHB]; 1 9, 
Ecuador, Br. 4.5.99, Haensch S. G. [ZMHB]; 1 9, Napo, [Ecuad.], R. Haensch S. G. [ZMHB]; 
1 S. Peru [DEIC]: 1 6,\ 9: Jumbatis, a. Huallaga, N. Peru, 350m, leg. C. Kulg 1932, ded. 
Nagel 1933, November [DEIC]; 1 o\ 1 9, Chaguta, Peru, mittl. Huallaga, leg. G. Kulg 1935, 
ded. Nagel 1935, März [DEIC]; 5 â, 1 9 , Ecuador, Prov. Napo, Sta. Rosa de Misahualli, 40 m, 
21.3.-26.3.1988, leg. Riede et al. [SMNS]; 2 ó\ 1 9, Santa Inez (Ecuad.), R. Haensch S. 
[ZMUH]; 1 c?. Ecuador, Barancas, 16.XII.1905 [ZMUH]; 1 9 , Ost-Ecuador, Riobamba-Macas 
und flussabwärts. E. Fever leg. [ZMUH]; 5 S, 5 9, several eggs, culture O. Zompro, origin: 
Peru [OZ no. 42-1-11]; 1 <5. Ecuador, Puyo Region. Collectedly Chris Râper. 10-12-1990 cî 
[Coll. P. E. Bragg. Accession-number PEB-461]. 

Diagnosis. Large Oreophoetini. Body less shiny than in O. mima, head with 

four black spots. Thorax and abdomen with red (â), orange or yellow ( 9 ) stripes. 9 

with a broad, black stripe beside yellow median line. Femora of females not annu- 

lated, but sometimes with a wide black area in the anterior half. A male in the 

t 

collection P. E. Bragg (Accession no. PEB-461 ) features a totally black body. 

Measurements (mm), males. Body: 53.2-62.5; head: 2.8-3.7; pronotum: 2.8- 
3.0; mesonotum: 13.8-17.0; metanotum: 9.0-9.7; mediansegment: 1.3-1.6; profemora: 
1.82-22.1; protibiae: 20.0-27.0; mesofemora: 13.5-16.9; mesotibiae: 16.2-22.0; 
metafemora: 18.5-22.2; metatibiae: 22.9-28.6. 

Measurements (mm), females. Body: 54.8-64.0; head: 3.9-5.0; pronotum: 2.7- 
3.3; mesonotum: 13.7-15.2; metanotum: 7.5-8.8; mediansegment: 1.7-2.0; profemora: 
16.5-19.6; protibiae: 20.6-24.0; mesofemora: 13.8-16.1; mesotibiae: 16.0-19.4; 
metafemora: 17.0-21.1; metatibiae: 22.1-26.7. 

Egg. General colour of capsule light brown to reddish brown, with irregular 
darker spots. Surface slightly punctured, in lower magnification leather-like (x 20). 
Capsule rounded rhombic in transverse cross-section, subround from lateral view. 
Micropylar plate projecting, elongated, marginated, reaching operculum anteriorly. 
Short median line present. Operculum elongated oval, with a group of light, short and 
thick bristles in its center. Measurements (mm). Length: 3.15; width: 1.65; height: 
2.55. 

Comments. The type-species of Oreophoetes is one of the most famous 
phasmids. It has been cultured since 1984 in the Phasmid Study Group (no. 84), 
culture stock was collected in Peru, Tarapoto, Valley of Rio Shilcayo and is easy to 
rear on various ferns. The temperuture shold not be higher than room-temperature. 
Because of its striking colours O. peruana is one of the favourite species for photo- 
graphers, and numerous pictures have been published in different kinds of magazines. 

Mottaz & Tudor (1989: 39) and Potvin (1998: 28) present brief descriptions, 
figures and notes on the culture of this species. 

Oreophoetes peruana nigripes (Scudder, 1875) stat. n. 

Bacteria nigripes Scudder, 1875: 278; 1896: 205, 217. 

Material examined. Syntypes: 2 6 , Bad. nigripes Scudd. Type Peruv. Andes [ANSP], 
examined from a colour print by P. D. Brock; 1 6: Chaguta, Peru, mittl. Huallaga, März, leg. 
G. Klug 1935, ded. Nagel 1935 [SMTD]. 



148 °- ZOMPRO 

Diagnosis. The head, pronotum, anterior and posterior part of meso- and 
metanotum and the terminal three abdominal segments are yellow, the rest of the 
body and the extremities are black, the tarsal segments are yellowish-black. 

Measurements (mm), taken from a â from SMTD: Body: 52.0; head: 3.0; 
pronotum: 2.9; mesonotum: 15.8; metanotum: 7.5; mediansegment: 1.1; profemora: 
20.5; protibiae: 28.5; mesofemora: 16.2; mesotibiae: 22.1; metafemora: 21.6; meta- 
tibiae: 32.0. 



Comments. This taxon differs from the variable O. peruana (Saussure, lì 
in the colouration and features slight differences in the form of the genitalia. There- 
fore it is reduced to subspecies level. 

The measurements published by Scudder, 1875: 278 (Body: 63; protibiae: 32; 
mesotibiae: 52; metatibiae: 37) are obviously incorrect, as it is clearly visible in 
colour print of the holotype, that the metatibiae are longer than the mesotibiae and not 
considerably shorter. Perhaps Scudder' s 52 should read 25. In all other respects the 
specimen from SMTD agrees perfectly with Scudder' s material. 

Oreophoetophasma gen. n. 

Diagnosis. Small Oreophoetini. Head globose (a) or subglobose (9), vertex 
raised, eyes projecting more than (6) or hemispherically (9). Antennae projecting 
beyond abdomen. Pronotum narrower than head, subrectangular. Mesonotum elon- 
gate, five times as long as pronotum, but narrower (8) or as wide as it (9). Meta- 
notum half as long as previous segment. Profemora straight (6) or basally slightly 
curved and impressed interiorly ( 9 ), subquadrate in cross-section, in 9 edges dis- 
tinctly produced. Meso- and metafemora rectangular in cross-section, in ? edges also 
produced. Tibiae quadrate in cross-section, longer than femora, basitarsi as long as 
following segments combined, these increasingly shorter, terminal tarsite as long as 
second. Median segment one third as long as metanotum, segments II to V in- 
creasingly longer and ( ? ) wider, V to VII increasingly shorter and ( 9 ) narrower. In 
â VIII shortest segment, IX longer, X shorter than IX but longer than VIII, vomer 
simple triangular, poculum bulgy, cerei simple, slightly curved. In 9 VIII to X 
increasingly shorter, cerei slightly flattened, subgenital plate swollen, not projecting 
terminal segment. 

Name. "Oreophoetophasma" mirrors the close relation to Oreophoetes Rehn, 
1904. The name is of neutral gender. 

Type-species, by present designation: Oreophoetophasma hennemanni sp. n. 

Oreophoetophasma hennemanni sp. n. 

Material. Holotype: o\ N-Peru, Rodriguez de Mendoza, 12.8.1994-10.10.1996 
[MUSM, ex coll. FHH no. 0318-4]. Paratypes: 1 <?, data as holotype [MHNG, ex coll. FHH no. 
5]; 1 S, 1 9, 2 eggs ex abdomen, data as holotype [FHH no. 1, 6]; 1 o\ 1 9, data as holotype, 
ex coll. FHH 0318-2, 3 [OZ no. 392-1, 2]; 1 9. Oxapampa XII. 1931 H. Knipper, Peru, 1800m. 
Ü. M. G. Schreiber [ZMHB, drawer 88/1]. 



A REVISION OF OREOPHOETES 



149 




juez de I 
L9Ö4-10jO^ 1996! 




1.2.6. 1994-1 0. 1 0. 1 996 



Figs 1-4 
Oreophoetes peruana peruana (Saussure, 1868). I. Ô, lectotype [MHNG]; 2. 9 [OZ no. 42-5]; 
Oreophoetophasma hennemanni Zompro sp. n. 3. S, holotype [MUSM]; 4. 9, paratype [OZ 
no. 392-2] 



150 °- ZOMPRO 

Description. Male: Head globose, yellow, with a black, quadrate spot on the 
occiput. Frons, labrum, mandibles and appendices of the head black, clypeus brown- 
ish red. Eyes projecting slightly more than hemispherically. Scapus flattened rectan- 
gular, pedicellus cylindrical, two thirds as wide and half as long, third antennite twice 
as long as scapus and pedicellus combined; following segments strongly elongated. 
Antennae consisting of about 33 segments, projecting beyond the tip of abdomen by 
the length of the mesonotum. 

Pronotum black, longer than wide, narrower but longer than head, with slightly 
curved lateral margin, deep transverse impression medially and an incomplete median 
line. Mesonotum reddish brown, strongly elongated, more than five times as long as 
pronotum, except for the anterior and posterior end distinctly narrower than pro- 
notum. Metanotum less than half as long as mesonotum, as wide as pronotum. 

Profemora straight, strongly elongated, rectangular in cross-section, red, base 
and tip darkened. Protibiae considerably longer, quadrate in cross-section, darker in 
colour. Probasitarsus as long as remaining segments combined, following ones 
increasingly shorter, terminal tarsite as long as second tarsite. Meso- and metafemora 
rectangular in cross-section, knees darkened. Tibiae and tarsi as in foreleg. 

Median segment and following abdominal segments coloured as meso- and 
metanotum, one quarter as long as metanotum and half as long as abdominal segment 
II. Segment II to V increasingly longer, V to VII increasingly shorter. Segments II to 
VII of equal width. VIII trapezoidal broadened posteriorly, with distinct median ridge. 
IX yellow, tectiform, with deep lateral impression which separates the broad lateral 
margins. X with distinct median ridge, posterior margin divided in two projecting 
lobes, these with distinct, black teeth interiorly. Abdomen black ventrally. Poculum 
bulbous, with wrinkled posterior margin. 

Vomer simple triangularly, yellow. Cerci black, slightly curved, bristled. 

Female: General colour green. Head subglobose, slightly longer than wide, 
vertex distinctly raised. Head and appendices green, clypeus light reddish. Eyes pro- 
jecting hemispherically. Antennae coloured as head, broken in the specimens exa- 
mined. Scapus rectangular, flattened, pedicellus two thirds as wide and half as long. 
Third antennite less than twice as long as previous two segments, following segments 
elongated. 

Pronotum subrectangular, with curved lateral margin and distinct longitudinal 
and transverse median impressions. Mesonotum five times as long as pronotum. 
Median ridge distinct, with another, undulated, sometimes interrupted ridge subme- 
dianly, and a broad bulge sublaterally. Metanotum half as long as mesonotum, with 
similar structure. 

Legs coloured as body. Profemora subquadrate in cross-section, edges pro- 
duced. Base slightly curved and distinctly depressed interiorly. Tibiae and tarsi as in 
male, tarsal segments yellow, claws black. Meso- and metafemora rectangular in 
cross-section, edges also distinct, tibiae and tarsi as in foreleg. 

Median segment one third as long as metanotum. with median keel and a 
converging carinae submedianly: Segments II to V increasingly longer and broader, V 
to VII shorter and narrower. Ill to VIII with two converging carinae submedially. VIII 



A REVISION OF OREOPHOETES 



151 












11 




Figs 5- 17 
Oreophoetes peruana peruana (Saussure, 1868). Terminal abdominal segments, 5. 6, 6. 9; 
Oreophoete s peruana nigripes (Scudder, 1895). Terminal abdominal segments, 7. 6; Oreo- 
phoetes mima (Giglio-Tos, 1898). Terminal abdominal segments, 8. 6,9. 9; Oreophoe- 
tophasma hennemanni Zompro sp. n. Terminal abdominal segments, 10. 6, 11, 9; Oreo- 
phoetes peruana peruana (Saussure, 1868), egg. 12. Lateral, 13. Dorsal; Oreophoetes mima 
(Giglio-Tos, 1898), egg. 14. Lateral, 15. Dorsal; Oreophoetophasma hennemanni Zompro sp. 
n., egg. 16. Lateral, 17. Dorsal. - Scale 5-1 1: 10 mm; scale 12-17: 1 mm. 



152 O- ZOMPRO 

tectiforme, as long as VI, IX shorter than VIII, as long as VII, with one converging 
carina submedianly. X shorter than IX, with sharp median carina. Subgenital plate 
yellow, acute posteriorly, anterior part depressed laterally, posterior half with distinct 
median keel. Cerci simple, slightly flattened, bristled. 

Measurements (mm), holotype a: Body: 37.0; head: 2.0; pronotum: 1.9; meso- 
notum: 10.6; metanotum: 4.8; mediansegment: 1.1; profemora: 14.3; protibiae: 17.3; 
mesofemora: 11.0; mesotibiae: 12.3; metafemora: 14.6; metatibiae: 18.2. 

Measurements (mm), Si: Body: 37.0-37.2; head: 1.8-2.0; pronotum: 1.8-2.0; 
mesonotum: 10.5-10.8; metanotum: 4.8-5.0; mediansegment: 0.9-1.1; profemora: 
13.7-14.8; protibiae: 17.0-17.5; mesofemora: 10.6-1 1.0; mesotibiae: 1 1.7-12.3; meta- 
femora: 14.3-14.8; metatibiae: 17.8-18.2. 

Measurements (mm), ? 9: Body: 40.1-40.3; head: 2.9-3.1; pronotum: 1.9-2.0; 
mesonotum: 9.2-10.0; metanotum: 4.8-5.0; mediansegment: 1.6-1.8; profemora: 12.7- 
12.9; protibiae: 13.0-14.2; mesofemora: 9.2-9.4; mesotibiae: 9.8-10.0; metafemora: 
12.0-12.2; metatibiae: 15.2-15.8. 

Egg: General colour uniformly black. Capsule strongly depressed laterally, 
braodest in middle, equally narrowed laterally, with leather-like structure. Micropylar 
plate strongly elongated, reaching operculum anteriorly. Operculum slightly curved, 
oval. The description is based on two almost fully developed eggs out of a female's 
abdomen. Measurements (mm). Length: 2.75; width: 1.15; height: 2.25. 

Name: This beautiful species is dedicated to Mr Frank Hennemann, Freins- 
heim, Germany, to acknowledge several years of good cooperation. 

ACKNOWLEDGEMENTS 

The author wants to thank the authors of the mentioned museums for their 
support. Mr Frank Hennemann (Freinsheim, Germany) and Mr Dirk Berger (Berlin, 
Germany) supplied specimens, Mr Paul Brock (Slough, England) sent a picture of a 
type of Bacteria nigripes Scudder. Dr E. Bragg has to be acknowledged for helpful 
discussions on the manuscript. Mrs Anke Teschke made linguistical annotations to the 
manuscript and rendered hospitality during the author's stay in Berlin. Prof. Dr 
Joachim Adis and Prof. Dr Wolfgang Junk have to be acknowledged for their support 
during the works on this paper. 

REFERENCES 

Bradley, J. C. & Galil, B. S. 1977. The taxonomic arrangement of the Phasmatodea with keys 

to the subfamilies and tribes. Proceedings of the Entomological Society, Washington 

79(2): 176-208. 
Brock, P. D. 1998. Catalogue of stick-insect (Insecta: Phasmida) type material in the Museo 

Regionale di Scienze Naturali, Torino. Bollettino del Museo Regionale di Scienze 

Naturali, Torino 15(2): 299-310, pi. 1-2. 
Brunner von Wattenwyl, C. 1907. Die Insektenfamilie der Phasmiden. II. Phasmidae 

Anareolatae (Clitumnini. Lonchodini, Bacunculini). W. Engelmann, Leipzig, pp. 181- 

340, pis. 7-15. 



A REVISION OF OREOPHOETES 153 



GiGLio-Tos, E. 1898. Viaggio del Dr. Enrico Festa nella Republica dell'Ecuador et regioni 
vicine. VI. Ortotteri. Bollettino dei Musei di Zooioga ed Anatomia comparata della 
Royale Università di Torino 13(31 1): 1-108. 

GiGLio-Tos, E. 1910. Fasmidi esotici del R. Museo zoologico di Torino e del Museo civico di 
Storia naturale di Genova. Bollettino dei Musei di Zooioga ed Anatomia comparata 
della Royale Università di Torino 25(625): 1-57. 

Heb ARD, M. 1924. Studies in Dermaptera and Orthoptera of Equador. Proceedings of the 
Academy of Natural Sciences, Philadelphia 76: 109-248, pi. 5-10. 

Kirby, W. F. 1904. A synonymic catalogue of Orthoptera. 1. Orthoptera Euplexoptera, Cur- 
soria et Gressoria. (Forficulidae, Hemimeridae, Blattidae, Mantidae, Phasmidae). Bri- 
tish Museum, London. 501 pp. 

Mottaz, D. & Tudor, D. 1989. PSG No. 84: Oreophoetes peruanas (Saussure). The Phasmid 
Study Group Newsletter 39: 15-17. 

Potvin, W. 1998. Soortbeschrijving van Oreophoetes peruana (Saussure). Phasma 8(32): 28- 
32. 

Redtenbacher, J. 1906. Die Insektenfamilie der Phasmiden. I. Phasmidae Areolatae. Leipzig. 
1-180. pis. 1-6. 

Rehn, J. A. G. 1904. Studies in the Orthopterous family Phasmidae. Proceedings of the 
Academy of Natural Sciences, Philadelphia 56: 38-107. 

Saussure, H. de 1868. Phasmidarum novarum species nonullae. Revue et Magazine de Zoo- 
logie (2)20: 63-70. 

Scudder, S. H. 1875. Notes on Orthoptera from Northern Peru, collected by Professor James 
Orton. Proceedings of the Boston Society of Natural History 17: 257-282. 

Scudder, S. H. 1896. List of exotic Orthoptera described by S. H. Scudder, 1868-1879, with a 
revision of their nomenclature. Proceedings of the Boston Society of Natural History 
27:201-218. 

Sellick, J. 1997. The range of egg capsule morphology within the Phasmatodea and its rele- 
vance to the taxonomy of the order. Italian Journal of Zoology 64: 97-104. 

Sellick, J. T. C. 1998. The micropylar plate of the eggs of Phasmida, with a survey of the 
range of plate form within the order. Systematic Entomology 23: 203-228. 

Zompro. O. 1996. Zum Sammeln, Transport, Konservieren und Züchten von Phasmiden. 
Entomologische Zeitschrift 106(5): 194-202. 

Zompro, O. 2001a. A generic revision of the insect order Phasmatodea: The New World genera 
of the stick insect subfamily Diapheromeridae: Diapheromerinae = Heteronemiidae: 
Heteronemiinae sensu Bradley & Galil, 1977. Revue suisse de Zoologie 108(1): 189- 
255. 

Zompro, O. 2001b. The Phasmatodea and Raptophasma n. gen., Orthoptera incertae sedis, in 
Baltic Amber (Insecta: Orthoptera). Mitteilungen des Geologisch-Pläontologischen 
Instituts der Universität Hamburg 85: 229-262. 

Zompro, O. 2001c. Critical notes on Heteronemia Gray, 1835 and Heteronemiidae (Insecta: 
Phasmatodea). Studies on Neotropical Fauna and Environment 36(3): 221-225. 



Revue suisse de Zoologie 109 (1): 155-165; mars 2002 



Bryconamericus uporas sp. n. (Characif ormes, Characidae), a new 
species from the rio Uruguay basin, in Argentina 

Jorge R. CASCIOTTA ; M. de las Mercedes AZPELICUETA 

& Adriana E. ALMIRON 

Division Zoologia Vertebrados, Facultad de Ciencias Naturales y Museo de La Plata, 

Paseo del Bosque, 1900 La Piata, Argentina. 

E-mail: jrcas@ museo. fcnym.unlp.edu. ar 



Bryconamericus uporas sp. n. (Characiformes, Characidae), a new 
species from the rio Uruguay basin, in Argentina. - A new species of the 
genus Bryconamericus is described from rio Uruguay basin in Misiones, 
Argentina. Bryconamericus uporas is distinguished by the following 
combination of characters: low body depth (28.9-32.2 in % of SL); all teeth 
of upper jaw broadest distally; premaxillary teeth of the inner series with 
seven cusps; premaxillary teeth of the outer row with five cusps, and 
maxillary teeth with five or seven cusps. Also, the new species has 18-20 
branched anal-fin rays, large subcircular black humeral spot, wide black 
lateral band, light violet upper half of flank, and lower half silvery. The 
new species was collected in the headwaters of the arroyos Once Vueltas, 
Toro, Fortaleza, and Yabotf-Mini. Those streams were born in the sierras, 
having rocky and sandy bottom, with clear and rapid water. 

Key- words: Characiformes - Characidae - Bryconamericus - Argentina - 
Misiones - Uruguay basin. 

INTRODUCTION 

Seven species of the characiform genus Bryconamericus are known from 
southern South America. This genus includes about 40 species (Malabarba & Kindell, 
1995), living in freshwaters from Central America (Eigenmann, 1927; Géry, 1977) to 
the south of Buenos Aires Province in Argentina (Menni et al, 1988; Casciotta et al, 
1999). 

Most of these species have been known since the beginning of the last century 
(Evermann & Kendall, 1906; Eigenmann et al, 1907; Fowler, 1940). However, in- 
creasing efforts in studies of characiform fishes have resulted in recent discoveries of 
new species (Malabarba & Kindell, 1995; Azpelicueta & Almirón, 2001). The 
purpose of this paper is to describe a new species of the genus from the rio Uruguay 
basin, in Argentina. 



Manuscript accepted 28.09.2001 



156 J- R- CASCIOTTA ET AL. 

MATERIAL AND METHODS 

Measurements are straight line distances taken with calliper. Standard length 
(SL) was measured from tip of snout to hypural joint, head length includes the oper- 
cular flap, caudal peduncle length is taken from last anal-fin ray to hypural joint. 
Specimens were cleared and stained (C&S) for cartilage and bone following Taylor & 
Van Dyke (1985). 

The specimens examined belong to Academy of Natural Sciences of Phila- 
delphia, USA (ANSP), Fundación Miguel Lillo, Tucumân, Argentina (FML), 
Muséum d'histoire naturelle, Genève, Switzerland (MHNG), and Facultad de Cien- 
cias Naturales y Museo, La Plata, Argentina (MLP). 

Comparative material examined. Bryconamericus agna Azpelicueta & Almirón, 2001: 
FML 3700, holotype, 61.5 mm SL, Argentina, Misiones, arroyo Tabay, Parana basin. ANSP 
177871, 4 ex., 50.4-57.3 mm SL, collecting data as holotype. MHNG 2611.46, 4 ex., 54.3-60.0 
mm SL, collecting data as holotype. 

Bryconamericus iheringi (Boulenger, 1887): MLP 9073, 1 10 ex., 39.9-44.3, Argentina, 
Buenos Aires, Sierra de la Ventana. MLP 9103, 15 ex., 34.8-49.2, Argentina, Buenos Aires, 
Berisso, Los Talas (man-made ponds connected to Rib de la Plata). 

Bryconamericus exodon Eigenmann, 1907: MLP 18-IX-80-1, 2 ex., 39.0-43.5 mm SL, 
Argentina, Buenos Aires, Rio de la Plata in Punta Lara. 

Bryconamericus thomasi Fowler, 1940: FML 1969, 94 ex. (5 measured, 2 males and 3 
females), 40.3-55.4 mm SL, Argentina, Salta, rio Piedras. 

Bryconamericus uporas sp. n. (non type): MLP uncat, 4 ex., 26.7-49.4 mm SL, 
Argentina, Misiones, arroyo Toro. MLP uncat, 3 ex., 42.9-45.0 mm SL, Argentina, Misiones, 
Arroyo Yabotf-Minf. 

RESULTS 

Bryconamericus uporas sp. n. Figs 1-8, table 1 

Holotype. MLP 9568, male, 51.5 mm SL, Argentina, Misiones, Municipio Leandro N. 
Alem, arroyo Once Vueltas (27° 38' S - 55° 12' W), Uruguay basin. Coll. J. Casciotta, A. 
Almirón & M. Donato, February-2001. 

Paratypes. MLP9583, 14 ex., 43.4-51.0 mm SL. collected with the holotype. MHNG 
2619.23, 5 ex., 41.0-47.8 mm SL, Argentina, Misiones, arroyo Fortaleza (26° 45' S - 54° 
10' W), coll. J. Casciotta, A. Cione & M. Donato, April-2000. 

Diagnosis. Biyconamericus uporas is distinguished from other species of the 
genus by the following combination of characters: low body (28.9-32.2 % of SL); 
premaxillary and maxillary teeth with distal portion broader than the base; teeth of 
premaxillary inner row heptacuspid and those of outer row pentacuspid; 18-20 
branched anal-fin rays; large subcircular black humeral spot, and wide black lateral 
band. The new species has the upper half of flank light violet and the lower half 
silvery. 

Description. Morphometries of holotype and 14 paratypes are presented in 
table 1. Body moderately elongate (Fig. 1). Greatest body depth approximately at 
dorsal-fin origin. Dorsal profile of body distinctly convex from upper lip to dorsal-fin 
origin, almost straight from dorsal-fin base to caudal peduncle. Ventral profile of 
body sligthly convex from mouth to anal-fin origin, straight from anal-fin origin to 
caudal peduncle. Dorsal and ventral profiles of caudal peduncle concave. Body 
laterally compressed between pectoral and anal fins. 



A NEW BRYCONAMERICUS FROM URUGUAY BASIN 



157 



Table 1 
Morphometries of the holotype and 14 paratypes of Bryconamericus uporas sp. n. Standard 
length is expressed in mm. SD: standard deviation. 





Holotype 


Range 


mean 


SD 


Standard length 


51.5 


43.4-51.5 






Percentage ofSL 










Body depth 


30.7 


28.9-32.2 


30.4 


0.984 


Head length 


27.1 


24.6-27. 1 


25.9 


0.763 


Predorsal length 


46.6 


46.6-50.5 


48.6 


1.261 


Preventral length 


41.3 


41.3-46.4 


43.9 


1.403 


Preanal length 


57.9 


55.0-60.9 


59.0 


1.513 


Dorsal-fin base 


13.2 


12.7-14.1 


13.3 


0.488 


Anal-fin base 


29.9 


25.5-29.9 


27.4 


1.122 


Pelvic-fin length 


13.0 


12.5-15.1 


14.1 


0.754 


Pectoral-fin length 


21.9 


20.2-22.4 


21.4 


0.632 


Caudal peduncle depth 


10.7 


10.4-11.7 


11.0 


0.375 


Caudal peduncle length 


16.9 


14.9-17.7 


16.4 


0.958 


Distance between pectoral 










and pelvic fin origins 


19.4 


19.4-22.5 


21.3 


0.763 


Distance between pelvic 










and anal fin origins 


15.9 


13.7-18.0 


15.9 


0.969 


Percentage of head length 










Interorbital width 


28.6 


28.6-34.2 


30.9 


1.747 


Head depth 


75.0 


75.0-86.3 


80.7 


2.889 


Orbital diameter 


35.7 


35.7-43.0 


39.2 


1.818 


Snout length 


20.0 


18.1-21.2 


19.6 


1.018 


Premaxillary+maxillary length 


33.5 


33.5-41.9 


37.4 


2.340 


Maxillary length 


25.0 


20.8-27.3 


23.9 


2.150 


Percentage of pectoral-pelvic fin origins 










Pectoral length 


113.0 


95.3-113.0 


100.0 


4.242 



Dorsal-fin origin nearer snout tip than base of caudal-fin rays, dorsal-fin origin 
behind vertical through last pelvic-fin ray insertion. Adipose fin present. Tip of pec- 
toral fin reaching or not pelvic-fin origin. Tip of pelvic fin never reaching anal-fin 
origin. 

Dorsal fin with ii,8 rays; posterior margin of dorsal fin straight, second un- 
branched and first branched dorsal-fin rays of same length. Holotype (largest spe- 
cimen) with small and few (about 3) hooks on two first branched dorsal-fin rays. 

Anal fin with iv, 18-20 rays (3 ex.= 18, 7 ex.= 19, 5 ex.= 20), some males with 
few small hooks on first branched rays. Most of specimens with last unbranched and 
first six branched rays forming an anterior lobe. 

Pectoral fin with i, 10-12 rays (2 ex.= 10, 7 ex.= 11, 6 ex.= 12), posterior 
pectoral-fin margin straight. Scattered hooks on dorsal surface. 

Pelvic fin with i,7 rays, with small hooks on ventral surface. 



158 J - R - C ASCIUTTA ETAL. 




Fig. 1 
Bryconamericus iiporas sp. n., holotype, MLP 9568, 51.5 mm SL. 

Caudal fin with one unbranched and 9 branched rays on upper lobe; one 
unbranched and 8 branched rays on lower lobe. Lower caudal lobe scarcely longer 
and more rounded. 

Dorsal profile of head gently convex, concave over supraoccipital. Snout 
rounded, upper jaw distinctly longer than lower jaw. Mouth placed at level of lower 
orbital margin. Maxilla surpassing anterior orbital margin. Maxilla with ascending 
process long, lateral process laminar. Usually 3 teeth, with 5 or 7 cusps; sometimes, a 
fourth posterior small tooth conic or bicuspid (Fig. 2). Premaxilla bearing two series 
of teeth, wider distally, compressed anteroposteriorly, with stronger median cusp 
(Fig. 3). Usually, outer series with 3 aligned teeth, all pentacuspid (1 ex. with 4 teeth, 
2 ex. with 2). Inner series of premaxillary teeth consisting of 4 teeth, with 7 cusps. 
Symphysial tooth narrower; third and fourth teeth broadest (Fig. 4). Dentary bearing 
7-10 teeth, first four anterior teeth large; last ones very small. Symphysial tooth 
broad. Distal area of each tooth compressed anteroposteriorly. Usually 5 cusps in 
large teeth, remaining teeth with 3 cusps (Fig. 5). 

Eyes large. Postero-ventral edge of third infraorbital not in contact, but very 
close, with sensory tube of preopercle. 

Scales cycloid. Lateral series with 37-40 perforated scales (2 ex.= 37, 4 ex., 
including holotype= 38, 7 ex.= 39, 2 ex.= 40). Five scales between dorsal-fin origin 
and lateral line, 4-5 scales between lateral line and anal-fin origin. Fourteen scales 
around caudal peduncle. Eleven to fourteen scales not forming a regular median series 
between supraoccipital process and dorsal-fin origin in most specimens. Nine to 
eleven scales in one row, covering proximal portion of eight to ten first anal-fin rays. 

Coloration upon capture: Upper half of flank light violet; lower half silvery 
(Fig. 6). 

Coloration in alcohol preserved specimens: Ground color pale yellow, with 
upper area of flanks darker; margin of scales with dark chromatophores forming a 
reticular pattern. Lower half of flanks with small isolated chromatophores, some of 
them concentrated over anal-fin and other ones following myosepta. Dorsum of head 
and snout with black chromatophores. Scattered chromatophores on opercular area, 



A NEW BRYCONAMERICUS FROM URUGUAY BASIN 



159 




Figs 2-5 
Bryconamericus uporas sp. n., 44.5 mm SL. 2, lateral view of left maxilla; 3, lateral view of 
left premaxilla; 4, detail of third and fourth teeth of inner premaxillary series, lingual view; 5, 
medial view of left lower jaw. Scale bar: 1 mm. 



cheek, and maxilla. Ventral region of head, and vent whitish. A large subcircular 
black humeral spot, sometimes ventrally elongate, placed behind third or fourth scales 
of longitudinal series. Next 2 or 3 scales lacking chromatophores. A wide dark lateral 
band 2 or 3 scales deep extended on middle flank, connected to a caudal spot. 
Usually, lateral band extending over medial caudal-fin rays. Posterior margin of eye 
with a silvery half-moon shaped spot. Dorsal fin with chromatophores, especially 
concentrated on distal half; dorsal-fin rays with chromatophores on their margins. 
Anal fin with chromatophores, those of distal area very small, larger chromatophores 
placed on basal region. Adipose with small scattered chromatophores. Caudal fin with 



160 



J. R. CASCIOTTA ET AL. 




Fig. 6 
Bryconamericus uporas sp. n., upon capture, arroyo Once Vueltas, Misiones, Argentina. 



chromatophores, specially concentrated on ray margins and close to distal edges. 
Pectoral and pelvic fins hyaline, with small chromatophores on their surfaces. 

Etymology. The specific name uporas is a guarani word meaning an animal- 
shaped ghost of the water, who care streams, ponds, falls, and swamps. 

Distribution. This species is known from headwaters of the arroyos Once 
Vueltas, Toro, Fortaleza, and Yaboti-Minf, rio Uruguay basin, Province of Misiones, 
Argentina (Fig. 7). The depth of the streams was irregular, about 80 cm (average); the 
substrate was formed by sand and stones; the course had small falls and pools, and 
clear water without vegetation (Fig. 8). The temperature of the water was 24-25 °C. 
Many specimens were observed moving upstream over small falls and falling in pools 
during the day, in February. 

DISCUSSION 

The traditional definition of the genus Bryconamericus done by Eigenmann 
(1927) included the species with four teeth in the inner row of the premaxilla, maxilla 
with few teeth along its anterior border, second suborbital expanded covering lower 
limb of the preopercle, a single series in the dentary, and absence of scales over the 
caudal-fin lobes and of a pouch scale on the base of the caudal fin in males. The 
validity of those characters were subsequently discussed by some authors (Malabarba 
& Malabarba, 1994) but a phylogenetic definition of the genus is still pendant and 
Eigenmann's definition is still in use for the generic placement of the new species. 

Géry (1977) identified two groups of Bryconamericus, B. diaphanus-group and 
B. peruanus-group. These groups are artificial, however Géry (1977) is the only 
available paper presenting an overview of the genus. Bryconamericus uporas may be 
included within the B. diaphanus-group because it has 15 to 25 anal-fin rays and 4-6 
transverse scales above the lateral line. However, the shape of premaxillary and 
maxillary teeth of B. uporas differs from that present in the species included in that 
group. 



A NEW BRYCONAMERICUS FROM URUGUAY BASIN 



161 




Fig. 7 
Map showing the left tributaries of no Parana and right tributaries of rio Uruguay, Misiones, 
Argentina, with the geographical distribution of Bryconamericus uporas sp. n.: 1, arroyo Once 
Vueltas (type locality); 2, arroyo Toro; 3, arroyo Fortaleza; 4, arroyo Yabotf-Minf. 

The following species of Bryconamericus were described from southern South 
America: B. iheringi, B. eigenmanni, B. lambari, B. thomasi, B. exodon, B. stramineus 
fa junior synonym of B. exodonl), B. sylvicola, and B. agna. 



162 



J. R. CASCIOTTA £7ML. 



.._,._ — . 




Fig. 8 
Habitat of Bryconamericus uporas sp. n.. arroyo Fortaleza, Misiones, Argentina. 



A NEW BRYCONAMERICUS FROM URUGUAY BASIN 163 

Bryconamericus uporas differs from B. exodon in having an aligned outer row 
of premaxillary teeth and deeper body (28.9-32.2 vs. 22.9-26.8 % in SL). Also a wide 
black lateral band is present in B. uporas, whereas a wide silvery band occurs in B. 
exodon. From B. eigenmanni, B. uporas is easily distinguished by the higher number 
of branched anal-fin rays (18-20 vs. 15- 17), lack of a dot on upper half of dorsal fin, 
and several morphometric characters such as longer caudal peduncle (14.9-17.7 vs. 
18.0-23.0) and shorter distance between origins of pelvic and anal fins (13.7-18.0 vs. 
16.0-25.8). Bryconamericus uporas differs from B. iheringi by lower body depth 
(28.9-32.2 vs. 33.7-38.1 % in SL), and the shorter predorsal distance (46.6-50.5 vs. 
55.5-56.8 % in SL). The wide dark lateral band present in B. uporas distinguishes this 
species from B. iheringi, B. sylvicola, and B. lambari, but a similar band occurs in B. 
agna and B. thomasi. The later species has a peculiar deeper caudal peduncle (15.6- 
17.4 vs. 10.4-11.0 % in SL in B. uporas). The lower body and number of branched 
anal-fin rays differ B. uporas from B. sylvicola (28.9-32.2 vs. 36.1-40.7 % in SL); 
(19-22 vs. 22-25). Also B. uporas has a wide black lateral band vs. a very narrow 
band. Bryconamericus uporas differs from B. agna in having lower body (28.9-32.2 
vs. 34.1-39.8 % in SL), longer caudal peduncle length (14.9-17.7 vs. 10.9-11.8 % in 
SL) and higher number of dentary teeth (7-10 vs. 6-7). 

Among the species described from southern South America, it is possible to 
find some species with a large geographical distribution while others have a very 
restricted geographical distribution. Bryconamericus iheringi and B. exodon occur in 
a wide geographical area, living in different types of environments of the Rio de la 
Plata basin and Laguna dos Patos system. The two species occur in large and small 
rivers, and the former also inhabits shallow and deep ponds. Besides, B. iheringi 
represents the southernmost record of the genus in the rio Sauce Grande drainage (38° 
45' S), in the south of Buenos Aires Province (Casciotta et al. ,1999). 

Among the species with restricted distribution, B. eigenmanni lives in 
endorrheic basins of the nos Primero and Pichanas, Province of Cordoba, Central 
Argentina (Miquelarena & Aquino, 1999). Bryconamericus thomasi occurs in the 
upper basin of the rio Bermejo, rio Pasaje-Juramento-Salado, and rio Lipeo, Provinces 
of Salta and Jujuy in Argentina, and Departamento Tarija in Bolivia (Miquelarena & 
Aquino, 1995). Bryconamericus lambari inhabits in tributaries of Laguna dos Patos 
system, in Brasil (Malabarba & Kindel, 1995). Bryconamericus sylvicola (Braga, 
1998) lives only in arroyo Urugua-i above the falls. Bryconamericus agna is only 
known from the type locality in the arroyo Tabay, rio Parana basin (Azpelicueta & 
Almirón, 2001). All those species live in streams and small rivers, with sandy and 
rocky bottom, shallow (0.5 m) or deep (2 m) pools, and current water. 
Bryconamericus uporas belongs to that group of species with restricted distribution, 
being present in headwaters of different streams which flow into the rio Uruguay in 
Misiones Province. 



164 J- R - CASCIOTTA ETAL. 

IDENTIFICATION KEY FOR SPECIES OF BRYCONAMERICUS FROM 
SOUTHERN SOUTH AMERICA 

1 Premaxillary teeth of the outer row not aligned B. exodon 

Premaxillary teeth of the outer row aligned 2 

2 Wide lateral band 3 

Narrow lateral band 5 

3 First four dentary teeth with similar size followed by other ones much 

smaller 4 

Six or seven dentary teeth decreasing in size anteroposteriorly B. agna 

4 Premaxillary and maxillary teeth expanded distally B. uporas sp.n. 

Premaxillary and maxillary teeth not expanded distally B. thomasi 

5 Caudal peduncle length 1 8-23 % of SL, dorsal fin brownish . . . B. eigenmanni 
Caudal peduncle length 11-18 % of SL, dorsal fin hyaline or with a 

dark distal area 6 

6 Body depth 24.6-29.8 % of SL B. lambari 

Body depth 3 1 .5-40.7 % of SL 7 

7 Anal-fin base 19.5-25.7 % of SL B. iheringi 

Anal fin-base 28.0-33.7 % of SL B. sylvicola 

ACKNOWLEDGEMENTS 

Authors thank C. Tremouilles for the drawings. This paper was supported by 
grants from Consejo Nacional de Investigaciones Cientificas y Técnicas (CONICET) 
and Comisión de Investigaciones Cientificas de la Provincia de Buenos Aires (CIC). 

REFERENCES 

Azpelicueta, M. & Almirón, A. 2001. A new species of Bryconamericus (Characiformes, 

Characidae) from Parana basin in Misiones, Argentina. Revue suisse de Zoologie 108: 

275-281. 
Braga, L. 1998. Una nueva especie de Bryconamericus (Ostariophysi, Characidae) del rio 

Urugua-i, Argentina. Revista del Museo Argentino de Ciencias Naturales Bernardino 

Rivadavia 8 (3): 21-29. 
Casciotta, J., Almirón, A., Cione, A. & Azpelicueta, M. 1999. Brazilian freshwater fish 

assemblages from southern pampean area. Argentina. Biogeographica 75: 67-78. 
Eigenmann, C. H. 1927. The American Characidae. Memoirs of the Museum of Comparative 

Zoology 43: 311-428. 
Eigenmann, C. H., McAtee, W. L. & Ward, D. P. 1907. On further collections of fishes from 

Paraguay. Annals of the Carnegie Museum 4: 1 10-157. 
Evermann, B. W. & Kendall, W. C. 1906. Notes on a collection of fishes from Argentina, 

South America; with descriptions of three new species. Proceedings of the United 

States National Museum 31: 67-108. 
Fowler, H. W. 1940. Zoological results of the second bolivian expedition for the Academy of 

Natural Sciences of Philadelphia 1936-1937. Part I. - The fishes. Proceedings of the 

Academy of Natural Sciences of Philadelphia 42: 43-103. 
GÉRY, J. 1977. Characoids of the World. TFH Publications Inc., Neptune Citv, New Jersey, 

672 pp. 



A NEW BRYCONAMERICUS FROM URUGUAY BASIN ]65 



Malabarba, L. R. & Kindel, A. 1995. A new species of the genus Bryconamericus Eigen- 
mann. 1907 from southern Brazil (Ostariophysi: Characidae). Proceedings of the Biolo- 
gical Society of Washington 108: 679-686. 

Menni, R. C. Lopez. H. L. & Arâmburu, R. H. 1988. Ictiofauna de Sierra de la Ventana y 
Chasicó (Prov. de Buenos Aires, Argentina). Zoogeografia y parâmetros ambientales. 
Anales del Museo de Historic! Natural de Valparaiso 19: 75-84. 

MiQUELARENA, A. M. & Aquino, A. E. 1995. Situation taxonómica y geografica de Bryco- 
namericus thomasi Fowler, 1940 (Teleostei, Characidae). Revista Brasileira de Bio- 
logia 55: 559-569. 

MiQUELARENA, A. M. & Aquino, A. E. 1999. Taxonomic status and geographic distribution of 
Bryconamericus eigenmanni Evermann & Kendall, 1906 (Characif ormes: Characidae). 
Proceedings of the Biological Society of Washington 1 13: 523-530. 

Taylor, W. R. & Van Dyke, G. C. 1985. Revised procedures for staining and clearing small 
fishes and other vertebrates for bone and cartilage study. Cybium 9: 107-1 19. 



Revue Suisse de Zoologie 109 (1): 167-176; mars 2002 



A contribution to the study of the genus Centromerus Dahl 
(Araneae: Linyphiidae) in caves of the Balkan Peninsula 

Christo DELTSHEV 1 & Bozidar P. M. CURCIC 2 

1 Institute of Zoology, Bulgarian Academy of Sciences, bid. Tsar Osvoboditel 1, 
1000-Sofia. Bulgaria. 

2 Institute of Zoology, Faculty of Science, Studentski Trg 16, 
11000-Belgrade, Yugoslavia. 

A contribution to the study of the genus Centromerus Dahl (Araneae: 
Linyphiidae) in caves of the Balkan Peninsula. - Centromerus serbicus 
Deltshev sp. n. is described from specimens previously attributed to C. pr. 
dacicus. The taxonomic relationships between C bulgarianus, C. dacicus 
and C serbicus sp. n. are discussed and new illustrations are presented. 
These are closely allied and strictly vicariant species forming a super- 
species. C. acutidentatus Deltshev, sp. n. is described from caves in Serbia, 
Bulgaria and Macedonia. This species is closely related to C obenbergeri 
Kratochvfl & Miller and to C gentilis Dumitrescu & Georgescu. The 
hitherto unknown female from C.obenbergeri is described. 

Key-words: Araneae - Centromerus - taxonomy - caves - Balkan 
Peninsula. 

INTRODUCTION 

Deltshev and Curcic (1997) analyzed the taxonomic and phylogenetic inter- 
relationships between the spiders of the C. europaeus species group (i.e. C. bulga- 
rianus Drensky, C. subcaecus Kulczyhski, C. dacicus Dumitrescu & Georgescu) from 
caves of the Balkan Peninsula. A troglobitic spider found in the Zlotska Pecina Cave 
(Serbia) was first identified as C. prope dacicus and the study of additional material in 
comparison with material of C. dacicus later showed that the species is new to 
science. It is here discussed as C. serbicus sp. n. Another new species C. acutiden- 
tatus sp. n., which is closely related to C. obenbergeri Kratochvfl & Miller and C. 
gentilis Dumitrescu & Georgescu. was collected from the leaf litters and from caves 
of Serbia, Bulgaria and Macedonia. Centromerus obenbergeri is redescribed and the 
hitherto unknown female of this species is illustrated for the first time. 

Abbreviations: Names of collectors are abbreviated as follow: R.N. Dimitri- 
jevic - RND; O.S. Karamata - OSK; L.R. Lucie - LRL; S.E. Makarov - SEM; A.M. 
Petrovic - AMP; V.M. Pesic - VMP; S.V. Stancovic - SVS; G.S. Stojanovic - GSS; 
E.A. Stojkoska; V.T. Tomic - VTT; B.P.M. Curcic - BPMC; S.B. Curcic - SBC; N.B. 
Curcic - NBC; B. Petrov - BP. 
All measurements are in mm. 



Manuscript accepted 18.08.2001 



168 C. DELTSHEV & B. P. M. CURCIC 



DESCRIPTIONS (AND REDESCRIPTION) 

Centromerus serbicus Deltshev sp. n. Figs 1, 2, 7, 8 

Centromerus europaeus (Simon): Fage, 1931 (partim; misidentification):170-171; Drensky, 
1936: 94; C. subcaecus (Kulczyfiski): Kratochvfl & Miller, 1938 (partim; misidentification): 
109-111; Thaler & Hoefer, 1987: 390-393; C. prope dacicus: Deltshev & Curcic, 1997: 49-55, 
figs 2, 5, 8, 10: Deltshev et al, 1997: 37P. 

Material examined: YUGOSLAVIA: Zlotska Pecina Cave (Lazareva Pecina Cave), 
vili. Zlot, near Bor, Serbia, 1 6 holotype, 3 S, 30 5 paratypes, 21-23 November 1995, collec- 
ted by R.N.D, S.E.M., L.R.L.; 2 S, 5 5 paratypes, 16-17 June 1996, coll. RND, OSK, SEM. 
VTT; 1 6, 16 ? paratypes, 26 October 1996, coll. RND, OSK, SEM, LRL; Vernjikica Cave, 
vili. Zlot near Bor, 1 9 paratype, 17 June 1996, 4 6, 16 9 paratypes, October 1996, coll. RND, 
OSK, SEM, VTT; Resavska Pecina Cave, near Despotovac, East Serbia. 2 S, 3 9, paratypes, 
14 June 1996, coll. RND. OSK. SEM. VTT. 

Depository: The collection of the Institute of Zoology. Faculty of Biology, University 
of Belgrade, Yugoslavia; 1 6 and 3 9 paratypes are deposited in the collection of the Muséum 
d'historié naturelle in Geneva, Switzerland. 

Comparative material: C. bulgarianus (lectotype, deposited in the collection of Zoo- 
logy, Sofia): Fig. 5 ; C. dacicus, Figs 3, 4, 6 (specimens from Pestua "E.A. Martel", 8 March 
1960, deposited in the collection of Institut de Spéologie "E. Racovitza"). 

Etymology: The species name is derived from Serbia. 

Diagnosis: C. serbicus sp. n. is a blind species, closely related to C. bulgaria- 
nus and C. dacicus, distinguished from them by the following differential features 
(figs 1-7): C. serbicus sp. n. is larger and possesses a larger palp; the embolic division 
is similar but the terminal apophysis is longer and larger (figs 5-7); the lamella (sensu 
Merret, 1963) appears to be similar to those of C. bulgarianus and C. dacicus, but 
differs in details (figs 1-4); females are almost indistinguishable, but there are diffe- 
rences in details of the epigyne. 

Description: Male/female: Total length 1.8/2.16. Céphalothorax, length 
0.79/.0.82, width 0.61/0.57; sternum, length 0.39/0.42, width 0.42/0.42; abdomen, 
length 0.97/1.26. Céphalothorax similar in both sexes, yellow to yellow brown. 
Abdomen grey to pale grey. Eyes completely absent. Chelicerae yellow brown, armed 
with 3 teeth on outer row and 3-4 denticles on inner row. Legs: IV-I-II-III. I: Fe. 
1.15/1.26 , Pl+Ti. 1.40/1.40. Mt. 0.9/0.9. Ta. 0.61/0.57. Femora I with a prolateral 
spine in apical half. Tibiae I-III with 2 dorsal spines. Tibia IV with 1 dorsal spine. 
Metatarsi I-II with 1 small dorsal spine. 

Male palp (figs 1. 2. 7). With a strong dorsal spine on patella. Cymbium with a 
postero-dorsal protuberance. Paracymbium large, with serrated inner margin; 3-4 
short hairs near proximal end. Embolic membrane well - presented, with strongly 
serrated outer margin. Terminal apophysis lamellar. Lamella characteristically built, 
continuous with radix. 

Epigyne presented on figure 8. 

Affinities: C. serbicus sp. n., C. bulgarianus and C. dacicus are closely allied 
and strictly vicariant species forming a superspecies. They belong to the europaeus- 
group of the genus Centromerus on the Balkan peninsula. All these taxa are similar, 
they have limited ranges and probably, represent the descendants of a common an- 
cestor; it is assumed that this ancestral form is no longer present in the epigean fauna 
and that it has been replaced by an extant Centromerus (Deeleman-Reinhold, 1976, 
Deltshev & Curcic. 1997). 



CENTROMERUS IN CAVES OF THE BALKANS 



169 




Figs 1-4 
1, Centromerus serbicus Deltshev sp. n., left male palp, external view; 2, ditto, internal view; 3, 
C. dacicus Dumitrescu & Georgescu, left male palp, external view; 4, ditto, internal view. Scale 
line 0.1 mm. 



170 



C. DELTSHEV & B. P. M. CURCIC 






5,6,7 




Figs 5-8 
5-7, terminal apophysis of Centromerus bulgarianus (Drensky) (5), C. dacicus Dumitrescu & 
Georgescu (6), C. serbicus Deltshev sp. n. (7); 8, C. Serbiens Deltshev sp. n. vulva and epigyne, 
ventral view. Scale lines 0.1 mm. 



CENTROMERUS IN CAVES OF THE BALKANS ] 7 ] 

Centromerus acutidentatus Deltshev sp. n. Figs 9-21 

Centromerus obenbergeri (Kratochvfl & Miller): Curcic et ai, 1999 (misidentifucation): 7P. 

Material examined: YUGOSLAVIA: Monastery Cave I (at the entrance in leaf litter), 
vili. Selacka near Minicevo, Serbia, 1 â holotype, 4 9 paratypes, 15 November 1997 (coll. 
RND, OSK, VTT. SBC, NBC); Avala, Carpivev Best (leaf litter) 1 9 paratype, 21-24.09.1997 
(coll. BPMC, SEM, LPvL, VTT); MACEDONIA: Ubava Cave, Matka, klisura Treske, 1 $ 
paratype, 1 juv, 13 July 2000 (coll. SEM, VTT, SBC, GSS, EAS, SVS); BULGARIA: 
Sturshelitsa Cave, vili. Goleshevo, 1 9 paratype, 14 April 1993 (coll. BP); Gaber Reserve, Mt. 
Pianets, Kjustendil, 1 â paratype, 06 April 2001 (coll. BP). 

Depositon". The holotype and 1 paratype (Monastery Cave I) are deposited in the 
collection of the Muséum d'historié naturelle in Geneva, Switzerland. The rest, 2 female 
paratypes from the same locality, as well as the material of Ubava Cave, Macedonia is in the 
collection of the Institute of Zoology, Faculty of Biology, University of Belgrade, Yugoslavia. 
The material from the Sturshelithsa Cave and Gaber Reserve, Bulgaria is in the collection of 
the Institute of Zoology, Sofia, Bulgaria. 

Etymology: Latin : acutus = sharp; dentatus = dentate.The specific name refers 
to the sharp teeth on the paracymbium. 

Diagnosis: C. acutidentatus sp. n. is clearly related to C. gentilis Dumitresco & 
Georgesco and to C. obenbergeri Kratochvfl & Miller known from the caves in 
Romania and Montenegro. The new species differs from them by the larger para- 
cymbium armed with the bigger and sharper teeth, by a caharacteristic lamella with 
two denticles on its posterior border and by a longer terminal apophysis (figs 9-17). 
Females are very similar, but there are differences in details of the epigynes and 
vulvae (figs 18-21). 

Description: Male/female: Total length 2.16/1.80. Céphalothorax, length 
1.0/0.72, width 0.79/0.51; sternum, length 0.54/0.54, width 0.42/0.42; abdomen, 
length 1.08/1.08. Céphalothorax similar in both sexes, yellow to yellow brown. 
Abdomen grey to pale grey. Eyes surrounded by a narrow black area; anterior 
medians almost in contact with each other and ca. 0.75 diam. apart from the laterals; 
posterior medians ca. 1.25 diam. apart from each other and ca. 0.75 diam. from 
laterals. Chelicerae yellow brown, armed with 3 teeth on outer row and 4 denticles on 
inner row. Legs: IV-I-II-III. I: Fe. 0.9/0.72, Pl.+Ti. 1.07/1.07, Mt. 0.61/0.50, Ta. 
0.54/0.42. Femora I with a prolateral spine in apical half. Tibiae I-III with 2 dorsal 
spines. Tibia IV with 1 dorsal spine. Metatarsi I-II with small 1 dorsal spine. 
Metatarsi I-III with a single trichobothrium. 

Male palp (figs 9-17). A strong dorsal spine on patella. Cymbium with a 
postero-dorsal protuberance. Paracymbium large, with 13-16 well - developed and 
sharp teeth on inner margin; 4-5 short hairs near proximal end. Embolic membrane 
well - presented, with strongly serrated outer margin. Terminal apophysis lamellar. 
Lamella characteristically built, continuous with radix. 

Epigyne and vulva presented on figures 18-21. 

Affinities: C. acutidentatus sp. n., C. gentilis and C. obenbergeri are closely 
allied. They belong to the sylvaticus-group of the genus Centromerus on the Balkan 
peninsula. C. obenbergeri and C. gentilis are known only from caves, C. acutiden- 
tatus was collected in caves and from forest. 



172 



C. DELTSHEV & B. P. M. CURCIC 




Figs 9-12 
9, Centromerus acutidentatus Deltshev sp. n.. right male palp, external view; 10, ditto, internal 
view (specimen from Monastery Cave I, Serbia); 11, ditto left male palp, external view; 12, 
ditto, internal view (specimen from Gabra Reserve, Bulgaria). Scale line 0.2 mm. 



CENTROMERUS IN CAVES OF THE BALKANS 



173 






13, 14 



15, 16, 17 



Figs 13-17 
13, Centromerus acutidentatus Deltshev sp. n., right male palp external view; 14, ditto, internal 
view (specimen from Ubava Cave, Macedonia); 15, C. acutidentatus Deltshev sp. n., terminal 
apopysis (Monastery Cave I, Serbia); 16, ditto (Gabra Reserve, Bulgaria); ditto (Ubava Cave, 
Macedonia). Scale lines 2.0. mm. 



174 



C. DELTSHEV & B. P. M. CURCIC 





Figs 18-21 
18, 20, Centromerus acutidentatus Deltshev sp. n., epigyne and vulva, ventral (18) and dorsal 
(20) view (specimen from Monastery Cave I, Serbia); 19, 21. ditto (Gabra Reserve, Bulgaria). 
Scale line 0.1mm. 



Centromerus obenbergeri Kratochvfl & Miller Figs 22-26 

Centromerus subcaecus (Kulczyriski): Kratochvfl, 1934 (misidentification): 188-189, fig. 6.; C. 
obenbergeri: Kratochvfl & Miller, 1938: 113. 

Material examined: MONTENEGRO: Lipska Pecina cave, vili. Lipa, Cetinje, 1 <?, 1 9 , 
8. September 2000 (coll. SBC, AMP, VMP). 

Depository: The collection of the Muséum d'histoire naturelle in Geneva, Switzerland. 

Description: Male/female: Total length 1.44/1.44. Céphalothorax, length 0.79/ 
0.72, width 0.64/0.54; sternum, length 0.34/0.42, width 0.46/0.42; abdomen, length 
0.72/0.9. Céphalothorax similar in both sexes, yellow to yellow brown. Abdomen 
grey to pale grey. Eyes, small and surrounded by a narrow black area; anterior 
medians almost in contact with each other and ca. 0.75 diam. apart from the laterals; 
posterior medians ca. 1.25 diam. apart from each other and ca. 0.75 diam. from 
laterals. Chelicerae yellow brown, armed with 3 teeth on outer row and 4 denticles on 
inner row. Legs: IV-I-II-III. I: Fe. 1.0/0.9, Pl.+Ti. 1.21/1.11, Mt. 0.82/0.72, Ta. 
0.54/0.54. Femora I with a prolateral spine in apical half. Tibiae I-III with 2 dorsal 



CENTROMERUS IN CAVES OF THE BALKANS 



175 






Figs 22-26 
22, Centromerus obenbergeri Kratochvfl & Miller, left male palp, external view; 23, ditto, 
internal view; 24, terminal apophysis; 25, epigyne, ventral view; 26, vulva, dorsal view. Scale 
line 0.1 mm (all figures). 



176 c - DELTSHEV & B. P. M. CURCIC 

spines. Tibia IV with 1 dorsal spine. Metatarsi I-II with 1 small dorsal spine. Meta- 
tarsi I-III with a single trichobothrium. 

Male palp (figs 22-24). A strong dorsal spine on patella. Cymbium with a 
postero-dorsal protuberance. Paracymbium large, with 12 teeth on inner margin; 4 
short hairs near proximal end. Embolic membrane well - presented, with strongly 
serrated outer margin. Terminal apophysis lamellar. Lamella characteristically built, 
continuous with radix. 

Epigyne and vulva presented on figures 25, 26. 

ACKNOWLEDGEMENTS 

We thank the following colleagues: R.N. Dimitrijevic, L.R. Lucie, S.E. 
Makarov, O.S. Karamata, V.T. Tomic; S.B. Curcic, N.B. Curcic (Belgrade), B. Petrov 
(Sofia), G.S. Stojanovic, E.A. Stojkoska, S.V. Stancovic (Scopie), A.M. Petrovic, 
V.M. Pesic (Podgorica) for presenting their materials. 

REFERENCES 

Curcic, B. P. M., Deltshev, C, Dimitrijevic, R. N., Karamata, O. S., Tomic, V. T.. Curcic, 
S. B. & Curcic, N. B. 1999. On some cave-dwelling spiders (Araneae, Arachnida) 
from Serbia, Yugoslavia. Part V. Archives of Biological Sciences 51 (1): 1P-&P. 

Deeleman-Reinhold, C 1976. Distribution patterns of European cave spiders. Proceedings of 
the International Symposium on Cave Biology, Oudtshoom: 25-35. 

Deltshev, C & Curcic, B.P. M. 1997. Contribution to the knowledge of the group europaeus 
of Centromerus Dahl (Linyphiidae. Araneae) in the Balkan Peninsula. Revue suisse de 
Zoologie 104 (10): 49-55. 

Deltshev, C, Curcic, B. P. M., Dimitrijevic. R. N., Makarov, S. E., Lucie, L. R. & Tomic, 
V. T. 1997. Additional report on cave-dwelling spiders (Araneae, Arachnida) from 
Serbia, Yugoslavia. Archives of Biological Sciences 49 (3-4): 37P-38P. 

Drensky, P. 1936. Katalog der echten Spinnen (Araneae) der Balkanhalbinsel. Sbornik of 
Bulgarian Academy of Sciences 32: 1-223. 

Fage, L. 193 1 : Araneae. Biospeologica 5 e série. Archives de la Zoologie expérimentale et gene- 
rateli: 1-291. 

Kratochvil, J. 1934. Liste générale des Araignées cavernicoles en Yougoslavie. Prirodo- 
slovne razprave Ljubljana 2: 165-226. 

Kratochvil. J. & Miller, F. 1938. Sur le problème des araignées cavernicoles du genre Cen- 
tromerus de la Péninsule balkanique. Mitteilungen aus königlichen naturwissenschaft- 
lichen Instituten in Sofia 1 1: 107-1 13. 

Merret, P. 1963. The palpus of male spiders of the family Linyphiidae. Proceedings of the 
Zoological Society of London 140 (3): 347-467. 

Thaler, K. & Hoefer, H. 1987. Eine weitere Art der Gattung Centromerus Dahl 1886 in 
Mitteleuropa: C sp. prope subcaecus Kulczyhski 1914 (Arachnida: Araneae: Liny- 
phiidae). Senkenbergiana biologica 68:389-396. 



Revue suisse de Zoologie 109 ( 1 ): 177-187; mars 2002 



Zwei Bythinini neu für Friaul-Julisch Venetien (Italien): 
Gasparobythus kahleni sp. n. und Tychobythinus xambeui manfredi 
ssp. n. (Coleoptera, Staphylinidae, Pselaphinae) 

Lorenz NEUHÄUSER-HAPPE 

Ökoteam - Institut für Faunistik und Tierökologie, Bergmanngasse 22, A-8010 Graz. 

Two Bythinini new to Italy: Gasparobythus kahleni sp. n. and Tycho- 
bythinus xambeui manfredi ssp. n. (Coleoptera, Staphylinidae, Psela- 
phinae). - Gasparobythus kahleni sp. n. is described from Friaul near to 
the borderline of Italy and Slovenia and is the second species of the genus. 
Tychobythinus xambeui manfredi ssp. n. is described from the riverbanks of 
the Tagliamento near to Udine. The main characters of T. xambeui 
(Guillebeau, 1888) are provided. 

Key-words: Coleoptera - Staphylinidae - Pselaphinae - Gasparobythus - 
Tychobythinus - new species - north-eastern Italy. 

EINLEITUNG 

Besuchet (1985) hat basierend auf der Kombination von drei Merkmalspaaren 
(Fehlen bzw. Vorhandensein von basaler Halsschildfurche sowie einem marginalen 
Basalgrübchen und Kielchen auf den Flügeldecken) vier, wenige Arten umfassende 
Gattungen mit ausschließlich troglobionten Vertretern innerhalb des Bythinini-Sub- 
tribus Machaeritina unterschieden und in einem Bestimmungsschlüssel dargestellt 
(Machaerites, Bathybythus, Priohobythus, Antrobythus). Im Unterschied zu den 
anderen Gattungen des Subtribus Machaeritina ist diesen zumindest das Fehlen eines 
der oben genannten Positivmerkmale gemeinsam. 

Aufgrund des Auftretens einer neuen Merkmalskombination wurde von Poggi 
(1992) erst vor wenigen Jahren mit der Typusart Gasparobythus tergestinus eine neue 
Gattung begründet. Diese ist durch das Fehlen der basalen Halsschildfurche bei 
gleichzeitigem Vorhandensein des marginalen Grübchens und Kielchens auf den 
Flügeldecken ausgezeichnet. 

Während eine der zwei in der vorliegenden Arbeit behandelten Arten des 
Subtribus Machaeritina der umfangreichen Gattung Tychobythinus angehört, konnte 
die zweite Art der neuen Gattung Gasparobythus zugeordnet werden und ist somit 
erst die zweite Art dieses bislang nur aus Friaul-Julisch Venetien bekannten Taxons. 
Aufgrund dieser Sachlage wurde die Beschreibung der neuen Art ausführlicher 
gestaltet und mit einer Habitusabbildung sowie mit weiteren Detailansichten ergänzt. 
Auch hinsichtlich der Tychobythinus- Avi erfolgt - aufgrund bislang ungenauer und 
fehlerhafter Angaben zu dieser Art - eine Kurzbeschreibung. 



Manuskript angenommen 08.08.2001 



178 L. NEUHÄUSER-HAPPE 

Die Belegtiere beider Arten stammen von Manfred Kahlen (Hall in Tirol), 

dessen systematische, nach detaillierten tiergeografischen und ökologischen Gesichts- 
punkten ausgerichteten Aufsammlungen in den Südalpen bereits in den vergangenen 
Jahren und Jahrzehnten zu zahlreichen bemerkenswerten Nachweisen geführt haben 
und auf dessen Initiative die nunmehr vorliegenden Untersuchungen zurückgehen. In 
beiden Fällen wurden im Gebiet der Fundlokalitäten - u.a. auch unter Beisein des 
Autors - Nachsuchen betrieben, die jedoch nur hinsichtlich der neuen Gasparobythus- 
Art zu einem weiteren Fund durch Kahlen führten. In diesem Zusammenhang ist 
anzumerken, dass aufgrund ihrer versteckten Lebensweise Nachweise dieser Arten 
nur ausgesprochen schwer zu erbringen sind; so konnten von beiden Arten der 
Gattung Gasparobythus bis heute noch keine Weibchen gefunden werden. Die spär- 
lichen Nachweise der zweiten im Folgenden behandelten Art der Gattung Tycho- 
bythinus gelangen bislang überhaupt nur in Sekundärlebensräumen, die eine autöko- 
logische Charakterisierung derzeit nicht zulassen. 

MATERIAL 

Das untersuchte Belegmaterial stammt aus folgenden Museen: 

TLMF = Tiroler Landesmuseum Ferdinandeum. Innsbruck (Coli. Kahlen) 

MHNG = Muséum d'histoire naturelle. Genève 

MSNG = Museum civico di Storia naturale "G. Doria'\ Genova 



Gasparobythus kahleni sp. n. 

Holotypus (<?): Val. Clabocgnac bei Prepotto-Bodigoi, W Udine (46°04'N, 13°31'E), 
150m. Bachgraben. Wurzelwerk in lehmigem Kies in Felsnischen, 5.5.1989. Manfred Kahlen 
leg. (TLMF). Paratypus (6): Val. Nestadiuzza bei Prepotto-Bodigoi, W Udine (46°03'N, 
13°30'E), 150m, Wurzeigesiebe, 3.5.1986, Manfred Kahlen leg. (TLMF). 

Habitusansicht: Abb. 1 . 

Beschreibung: Länge: 1,08-1,14 mm; Färbung gelblich-rotbraun; Fühler, Ma- 
xillarpalpen und Beine gelbbraun; grobe Behaarung goldglänzend und aus ca. 0,06 
mm langen Haaren zusammengesetzt; zusätzlich zur Grundbehaarung auf der Ober- 
seite von Kopf, Halsschild. Flügeldecken und Hinterleib mit bis zu 0,25 mm langen, 
abstehenden Borsten, diese an ihrer Spitze mit einem abgesetzten dünnen Endteil. 

Kopf: deutlich breiter (0,26-0,3 mm) als lang (0,22-0,23 mm); Stirnbreite 
(0,13-0,14 mm) ca. halb so groß wie die maximale Kopfbreite, diese nur wenig vor 
dem Hinterrand gelegen, sodass die Kopfform eine charakteristische, an Arten der 
Gattung Decatocerus erinnernde dreieckige Grundform erhält; Medianeindruck der 
Stirn mit vereinzelten oberflächlichen Punkten besetzt und glänzend, doppelt so lang 
wie breit und gegen den Vorderrand stärker vertieft; die Außenkanten der Stirn 
scharfkantig, auf einer Strecke von 0,05 mm subparallel verlaufend, danach konti- 
nuierlich erweitert: über der Fühlereinlenkung mit jeweils einer kleinen, aber mar- 
kanten Aufwölbung; Kopfoberseite beidseitig der Stirnfurche punktiert, gegen den 
Kopfhinterrand Punktierung oberflächlich: Scheitel mit breit abgeflachter Beule, die 
gegen die Stirnfurche, die tentorialen Grübchen und den schmalen jeweils von diesen 



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179 




Abb. 1 
Gasparobythus kahleni sp. n., Dorsalansicht. Maßstab: 0,5 mm. 



1 go L - NEUHÄUSER-HAPPE 

zur Stirnfurche verlaufenden Rinnen steil abfällt; Hinterrand des Kopfes deutlich 
konkav, median mit kleinem Kielchen, das dorsal nach kurzem Verlauf verflacht und 
mit einem kleinen aber distinkten Körnchen endet; Augen fehlen; Schläfen mit etwas 
verdichteter nach hinten gegen den Hals gerichteter längerer Behaarung; Unterseite 
des Kopfes mit tiefer Querfurche, dahinter jeweils ein kurzes von den Seiten schräg 
nach vorne gegen die Mitte und bis zur Basis der Querfurche verlaufendes Kielchen; 
die Kielchen voneinander durch ein medianes, von hinten in die Querfurche mün- 
dendes Grübchen getrennt, zwischen den Kielchen beidseitig des medianen Grüb- 
chens einige kurze Borsten; unterseits vor dem Kopfhinterrand mit einer Querreihe 
von 10 langen, geschwungenen Borsten (Abb. 2); Maxillarpalpen (Abb. 4): 2. Glied 
(Länge: 0,15 mm) ventral mit 13 in schrägen Querreihen angeordneten Tuberkeln, 

3. Gld. (Länge: 0,05-0,055 mm) in seiner Vorderhälfte ventral mit 3 Tuberkeln, 

4. Gld. von normalem Aussehen (0,17-0,18 mm lang, max. Breite 0,065 mm); Fühler: 
so lang wie Kopf und Halsschild zusammen, 1. Glied 0,15 mm lang, ca. 4 mal so lang 
wie maximal breit, in seinem ersten Drittel bei dorsaler Aufsicht nur von halber Breite 
und apical keulenförmig verdickt, 2. Gld. so breit wie das vorhergehende, deutlich 
länger (0,055 mm) als breit (0,04 mm), 3. Gld. schmäler als das vorhergehende, 
geringfügig länger (0,03 mm) als breit (0,025 mm), 4. bis 8. Gld. etwa so breit wie 
das 3. Glied, etwa gleich lang wie breit, davon das 4. und 6. kaum merklich kleiner 
dimensioniert, 9., 10. und IL Gld. breiter als das jeweils vorhergehende, 9. fast, 10. 
deutlich doppelt so breit wie lang, 1 1 . Gld. um ein Viertel länger als breit. 

Halsschild: deutlich breiter (0,29-0,32 mm) als lang (0,25-0,27 mm); glatt und 
glänzend, nur an den Seiten mit sehr oberflächlicher Punktierung; in der Mitte auf 
einer breiten, sich vom Vorder- bis Hinterrand erstreckenden Fläche unbehaart und 
mit Spiegelglanz; Halsschildseiten gegen den Vorderrand in steilem und etwas 
kantigem Bogen verschmälert und vor dem Vorderrand seitlich eingeschnürt, gegen 
die Basis sanft und leicht konkav verengt; im hinteren Drittel der Halsschildseiten mit 
einem tiefen Grübchen; basale Halsschildfurche fehlt; Hinterrand gerandet und leicht 
doppelbuchtig. 

Flügeldecken: miteinander verwachsen, zusammen deutlich breiter als lang 
(max. Breite: 0,46-0,52 mm, Nahtlänge: 0,36-0,41 mm); entlang der Naht glatt und 
nur verstreut punktiert, ansonsten besonders gegen die Seiten mit dichterer und 
gröberer Punktierung, wobei der Abstand der Punkte zueinander meist nicht kleiner 
als ihr Durchmesser ist; an der Flügeldeckenbasis mit 2 tiefen ovalen Grübchen, 
wovon das äußere etwas breiter ist; zwischen beiden mit kurzer Dorsalfalte; 
Humeralfalte fehlt; seitlich hinter dem äußeren Basalgrübchen mit einem kleineren, 
ebenso tiefen marginalen Grübchen, von dessen Außenrand ein Kielchen gegen den 
umgeschlagenen Hinterrand der Flügeldecken verläuft. 

Hinterleib: glänzend und gegen die Seiten mit sehr verstreuter und nur ange- 
deuteter körnchenartiger und flachgrubiger Punktierung. 

Flügel: fehlen. 

Beine: schlank, besonders der Metafemur von auffällig geringer Dicke (0,065 
mm); Metatibia am apicalen Ende leicht nach innen gebogen. 



ZW'HI BYTHININI NIL' FUR ITALIAN 



181 






Abb. 2-4 
Lateralansicht des Kopfes von Gasparobythiis kahleni sp. n. (2) und Tychobythinus xambeui 
manfredi ssp. n. (3); Maxillarpalpen von Gasparobythiis kahleni sp. n., Ventralansicht (4). 
Maßstab: 0,1 mm. 

Aedeagus (Abb. 5): Innensackstrukturen asymmetrisch und schwach skiero- 
tisiert; ihr zentraler Teil terminal breit abgerundet und mit horizontalen Zähnchen- 
reihen versehen; laterale Teile in geschwungene abgerundete Spitzen auslaufend. 

Weibchen: bisher unbekannt. 

Artdiagnostische Merkmale: Die Art ist von Gasparobythiis tergestinus Poggi 
nur genitalmorphologisch einwandfrei zu unterscheiden. Während die Innensack- 
strukturen von G. tergestinus aus einem weit vorgestreckten, spitz zulaufenden und 
mit zahlreichen Zähnchen versehenen zentralen Endteil bestehen, sind diese bei G. 
kahleni sp. n. durch einen breit abgerundeten, niemals so weit herausragenden und mit 
weniger Zähnchen besetzten zentralen Endteil ausgezeichnet. Äußerlich bestehen 
zwischen den Individuen beider Arten nur ausgesprochen geringe Merkmalsunter- 
schiede, die aufgrund der geringen Zahl an bisher verfügbaren Individuen nicht mit 
Sicherheit als konstante Merkmale bewertet werden können. Die Kopfoberseite von 
G. kaheni sp. n. ist demnach ein wenig flacher und an ihren Hinterseiten geringfügig 
deutlicher punktiert als bei G. tergestinus. 

Etymologie: Benannt zu Ehren von Manfred Kahlen (Hall in Tirol), der die Art 
gesammelt hat und sie mir bereits in Kenntnis ihres Status als neue Art zur Unter- 
suchung und Beschreibung bereitgestellt hat. Kahlen gelang darüber hinaus mit dieser 
Art 1986 die Entdeckung des ersten bekannten Individuums dieser Gattung. 



1 g2 L. NEUHÀUSER-HAPPE 

Ökologie: Die bisherigen Funde deuten auf eine Lebensweise im ausgedehnten 
unterirdischen Spaltensystem hin. Nachweise dieser Art stellen ein schwieriges Unter- 
fangen dar, wobei die Gesiebeentnahme von tief in Spalten eindringenden Wurzeln 
bei entsprechend lockerem Substrat nach Kahlen (in lit.) die vermutlich geeignetste 
Sammelmethode sein dürfte. Die nah verwandte Art G. tergestinus konnte nach Poggi 
(1992) zwar bislang nur in Höhlen gefunden werden (Grotta Moser, Grotta Cosmini), 
ist jedoch vermutlich ebenso wie G. kahleni sp. n. ein typischer Spaltenbewohner. 
Das lockere und hohlraumreiche Substrat am Boden und in den Spalten der engen, 
nur auf ca. 20 Meter begehbaren Grotta Moser (Locus typicus von G. tergestinus) 
sprechen für diesen Sachverhalt. 

Verbreitung: Die Fundlokalitäten sowohl von G. tergestinus als auch G. 
kahleni sp. n. liegen in unmittelbarer Nachbarschaft zueinander und sind auf der 
Verbreitungskarte daher jeweils nur als ein Punkt dargestellt (Abb. 9). 

Tychobythinus xambeui manfredi ssp. n. 

Holotypus (c?):Tagliamento-Ufer bei Cornino, NE Udine (46°13'N, 13°00'E), 160m, 
rechtsufriger Auwald, Hochwassergenist, 23.6.1996, Manfred Kahlen leg. (TLMF). 

Die von Kahlen anlässlich der großen Hochwässer im Jahre 1996 durchge- 
führten Aufsammlungen entlang des Tagliamento führten zu dem sehr überraschen- 
den Nachweis einer neuen Unterart von Tychobythinus xambeui (Guillebeau, 1888). 
Der aktuelle Nachweis liegt mehr als 500 km vom bisher östlichsten bekannten 
Vorkommen dieser Art in den Französischen Seealpen entfernt. Bisher waren von 
Tychobythinus xambeui nur wenige Funde aus Südostfrankreich aus den Regionen 
Drôme, Vaucluse, Alpes-Maritimes bekannt (Jeannel, 1950; Poggi, 1977). Die Art 
wurde basierend auf einem weiblichen Exemplar von den Ufern der Rubion bei 
Montélimar (Drôme) beschrieben (Guillebeau, 1888; vgl. auch Ganglbauer, 1895). 
Jeannel (1950) konnte erstmals ein Männchen dieser Art untersuchen, wobei seine 
Ausführungen jedoch ungenau und fehlerhaft sind. Aufgrund des schlechten Prä- 
parationszustandes des Aedeagus des einzigen von Jeannel untersuchten Exemplars 
ist darüber hinaus auch die Beschreibung des männlichen Geschlechtsapparates 
gerade hinsichtlich seiner besonderen Auszeichnungen unvollständig geblieben. Da 
eine Identifizierung der Art anhand der bisherigen Beschreibungen daher nahezu un- 
möglich ist, werden - neben den genitalmorphologischen Unterscheidungsmöglich- 
keiten beider Unterarten - auch die wichtigsten Merkmale der Art nochmals zu- 
sammenfassend dargestellt. Für die nachstehende Kurzbeschreibung standen dem 
Autor neben dem männlichen Individuum vom Tagliamento noch drei weitere von 
Besuchet kontrollierte Belege aus Südostfrankreich ( 9 : "Inond. Rhône, Avignon II, 
Ch. Fagniez", 8 : "Drôme"; 6 : "In. Loup, A.M. X.53", alle Coll. MHNG) sowie drei 
von Poggi determinierte Individuen der nahe verwandten süditalienischen Art 
Tychobythinus gularis (Dodero, 1919) (cT & 9: "Puglia, Gioia del Colle: Grotta della 
Chiesa di M. Sannace, 24.2.1990, Angelini leg.", 6: "Basilicata, Oasi WWF Lago 
Pantano di Pignola, 14.-31.7.1991, Angelini leg.", alle Coll. MSNG) zur Verfügung. 

Beschreibung: Länge: 1,3-1,4 mm; Körper rotbraun; Flügeldecken etwas heller 
gefärbt; Behaarung normal. 



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183 





Abb. 5-8 
Dorsalansicht des Aedeagus von Gasparobythus kahleni sp. n. (5), Tychobythinus gularis 
(Dodero) (6), T. xambeui manfredi ssp. n. (7) und T. xambeui xambeui (Guillebeau) (8). 
Maßstab: 0,1 mm. 



184 



L. NEUHAUSER-HAPPE 




Abb. 9 
Bisher bekannte Fundlokalitäten von Gasparobythus kahleni sp. n. (Punkt) und G tergestinus 
Poggi (Kreis). 

Kopf: ein Viertel breiter als lang, stark und grob punktiert; der tiefe und breite 
Medianeindruck der Stirn hingegen größtenteils glatt und glänzend; Stirn halb so breit 
wie der Kopf über den Augen und auf einer Strecke von ca. einem Drittel der Kopf- 
länge scharfkantig und parallel zueinander nach hinten verlaufend, Kopf danach 
gegen die Augen erweitert; Augen bei den untersuchten Männchen aus 5-7 nicht 
immer zur Gänze pigmentierten Ommatidien bestehend (nach Jeannel bis ca. 20 
Ommatidien!); Weibchen mit normal ausgebildeten Augen (nach Guillebeau und 
Jeannel hingegen microphthalm!); Kopfunterseite der Männchen mit einer nach vorne 
gebogenen, die ganze Breite der Kopfunterseite einnehmenden Lamelle; diese an 
ihren Seiten halbkreisförmig ausgeschnitten und median mit einem dreikieligen nach 
hinten verlängerten Teil; Vorderteil der Lamelle median flach ausgerandet und in der 
Mitte mit einem nach vorne gerichteten Vorsprung auf dem ein Trichombüschel 
entspringt (Abb. 3); Maxillarpalpen: 2. Glied ventral mit 13 kleinen Tuberkeln, die 
sich größtenteils im stark erweiterten distalen Drittel befinden, an der Außenseite 
beim Männchen mit deutlichem eckigen Vorsprung und auf der Innenseite ventro- 
lateral mit oberflächlicher Eindellung, 3. Gld. mit 3-4 Tuberkeln, 4. Gld. von der 
Länge des Kopfes und von normalem Aussehen; Fühler: lang und schlank, deutlich 
länger als Kopf und Halsschild zusammen, 1 . Fühlerglied ca. 3,5 mal so lang wie 
breit, in seinem ersten Drittel kaum schmäler, 2. Gld. so breit wie das vorhergehende, 
deutlich länger als breit, 3. Gld. schmäler als das vorhergehende, ebenfalls länger als 
breit, 4. bis 8. Gld. etwa so breit wie das 3. Glied und etwa gleich lang wie breit. 

Halsschild: größtenteils glatt und glänzend; an den Seiten mit wenigen ober- 
flächlichen Punkten; zwischen Hinterrand und der basaler Halsschildfurche hingegen 
mit sehr deutlicher, dichter und grober Punktierung. 



ZWEI BYTHININI NEU FUR ITALIEN 



185 




Abb. 10 
Bisher bekannte Verbreitung von Tychobythinus xambeui xambeui (Guillebeau) (Punkte), 
T. xambeui manfredi ssp. n. (Kreis mit Punkt) und T. gularis (Dodero) (Quadrate). Die Nach- 
weise von T. gularis stammen aus Poggi (1994). 

Flügeldecken: nicht miteinander verwachsen; nur um ca. 1/10 breiter als lang; 
mit grober und tiefer Punktierung, der Abstand der Punkte zueinander ca. 2-3 mal so 
groß wie ihr Durchmesser. 

Hinterleib: glatt und glänzend. 

Flügel: vorhanden. 



186 L. NEUHÄUSER-HAPPE 

Beine: auffallend schlank; Metatibia im apicalen Drittel leicht nach innen 
gebogen. 

Aedeagus: Die asymmetrischen Innensackstrukturen sind durch einen charak- 
teristischen zurückgebogenen Sklerit ausgezeichnet. Die zwei Unterarten unter- 
scheiden sich in der Ausprägung der Innensackstrukturen und Parameren sowie in der 
Größe des Aedeagus (Abb. 7, 8). Die Paramerenenden sind bei T. xambeui manfredi 
ssp. n. demnach schmäler und länger als bei T. xambeui xambeui, die terminale Spitze 
des linken Sklerites ist im Unterschied zur Nominatunterart nach dem zurück- 
gebogenen Skleritast weniger stark verlängert und der rechte Sklerit ist nicht winkelig 
gebogen. Der Aedeagus von T. xambeui manfredi ssp. n. ist darüber hinaus größer als 
der von T. xambeui s. str. 

Verwandtschaftsverhältnisse: Im Grundbauplan des Aedeagus bestehen auf- 
fällige Übereinstimmungen mit der süditalienischen, bisher in ihren verwandt- 
schaftlichen Verhältnissen vermeintlich isoliert stehenden T. gularis (Abb. 6). Alle 
drei Taxa sind durch einen charakteristischen zurückgebogenen Sklerit ausgezeichnet. 
Unterschiede bestehen hinsichtlich der Parameren und den Innensackproportionen. 
Während bei T. xambeui s. str. der linke Sklerit der Innensackstrukturen nach seinem 
zurückgebogenen Ast stark verlängert und spitz ist, ist dieser Teil bei T. xambeui 
manfredi ssp. n. zwar ebenso spitz aber deutlich kürzer. Bei T. gularis ist dieser 
Abschnitt nur mehr sehr undeutlich als abgerundeter Fortsatz erkennbar. Der rechte 
Sklerit von T. xambeui s. str. ist im Gegensatz zu T. xambeui manfredi ssp. n. ge- 
winkelt, bei letzterer Unterart flacher gebogen, der von T. gularis ist deutlich 
gestreckter. Auch hinsichtlich der Parameren besteht eine Abfolge von breiten, schräg 
abgestutzten (T. xambeui s. str.), schmalen und langen schräg abgestutzten (T. 
xambeui manfredi ssp. n.) sowie spitz ausgezogenen Enden (T. gularis). Die große 
Ähnlichkeit der sekundären Geschlechtsmerkmale auf der Kopfunterseite der Männ- 
chen beider Arten kann als weiteres Indiz für die nahe Verwandtschaft von T. 
xambeui und T. gularis gedeutet werden (vgl. Abb. 1 in Karaman, 1959; Abb. 3). 

Etymologie: Benannt nach Manfred Kahlen (Hall in Tirol), der die Art 
gesammelt und für das Studium zur Verfügung gestellt hat. 

Angaben zur Autökologie von T. xambeui fehlen. Da beide Unterarten bislang 
nur in Hochwassergenisten gefunden wurde, bleibt der Primärlebensraum vorerst 
ungeklärt. Ihre nahe Verwandtschaft mit T. gularis, der mit einer Ausnahme (Licht- 
fang!) bisher ausschließlich in Höhlen gefunden wurde, legt die Vermutung nahe, 
dass auch von T. xambeui s. str. und T. xambeui manfredi ssp. n. - zumindest 
temporär - ähnliche Lebensräume besiedelt werden könnten. Die Kleinäugigkeit bei 
Tychobythinus spricht in erster Linie für eine verborgene Lebensweise im Spalten- 
system des Bodens. Erst weitere gezielte Nachsuchen können zur Klärung des Primär- 
lebensraumes von T. xambeui s. str. und T. xambeui manfredi ssp. n. sowie ihrer 
Verbreitung führen. Da zwischen den bekannten Vorkommen dieser Taxa weite 
Distanzen liegen (vgl. Abb. 10), ist auch die Möglichkeit des Auftretens weiterer 
Arten aus dieser Verwandtschaftsgruppe nicht auszuschließen. 



ZWEI B YTHININI NEU FÜR ITALIEN 1 87 

DANKSAGUNG 

Für kritische Anmerkungen und die Möglichkeit des Studiums von Beleg- 
material am Muséum d'histoire naturelle in Genf ergeht mein besonderer Dank and 
Dr. Claude Besuchet. Dr. Roberto Poggi danke ich für die zuvorkommende Unter- 
stützung und für wichtige Anregungen während meines Aufenthaltes am Museo 
civico di Storia naturale "G. Doria" in Genua. Für die gemeinsamen Exkursionen, die 
vielen Sammelhinweise und mitgeteilten Erfahrungen sowie für das bereitgestellte 
Material bin ich Herrn Manfred Kahlen (Hall in Tirol) zu großem Dank verpflichtet. 

LITERATUR 

Besuchet, C. 1985. Bythinini cavernicoles nouveaux de France et d'Espagne (Coleoptera: 

Pselaphidae). Revue suisse de Zoologie 92: 509-517. 
Ganglbauer, K. 1895. Die Käfer von Mitteleuropa. Zweiter Band. Familienreihe Staphy- 

linoidea. 1. Theil: Staphylinidae, Pselaphidae. Carl Gerold' s Sohn, Wien, VI + 881 pp. 
Guillebeau, F. 1888. Notes pour servir à l'étude de Psélaphiens. Revue d'Entomologie 7: 203- 

220, 368-380. 
Jeannel, R. 1950. Faune de France. 53. Coléoptères Psélaphides. Paul Lechevalier, Paris, 

421 pp. 
Karaman, Z. 1959. Su due nuovi Bitinini italiani (Coleotteri Pselafidi). Le Grotte d'Italia, 

serie 3.2: 71-75. 
Poggi, R. 1977. Studio segli Pselaphidae della Liguria (Coleoptera). Memorie della Società 

Entomologica Italiana 55: 1 1-100. 
Poggi, R. 1992. Forme nuove o poco note di Pselaphidae cavernicoli del Friuli-Venezia Giulia 

e della Jugoslavia (Coleoptera). Memorie della Società Entomologica Italiana 70: 207- 

224. 

Poggi, R. 1994. Appunti sinonimici su Tychobythinus gularis (Dod.) e T. anelili (Kar.) 
(Coleoptera Pselaphidae). Bolletino della Società Entomologica Italiana, Genova 126: 
141-144. 



Revue suisse de Zoologie 109 (1): 189-240; mars 2002 



Nuovi generi di Aleocharinae del Borneo 
(Coleoptera, Staphylinidae)* 

Roberto PACE 

Via Vittorio Veneto, 13, 1-37032 Monteforte d' Alpone (Verona), Italia. 



New genera of Aleocharinae from Borneo (Coleoptera, Staphyli- 
nidae). - Twenty-three genera and 35 species are described as new. The 
genera are distributed in 11 tribes. Hypocyphtini: Akanthoystera n. gen. 
(4 new species); Gyrophaenini: Mesophaena n. gen. (In. sp.); Homa- 
lotini: Psephothetemusa n. gen. (2 n. spp.), Aistenthusa n. gen. (4 n. spp.), 
Metechonica n. gen. (2 n. spp.), Megaparaglossa n. gen. (In. sp.), 
Apatelomixidota n. gen. (3 n. spp.), Episkilepta n. gen. (1 n. sp.); Diesto- 
tini: Anamignusa n. gen. (In. sp.); Bolitocharini: Antithetusa n. gen. (In. 
sp.), Panbrachyna n. gen. (1 n. sp.); Falagriini: Borneopora n. gen. (In. 
sp.); Athetini: Planadota n. gen. for Atheta borneensis Cameron, Para- 
nomusa n. gen. (1 n. sp.), Dikraspedella n. sp. (2 n. spp.), Trigonoglossa 
n. gen. (In. sp.), Serikasomina n. gen. (3 n. spp.), Ektasitrachela n. gen. 
(In. sp.); Thamiaraeini: Diabainella n. gen. (In. sp.); Lomechusini: 
Borneozyras n. gen. (1 n. sp.); Hoplandriini: Borneusa n. gen. (In. sp.); 
Oxypodini: Apatelieida n. gen. for Neosilusa stricticollis Cameron, Syn- 
temusa n. gen. (2 n. spp.). New combinations are proposed for two 
species: Atheta borneensis Cameron and Neosilusa stricticollis Cameron. 
Keys to new genera and species are provided. Each new genus and 
species is described and illustrated. 

Key-words: Coleoptera - Staphylinidae - Aleocharinae - new genera and 
species - taxonomy - Borneo. 



INTRODUZIONE 

Le accurate e ripetute ricerche effettuate sul Monte Kinabalu (Borneo, Sabah) 
nel corso di spedizioni zoologiche del Dr. Ales Smetana del "Centre for Land and 
Biological Resources Research" di Ottawa, del Dr. Daniel Burckhardt e del Dr. Ivan 
Lobi, entrambi del Museo di Storia Naturale di Ginevra, hanno riportato per la 
conservazione in questo Museo, un importantissimo, abbondante e talvolta inatteso 
materiale di Aleocharinae. Affidatomi in esame, mi ha permesso di riconoscere come 
nuovi per la Scienza ben 23 generi. La loro descrizione è lo scopo del presente lavoro. 



164° Contributo alla conoscenza delle Aleocharinae. 
Manoscritto accettato il 26.03.2001 



190 R - PACE 

Il sospetto che anche in materiale di Aleocharinae diverso dal termitofilo, 
questo già studiato e pubblicato da Kistner tra il 1969 e il 1976 (vedi bibliografia in 
Hammond, 1984), fosse possibile scoprire nuovi generi per la Scienza, mi venne 
allorché esaminai i tipi di Leptusa borneensis Cameron, 1933, da me riconosciuta 
appartenere al nuovo genere Mimopisalia Pace, 1984. E' seguito, nel corso dell'esame 
di tipi di altre specie di Aleocharinae del Borneo, il riconoscimento e la pubblicazione 
del nuovo genere Borneoxenia Pace, 1993, per Atheta (Aerofono) mjoebergi 
Cameron, 1928. Il fatto che Cameron non abbia riconosciuto i nuovi generi, è che 
questo autore, in presenza di esemplari di taglia troppo ridotta, non procedeva 
all'esame dei caratteri generici e si affidava al solo aspetto dell'habitus. Purtroppo le 
convergenze morfologiche relative a questo rendono ingannevole la comprensione di 
ciò che si ha sotto gli obiettivi del microscopio. E' questo anche il caso della specie 
Neosilusa stricticollis Cameron, 1943, che non solo va attribuita a nuovo genere, ma 
anche a tribù differente (da Homalotini a Oxypodini). 

Anche Sawada (1980) si era reso conto che alcune specie di Aleocharinae del 
Borneo descritte da Cameron e altri autori, non appartenevano al genere loro attribuito 
e talvolta andavano assegnati a generi inediti. 

Gli olotipi delle nuove specie sono conservati nel Museo di Storia Naturale di 
Ginevra (MHNG). Due olotipi provenienti dalle raccolte di Guillaume de Rougemont e 
Assing sono conservati nel Museo Regionale di Scienze Naturali di Torino (MRSN). 
Paratipi si conservano nei medesimi Musei e in collezione V. Assing, Hannover. 

METODO 

Kraatz (1856, 1857) fu tra i primi stafilinidologi che compresero l'importanza 
dell'esame delle parti boccali anche per le specie della sottofamiglia Aleocharinae, al 
fine di individuare o delimitare i generi. Se l'habitus spesso uniforme, indurrebbe a 
credere a una uniformità di nicchie ecologiche, ci si trova invece di fronte a Stafilinidi 
che occupano un gran numero di nicchie ecologiche differenti, anche non note. Si 
trovano specie fungicole, ripicole, cacciatrici di larve di Ditteri o di Coleotteri. Per- 
tanto l'anatomia delle parti boccali è soggetta a differenziazioni morfologiche impor- 
tanti, secondo la nicchia ecologica occupata. La forma della ligula e parti annesse 
pertanto assumono un'importanza basilare al fine del riconoscimento dei vari generi. 
Nel presente lavoro però, nel delineare i generi, non sono stati trascurati i minuti 
caratteri dell'habitus o il tipo di edeago e di spermateca. Questi due organi presentano 
caratteri tali da essere decisivi nel delimitare i generi, quando si è in presenza di 
possibili convergenze morfologiche delle parti boccali o dell'habitus. 

Di ogni genere qui descritto si dà una breve diagnosi e occasionalmente una 
chiave dei generi affini. Per i generi politipici si dà una chiave delle specie. 

Tribù HYPOCYPHTINI Laporte, 1835 (= Oligotini) 

Akanthoystera gen. n. Figg. 1-19 

Diagnosi. Per le antenne di 10 antennomeri (uno vestigiale sta alla base del 
primo antennomero), il corpo minuto e fusiforme, il nuovo genere appare affine al 



ALEOCHARINAE DEL BORNEO 191 

genere Oligota Mannerheim, 1831. Tuttavia per la presenza di formula tarsale 4-4-5-, 
invece di 4-4-4, va attribuito provvisoriamente a questa tribù. 

Descrizione. Taglia corporea e habitus di molte specie del genere Oligota 
Mannerheim; antenne di 10 antennomeri, uno vestigiale supplementare sta alla base; 
antennomeri 4 a 9 trasversi, fig. 10; tempie robustamente marginate; collo stretto; 
palpi labiali di 2 articoli poco allungati; ligula intera e lunga, fig. 6; palpi mascellari di 
4 articoli, di cui il terzo assai lungo e stretto, fig. 7; mento trapezoidale, con margine 
anteriore incavato, fig. 5; mesosterno carenato alla base; processo mesosternale acuto; 
mesocoxe contigue tra loro; addome fortemente ristretto posteriormente e ornato di 
robuste setole lunghe; formula tarsale 4-4-5; primo tarsomero posteriore lungo quanto 
i due tarsomeri seguenti riuniti; edeago fortemente strozzato a metà, figg. 2-3, 11-12, 
14-15 e 18-19; bulbo distale della spermateca con appendice apicale ricurva, figg. 4, 
9 e 16. 

Tipo del genere. Akanthoystera smetanai sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Irta posterior- 
mente". 

Comparazioni. Il nuovo genere si distingue per i caratteri dati nella seguente 
chiave. 

1. Formula tarsale 4-4-4; ligula divisa in due lobi; mesocoxe separate 

Oligota Mannerheim, 1831; Liophaena Sharp, 1880; 

Nematoscelis Wollaston, 1 867 
Formula tarsale 4-4-5; ligula intera; mesocoxe contigue Akanthoystera gen. n. 

Akanthoystera smetanai sp. n. Figg- 1-7 

Holotypus ó\ Borneo, Sabah, Mt. Kinabalu N.P., summit tr. Pondok Lowii, 2300-2400 
m. 28.IV. 1987, A. Smetana leg. (MHNG). 

Paratypi: 1 9, stessa provenienza; 4 es., Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot 
Springs, 500 m, 7.V.1987, Burckhardt & Lobi leg.; 1 es., Borneo, Sabah, Mt. Kinabalu N.P., 
Poring Hot Springs, 500 m, 11.V.1987, Burckhardt & Lobi leg.; 2 es., Borneo, Sabah, Mt. 
Kinabalu N.P., Poring Hot Springs, 550-600 m, 9. V. 1987, Burckhardt & Lobi leg. 

Descrizione. Lungh. 1,4 mm. Corpo lucido e rossiccio con elitre e una fascia 
addominale brune; antenne giallo-brune con i tre antennomeri basali giallo-rossicci; 
zampe gialle. Non è presente reticolazione sulla superficie corporea. La granulosità 
del capo è distinta, quella del pronoto e delle elitre è saliente. Addome coperto di 
rughe longitudinali. Edeago figg 2-3, spermateca fig. 4. 

Etimologia. Specie dedicata a uno dei suoi raccoglitori, il Dr. Ales Smetana 
del "Centre for Land and Biological Resources Research" di Ottawa. 

Akanthoystera punctum sp. n. Figg- 8-12 

Holotypus ó\ Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs. 510 m, 
30.VIII.1988, A. Smetana leg. (MHNG). 

Paratypus: 1 9 , stessa provenienza. 

Descrizione. Lungh. 1,1 mm. Corpo lucido e giallo-rossiccio, con elitre 

rossicce e i quattro uriti basali bruno-rossicci; antenne giallo-brune con i tre anten- 



192 



R. PACE 




FlGG. 1-12 

Habitus, edeago in visione laterale e ventrale, spermateca, mento, labio con palpo labiale, 
maxilla con palpo mascellare e antenna. 1-7: Acanthoystera smetanai gen. n., sp. n.; 8-12: 
Acanthoystera punctum gen. n., sp. n. 



ALEOCHARINAE DEL BORNEO 193 

nomeri basali gialli; zampe gialle. La reticolazione del capo è assente, quella del 
pronoto e delle elitre è estremamente superficiale. L'addome mostra dei granuli 
allungati. Il capo è privo di punteggiatura o di granulosità. Sul pronoto sono presenti 
quattro grossi granuli a ciascun lato della base: sul resto del pronoto i granuli sono 
fini. La granulosità delle elitre è molto superficiale e fine, L'addome mostra granuli 
allungati. Spermateca fig. 9, antenna fig. 10, edeago figg. 11-12. 

Etimologia. La nuova specie, prende nome di punctum (=punto), perché pre- 
senta un corpo piccolissimo, simile alla forma di un punto. 

Akanthoystera kinabaluensis sp. n. Figg. 13-16 

Holotypus 9, Borneo. Sabah, ML Kinabalu N.P.. 1550 m, 29.IV. 1987, Burckhardt & 
Lobi leg. (MHNG). 

Paratypi: 6 es., stessa provenienza, ma 27.IV. 1987; 5 es., Borneo, Sabah, Mt. Kinabalu 
N.P.. Poring Hot Springs, nr. Bat Cave, 600 m, 10.V.1987, Burckhardt & Lobi leg. 

Descrizione. Lungh. 1,3 mm. Corpo lucido e rossiccio; antenne rossicce con i 
tre antennomeri basali gialli; zampe gialle. Il corpo non è reticolato. Il capo è privo di 
punteggiatura. La granulosità del pronoto e delle elitre è grossolana e fitta e sulle 
elitre in più è allungata, come quella dell'addome. Edeago figg. 14-15, spermateca 
fig. 16. 

Akanthoystera mìnima sp. n. Figg- 17-19 

Holotypus (5, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 500 m, 
1 I.V. 1987, Burckhardt & Lobi leg. (MHNG). 

Descrizione. Lungh. 1,1 mm. Corpo lucido e giallo-rossiccio, con elitre e 
addome giallo-bruni; antenne di un giallo sporco alla base, resto delle antenne per- 
duto. La reticolazione del capo è assente, quella del pronoto e delle elitre è estre- 
mamente superficiale. La punteggiatura del capo è molto svanita. La granulosità del 
pronoto è fine e saliente sulla metà posteriore e superficiale sull'anteriore, quella delle 
elitre è saliente. Una rugosità allungata copre gli uroterghi. Edeago figg. 18-19. 

Comparazioni. Chiave delle specie del nuovo genere Akanthoystera. 

1 Pronoto coperto di granulosità forte e distribuita su tutta la sua superficie ... 2 
Pronoto coperto di granulosità fine: solo alcuni granuli posteriori sono forti . 3 

2 Corpo uniformemente rossiccio; edeago senza lamina ventrale, con uno 
spigolo ottuso al suo posto; appendice del bulbo distale della sper- 
mateca robusto. Lungh. 1,3 mm. M. Kinabalu A. kinabaluensis sp. n. 

Corpo rossiccio, con elitre e una fascia addominale brune; edeago con 
lamina ventrale; appendice del bulbo distale della spermateca esile. 
Lungh. 1,4 mm. M. Kinabalu A. smetanai sp. n. 

3 Capo, pronoto e pigidio giallo-rossicci; pronoto con una fila posteriore 
di granuli forti; edeago più robusto e più largo, in visione ventrale. 

Lungh. 1,1 mm. M: Kinabalu A. punctum sp. n. 

Capo e pronoto rossicci; pigidio giallo-bruno; metà posteriore del 
pronoto con granuli più salienti di quelli della metà anteriore; edeago 
esile. Lungh. 1,1 mm. M. Kinabalu A. minima sp. n. 



194 R - PACE 

Tribù GYROPHAENINI Kraatz. 1856 

Mesophaena gen. n. Figg. 20-24 

Diagnosi. Genere sicuramente appartenente alla tribù Gyrophaenini, per la 
presenza di corti denti in più file all'apice della lacinia. E' simile al genere Adelarthra 
Cameron, 1920, ma nettamente distinto per la presenza di una ligula intera (divisa in 
Adelarthra). 

Descrizione. Undici antennomeri. Corpo fusiforme, con robuste setole laterali 
delle elitre e dell'addome. Tempie robustamente marginate; collo largo; palpi labiali 
di due articoli; ligula intera, larga e corta, fig. 21; apice della lacinia con corti e 
robusti denti in più file; mesosterno carenato alla base; mesocoxe distanziate fra loro; 
formula tarsale 4-4-5; primo tarsomero posteriore corto; edeago figg 22-23, 
spermateca fig. 24. 

Tipo del genere. Mesophaena fragilis sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Manifestazione 
intermedia". Infatti sembra che il nuovo genere occupi posizione intermedia tra il 
genere Adelarthra e Pseudoligota Cameron, 1920. 

Comparazioni. Chiave dei generi affini al nuovo genere Mesophaena. 

1 Ligula profondamente divisa in due parti 2 

Ligula intera Mesophaena gen. n. 

2 Ligula divisa in due larghi lobi Adelarthra Cameron, 1920 

Ligula divisa in due lobi stretti e acuti 3 

3 Mesosterno carenato e diviso dal metasterno .... Sternotropa Cameron, 1920 
Mesosterno non carenato, fuso con il metasterno Pseudoligota Cameron, 1920 

Mesophaena fragilis sp. n. Figg. 20-24 

Holotypus a, Borneo, Sabah, Mt. Kinabalu N.P.. Poring Hot Springs, 500 m, 6.V.1987, 
Burckhardt & Lobi leg. (MHNG). 

Paratypi: 1 â, stessa provenienza; 1 9, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot 
Springs, 500 m, 1 I.V. 1987, Burckhardt & Lobi leg. (MHNG). 

Descrizione. Lungh. 1,4 mm. Corpo lucido e giallo rossiccio; antenne giallo- 
rossicce con base e undicesimo antennomero gialli; zampe gialle. La reticolazione del 
capo è assente, quella del pronoto è estremamente superficiale e quella delle elitre è 
distinta. La distinta granulosità del pronoto è piuttosto rada, quella delle elitre è 
saliente. Edeago figg. 22-23, spermateca fig. 24. 

Tribù HOMALOTINI Heer, 1839 

Psephothetemusa gen. n. Figg. 25-33 

Diagnosi. I denti della lacinia indicano che il nuovo genere non appartiene alla 
tribù Gyrophaenini. nonostante che l'edeago e l'habitus indichino una forte affinità 
con i generi di questa tribù. Per la forma della ligula, mai osservata nei generi della 
tribù Homalotini è genere nuovo per la Scienza. 



ALEOCHARINAE DEL BORNEO 



195 




FlGG. 13-24 

Habitus, edeago in visione laterale e ventrale, spermateca e mento. 13-16: Acanthoystera 
kinabaluensis gen. n., sp. n.; 17-19: Acanthoystera minima gen. n., sp. n.; 20-24: Mesophaena 
fragilis gen. n., sp. n. 



196 R - PACE 

Descrizione. Undici antennomeri; tempie marginate; palpi labiali di 2 articoli, 
non stiliformi; ligula intera, con una papilla apicale; paraglosse non sporgenti, fig. 28; 
palpi mascellari di 4 articoli; estremità della galea con lunghe setole; lacinia con una 
fila di spine interne, fig. 30; mento trapezoidale, con margine anteriore retto, fig. 29; 
mesosterno non carenato; processo mesosternale acuto; mesocoxe contigue; formula 
tarsale 4-4-5; primo tarsomero posteriore allungato; edeago figg. 26-27, spermateca 
fig. 32. 

Tipo del genere. Psephothetemusa problematica sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Colei che è un 
mosaico". Infatti il nuovo genere mostra caratteri che si riscontrano in altri generi, 
anche di tribù differenti, tanto da sembrare un mosaico di caratteri presi a prestito. 

Psephothetemusa problematica sp. n. Figg. 25-31 

Holotypus ó\ Borneo, Sabah, Mt. Kinabalu N.P., HQ Silau-Silau trail, 1560 m, 
23.V.1987, A. Smetana leg. (MHNG). 

Descrizione. Lungh. 1,0 mm. Corpo lucido e bruno-rossiccio, con elitre brune; 
antenne brune con base del terzo antennomero giallo chiaro; zampe bruno-rossicce. 
La reticolazione del capo è assente, quella del pronoto e delle elitre è distinta. La 
punteggiatura del capo è distinta assai superficiale. La granulosità del pronoto e delle 
elitre sono molto salienti e forti. Edeago figg. 26-27. 

Psephothetemusa introflexa sp. n. Figg. 32-33 

Holotypus 9, Borneo, Sabah, Mt. Kinabalu N.P., 1580 m, 27.IV. 1987, Burckhardt & 
Lobi leg. (MHNG). 

Descrizione. Lungh. 1,4 mm. Corpo lucido e bruno-rossiccio; antenne brune 
con gli antennomeri secondo e terzo giallo chiaro; zampe giallo-brune con tarsi gialli. 
La reticolazione del capo e del pronoto è distinta, quella delle elitre è superficiale e 
quella dell'addome è assente. La punteggiatura del capo è piuttosto fitta e 
superficiale. La granulosità del pronoto è forte e saliente, quella delle elitre è distinta. 
Gli uroterghi liberi terzo e quarto presentano striatura longitudinale superficiale, il 
quinto la presenta netta. Spermateca fig. 32. 

Comparazioni. Chiave delle specie del nuovo genere Psephothetemusa. 

1 Primo e secondo antennomero basale bruni; capo senza reticolazione, 

poco più stretto del pronoto; reticolazione delle elitre distinta. Lungh. 

1,1 mm. M. Kinabalu P. problematica sp. n. 

Primo antennomero basale bruno, secondo giallo pallido; capo distin- 
tamente reticolato e molto più stretto del pronoto; reticolazione delle 
elitre superficiale. Lungh. 1,4 mm. M. Kinabalu P. introflexa sp. n. 

Aisthentusa gen. n. Figg. 34-51 

Diagnosi. Per la ligula intera e corta, il nuovo genere sembra affine al genere 
Placusa Erichson, 1837, ma le mesocoxe sono tra loro separate e il mesosterno è 
carenato. 



ALEOCHARINAE DEL BORNEO 



197 




FlGG. 25-33 

Habitus, edeago in visione laterale e ventrale, labio con palpo labiale, mento, maxilla con palpo 
mascellare, sesto urotergo libero del 6, spermateca. 25-31: Psephothetemusa problematica 
gen. n., sp. n.; 32-33: Psephothetemusa introflexa gen. n., sp. n. 



Descrizione. Undici antennomeri; lati del corpo con lunghe e robuste setole 
isolate; tempie marginate; palpi labiali di 2 articoli piuttosto corti, fig. 40; palpi 
mascellari di 4 articoli, di cui il terzo è molto tozzo e il quinto è sottile e molto più 
lungo del terzo, fig. 39; mesosterno carenato, ma non alla base; mesocoxe separate fra 
loro; uroterghi fortemente striati longitudinalmente; formula tarsale 4-4-5; primo 
tarsomero posteriore allungato; edeago con un lobo preapicale distale, figg. 35, 41 e 
47, spermateca fig. 37. 



198 R - PACE 

Tipo del genere. Aisthentusa bomeensis sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Colei che è 
sensitiva" a motivo della presenza di lunghe setole laterali del corpo e di lunghe spine 
al margine posteriore del sesto urotergo libero del 6 . 

Comparazioni. Chiave del nuovo genere Aisthentusa e del genere Placusa. 

1 Ligula intera a lati arrotondati; terzo articolo dei palpi mascellari più 

lungo del secondo e debolmente rigonfio; quarto articolo dei palpi 
mascellari lungo circa la metà del terzo; pronoto pubescente; mesocoxe 

contigue Placusa Erichson, 1837 

Ligula intera a lati acuti; terzo articolo dei palpi mascellari più corto del 
secondo e fortemente rigonfio; quarto articolo dei palpi mascellari 
nettamente più lungo del terzo; pronoto nudo; mesocoxe fra loro 
separate Aisthentusa gen. n. 

Aisthentusa bomeensis sp. n. Figg. 34-40 

Holotypus 8, Borneo, Sabah, Mt. Kinabalu N.P.. below Layang Layang, 2590 m, 
1 .V. 1 987, A. Smetana leg. (MHNG). 

Paratypi: 1 <3\ stessa provenienza; 19 es., Borneo, Sabah, Mt. Kinabalu N.P., 2600 m, 
1. IV. 1987, Burckhardt & Lobi leg. 

Descrizione. Lungh. 2,1 mm. Corpo lucido e bruno; antenne brune con i 
quattro antennomeri basali giallo-rossicci; zampe giallo-brune con tarsi rossicci. La 
reticolazione del capo e dell'addome è assente, quella del pronoto e delle elitre è 
distinta, composta di maglie trasverse e ondulate. Il capo e il pronoto sono senza 
punteggiatura. Le elitre sono coperte di microgranuli sparsi e distinti. 

Aisthentusa hystrix sp. n. Figg. 41-45 

Holotypus S, Borneo, Sabah, Mt. Kinabalu N.P.. 29. X. 1990, G. de Rougemont leg. 
(MRSN). 

Paratypi: 4 es. Borneo, Sabah, Mt. Kinabalu N.P., 1550-1650 m, 24.IV. 1987, 

Burckhardt & Lobi lea.; 2 es.. Borneo, Sabah, Mt. Kinabalu N.P.. 1500 m. 25.IV. 1987, 

Burckhardt & Lobi leg.; 1 S. Borneo, Sabah, Mt. Kinabalu N.P., 1580 m, 27.IV. 1987, 

Burckhardt & Lobi leg.; 1 es. Borneo, Sabah, Mt. Kinabalu N.P.. 2600 m, 28.IV. 1987, 

Burckhardt & Lobi leg.; 2 Sa, Borneo, Sabah, Mt. Kinabalu N.P., 1550 m, 29.IV.1987, 
Burckhardt & Lobi leg. 

Descrizione. Lungh. 1,8 mm. Corpo lucidissimo e bruno, con addome bruno- 
rossiccio, avente gli uriti liberi 3°, 4° e 5° bruni; antenne rossicce con i tre antenno- 
meri basali gialli e gli antennomeri 9°, 10° e 11° giallo-bruni; zampe rossicce. La 
reticolazione è presente solo sulle elitre dove è superficiale, composta di maglie molto 
trasverse e ondulate. Edeago figg. 41-42, sesto urotergo libero del S fig. 44, sper- 
mateca fig. 45. 

Aisthentusa pacifica sp. n. Figg. 46-49 

Holotypus 6, Borneo. Sabah. Mt. Kinabalu N.P.. 1550 m, 29.IV. 1987, Burckhardt & 
Lobi leg. (MHNG). 



ALEOCHARINAE DEL BORNEO 



199 




FlGG. 34-42 

Habitus, edeago in visione laterale e ventrale, spermateca, sesto urotergo libero del 6 , labio 
con palpo labiale, maxilla con palpo mascellare. 34-40: Aisthentusa borneensis gen. n., sp. n.; 
41-42: Aisthentusa hystrix gen. n., sp. n. 



200 R - PACE 

Descrizione. Lungh. 1,4 mm. Corpo lucido e nero-bruno; antenne nero-brune 
con i due antennomeri basali di un giallo sporco. La reticolazione del capo è assente, 
quella del pronoto è estremamente superficiale e quella delle elitre è distinta, 
composta di maglie ondulate trasverse. Il capo è senza punteggiatura. Le elitre 
mostrano alcuni distinti granuli sparsi. Edeago figg. 47-48, sesto urotergo libero del 3 
fig. 49. 

Aisihentusa intermedia sp. n. Figg. 50-51 

Holotypus 9, Borneo, Sabah. Crocker Ra.. 1550-1650 m, 16.V.1987. Burckhardt & 
Lobi leg. (MHNG). 

Paratypi: 3 9 9, stessa provenienza; 1 9, Borneo, Sabah, Mt. Kinabalu N.P., Poring 
Hot Springs, 500 m, 1 1 .V. 1 987, Burckhardt & Lobi leg. 

Descrizione. Lungh. 1,4 mm. Corpo lucido e nero-bruno; antenne brune con i 
due antennomeri basali di un giallo sporco; zampe brune con tarsi rossicci. Il capo è 
senza reticolazione e senza punteggiatura. La reticolazione del pronoto è presente 
solo sul disco dove è estremamente superficiale. La punteggiatura delle elitre è netta. 
Spermateca fig. 50. 

Comparazioni. Chiave delle specie del nuovo genere Aisthentusa. 

1 Taglia maggiore, 2,1 mm; pronoto con reticolazione ondulata trasversa 
distinta; lobo preapicale dell'edeago poco sviluppato. M. Kinabalu 

A. bomeensis sp. n. 

Taglia minore, 1,4-1,8 mm; pronoto senza reticolazione o con retico- 
lazione estremamente svanita; lobo preapicale dell' edeago mediamente 

o molto sviluppato 2 

2 Antennomeri 4° a 10° trasversi. Lungh. 1,4 mm A. intermedia sp. n. 

Antennomeri 4° a 10° lunghi quanto larghi o più lunghi che larghi 3 

3 Taglia corporea maggiore: 1,8 mm; base dell'addome e pigidio rossicci; 
pronoto fortemente trasverso; spine esterne del 6° urotergo libero del S 
lunghissime e robuste; lobo preapicale dell' edeago poco sviluppato. 

M. Kinabalu A. hystrix sp. n. 

Taglia corporea minore: 1,4 mm; addome uniformemente nero-bruno; 
pronoto meno trasverso; spine esterne del 6° urotergo libero del 6 corte 
ed esili; lobo preapicale dell'edeago enormemente sviluppato. M. Kina- 
balu A. pacifica sp. n. 

Metechonica gen. n. Figg- 52-60 

Diagnosi. Habitus del genere Stenomastax Cameron, 1933, ma gli articoli dei 
palpi labiali non sono stiliformi e la ligula termina con due larghi lobi. 

Descrizione. Undici antennomeri; corpo fittamente pubescente; tempie robus- 
tamente marginate; palpi labiali di due articoli, fig. 55; palpi mascellari di 4 articoli; 
mento trapezoidale, con margine anteriore incavato fig. 56; processo mesosternale 
acuto; mesocoxe contigue tra loro; formula tarsale 4-4-5; primo tarsomero posteriore 
corto. 



ALF.OCÏIARINAH DHL BORNI-IO 



201 




Figo. 43-51 
Habitus, sesto urotergo libero del S , edeago in visione laterale e ventrale e spermateca. 43-45: 
Aisthentusa hystrix gen. n., sp. n.; 46-49: Aisthentusa pacifica gen. n., sp. n.; 50-5 1 : Aisthentusa 
intermedia gen. n., sp. n. 



Tipo del genere. Metechonica nova sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Immagine 
partecipe", nel senso che il nuovo genere mostra dei caratteri che fanno parte anche 
del genere Stenomastax. 

Comparazioni. Chiave dei generi affini al nuovo genere Metechonica. 



202 R - PACE 

1 Palpi labiali stiliformi 2 

Palpi labiali non stiliformi, corti 4 

2 Ligula divisa all'estremità Stenomasiax Cameron 1933 

Ligula intera 3 

3 Ligula che in lunghezza raggiunge al massimo il livello del primo 

articolo dei palpi labiali Silusa Erichson 1837 

Ligula che in lunghezza supera il livello del primo articolo dei palpi 
labiali Taraktmora Pace, 1998 

4 Ligula corta, essa raggiunge il livello del primo articolo dei palpi labiali 

Coenonica Kraatz, 1 857 

Ligula lunghissima, essa raggiunge l' apice del secondo articolo dei 

palpi labiali Meiechonica gen. n. 

Metechonica nova sp. n. Figg. 52-56 

Holotypus S. Borneo, Sabah, Mt. Kinabalu N.P., HQ Liwagu Riv., 1490 m, 
5.VIII.1988, A. Smetana leg. (MHNG). 

Descrizione. Lungh. 2,1 mm. Corpo lucido e giallo-rossiccio, con capo ed 
elitre bruno-rossicci; antenne rossicce con i due antennomeri basali e l'undicesimo 
giallo-rossicci; zampe gialle. La reticolazione del capo e delle elitre è superficiale, 
quella del pronoto è distinta e quella dell'addome è assente. Sul capo e sul pronoto 
non sono visibili né punteggiatura, né granulosità. La punteggiatura delle elitre è 
distinta, fine e fitta. Edeago figg. 53-54. 

Metechonica temburongensis sp. n. Figg- 57-60 

Holotypus <?, Borneo, Brunei. Temburong, Kuala Belalong (West), 11.11.1995, 
Borcherding leg. (MRSN). 

Paratypi: 2 9 9, stessa provenienza; 10 es., Borneo, Sabah, Mt. Kinabalu N.P., Poring 
Hot Springs, 480 m, 8.V.1987. A. Smetana leg.; 1 9, Borneo. Sabah, Mt. Kinabalu N.P., 
Poring Hot Springs, 550-600 m, 9. V. 1987. Burckhardt & Lobi leg. 

Descrizione. Lungh. 2,0 mm. Avancorpo debolmente lucido, addome lucido. 
Capo bruno, pronoto rossiccio, elitre brune con base rossiccia, addome giallo- 
rossiccio; antenne rossicce con i due antennomeri basali e apice dell'undicesimo 
giallo-rossicci; zampe gialle. La reticolazione del capo e del pronoto è distinta, quella 
delle elitre e dell'addome è superficiale. La granulosità del capo e del pronoto è 
confusa nella reticolazione, quella delle elitre è fine, fitta e distinta e quella 
dell'addome è saliente a raspa. Edeago figg. 58-59, spermateca fig. 60. 

Comparazioni. Chiave delle specie del nuovo genere Metechonica. 

1 Elitre interamente bruno-rossicce; penultimi antennomeri lunghi quanto 
larghi; edeago maggiore, meno profondamente ricurvo e con lunghis- 
simo pezzo copulatore interno. Lungh. 2,1 mm. M. Kinabalu . . M. nova sp. n. 
Elitre brune, con base rossiccia; penultimi antennomeri nettamente 
trasversi; edeago minore, profondamente ricurvo e con profondo pezzo 
copulatore interno. Lungh. 2,0 mm. Brunei M. temburongensis sp. n. 



ALEOCHARINAE DEL BORNEO 



203 




Figo. 52-60 
Habitus, edeago in visione laterale e ventrale, labio con palpo labiale, mento e spermateca. 
52-56: Metechonica nova gen. n., sp. n.; 57-60: Metechonica temburongnsis gen. n., sp. n. 



204 R. pace 



Megaparaglossa gen. n. Figg. 61-67 

Diagnosi. Per l'enorme sviluppo delle paraglosse, il nuovo genere risulta 
isolato, dato che l'habitus è "normale", cioè non ultraevoluto, come si riscontra nelle 
specie di Aleocharinae termitofile o mirmecofile. Queste paraglosse oltremodo 
sviluppate si riscontrano in certi generi termitofili o mirmecofili. 

Descrizione. Undici antennomeri; addome lievemente ristretto all'indietro; 
tempie marginate; palpi labiali di 2 articoli; ligula cortissima e incisa all'apice, quasi 
nascosta tra le paraglosse enormemente sviluppate e molto spinte in avanti, fig. 65; 
palpi mascellari di 4 articoli; galea ampiamente setolosa all'apice; denti della lacinia 
lunghissimi, fig. 66; mento trapezoidale, con margine anteriore retto, fig. 67; 
mesosterno non carenato; processo mesosternale acuto, mesocoxe appena distanziate 
fra loro; formula tarsale 4-4-5; primo tarsomero posteriore corto; edeago figg. 62-63, 
spermateca fig. 64. 

Tipo del genere. Megaparaglossa annularis sp. n. 

Etimologia. Il nome femminile della nuova specie significa "Grande para- 
glossa". 

Megaparaglossa annularis sp. n. Figg- 61-67 

Holotypus o\ Borneo, Sabah, Mt. Kinabalu, 1750 m, 21. IV. 1987, Burckhardt & Lobi 
leg. (MHNG). 

Paratypi: 1 S e 1 9, stessa provenienza; 2 SS, Borneo, Sabah, Mt. Kinabalu N.P., 
Poring Hot Springs, 550 m, 7.V.1987, Burckhardt & Lobi leg. 

Descrizione. Lungh. 1,7 mm. Corpo lucido e giallo-rossiccio, con capo ros- 
siccio, elitre brune con base giallo-rossiccia e una fascia addominale bruna; antenne 
brune con i tre antennomeri basali gialli; zampe gialle. La reticolazione del capo è 
estremamente superficiale, quella del pronoto è assente e quella delle elitre è svanita. 
La punteggiatura del capo è fine e superficiale. La granulosità del pronoto è poco 
saliente, quella delle elitre è netta. Edeago figg. 62-63, spermateca fig. 64. 

Apatelomixidota gen. n. Figg. 68-84 

Diagnosi. In base all'habitus e alla forma della spermateca, il nuovo genere 
mostra affinità con varie specie del sottogenere Datomicra Mulsant & Rey, 1874 di 
Aiheta Thomson, 1859. Ma la formula tarsale è 4-4-5 (invece di 4-5-5), gli articoli del 
palpi labiali sono 2 e la ligula di forma finora mai osservata. 

Descrizione. Undici antennomeri; corpo simile ad Atheta Thomson; tempie 
marginate; palpi labiali di 2 articoli, tra essi vi è un articolo vestigiale; ligula divisa 
all'apice in due rami, ciascuno dei quali porta un lobo, fig. 73; palpi mascellari di 4 
articoli; apice della galea molto lungo e coperto di setole cortissime, fig. 74; mento 
fig. 72; mesosterno non carenato; processo mesosternale acuto; mesocoxe tra loro 
contigue; formula tarsale 4-4-5; primo tarsomero posteriore lungo quanto i due 
seguenti riuniti; edeago figg. 69-70, spermateca fig. 71. 

Tipo del genere. Apatelomixidota borneensis sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Colei che dà una 
mescolanza ingannevole". Mescolanza è in riferimento ai caratteri di altri generi che 



AL.HOt'HARINAH DHL BORNKO 



205 




Figo. 61-67 
Habitus, edeago in visione laterale e ventrale, labio con palpo labiale, maxilla con palpo 
mascellare, spermateca e mento. 61-67: Megaparaglossa annularis gen. n., sp. n. 



206 R - PACE 

convergono in parte a comporre i caratteri del nuovo, che così risulta ingannevole per 
colui che si limita a un'osservazione superficiale. 

Apatelomixidota borneensis sp. n. Figg. 68-75 

Holotypus 9, Borneo, Sabah, Mt. Kinabalu N.P., above Poring Hot Springs, 520 m, 
9. V. 1 987, A. Smetana leg. (MHNG). 

Paratypi: 2 9?, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 550-600 m, 
9.V.1987, Burckhardt & Lobi leg.; 2 9 9, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot 
Springs, Bat Cave. 600 m, 1 O.V.I 987, Burckhardt & Lobi leg.; 1 <?, Borneo, Sabah, Mt. 
Kinabalu N.P., Poring Hot Springs, Langanan Falls, 900-950 m, 12.V.1987, Burckhardt & Lobi 
leg.; 1 9, Borneo, Sabah, Mt. Kinabalu N.P.,1750 m, 27.IV. 1987, Burckhardt & Lobi leg. 

Descrizione. Lungh. 1,8 mm. Corpo lucido e giallo-rossiccio sporco, con 
elitre e quarto urite libero bruno-rossicci; antenne brune, con i due antennomeri basali 
e l'undicesimo giallo-rossicci; zampe gialle. La reticolazione del capo è distinta, 
quella del pronoto è assente e quella delle elitre è superficiale. La granulosità del capo 
e del pronoto è saliente, fine e fitta, quella delle elitre è distinta. Edeago figg. 69-70, 
spermateca fig. 71, sesto urotergo libero del S fig. 75. 

Apatelomixidota burckhardti sp. n. Figg- 76-79 

Holotypus o\ 1 9, Borneo, Sabah, Mt. Kinabalu N.P., 1750 m, 27.IV. 1987, Burckhardt 
& Lobi leg. (MHNG). 

Paratypus: 1 S, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 500 m, 
1 1 .V. 1 987, Burckhardt & Lobi leg. 

Descrizione. Lungh. 2,0 mm. Avancorpo debolmente lucido, addome lucido. 
Corpo giallo-bruno, con capo bruno-rossiccio ed elitre e una fascia addominale brune; 
antenne brune con i due antennomeri basali e V undicesimo di un giallo sporco; zampe 
gialle. La reticolazione del capo e delle elitre è netta, quella delle elitre è distinta e 
quella dell'addome è assente. La granulosità del capo e del pronoto è appena distinta, 
quella del pronoto e dell'addome è distinta. Edeago figg 77-78, sesto urotergo libero 
del S fig. 79. 

Etimologia. Specie dedicata a uno dei suoi raccoglitori, il noto entomologo 
del Museo di Ginevra Dr. Daniel Burckhardt. 



Apatelomixidota intermedia sp. n. Figg- 80-84 

Holotypus 6 , Borneo. Sabah, Mt. Kinabalu N.P., Poring Hot Springs, nr. Bat Cave, 
600 m, 10.V.1987, Burckhardt & Lobi leg. (MHNG). 

Paratypi: 19 es., Borneo, Sabah. Mt. Kinabalu N.P., Poring Hot Springs, 600 m, 
7.V.1987. Burckhardt & Lobi leg.; 3 es.. Borneo. Sabah, Mt. Kinabalu N.P.. Poring Hot 
Springs, 550-600 m, 9.V.1987, Burckhardt & Lobi leg. 

Descrizione. Lungh. 1,6 mm. Corpo lucido e giallo-bruno, con elitre e quarto 
urite libero bruni; antenne bruno-rossicce, con i due antennomeri basali e 
l'undicesimo gialli; zampe gialle. La reticolazione dell'avancorpo è distinta, quella 
dell'addome è assente. La granulosità dell'avancorpo è saliente, quella dell'addome è 
distinta. Edeago figg. 81-82, sesto urotergo libero del S fig. 84. 

Comparazioni. Chiave delle specie del nuovo genere Apatelomixidota. 



ALEOCHARINAE DEL BORNEO 



207 




Figg. 68-75 
Habitus, edeago in visione laterale e ventrale, spermateca, mento, labio con palpo labiale, 
maxilla con palpo mascellare, sesto urotergo libero del 6 . 68-75: Apatelomixidota borneensis 
gen. n., sp. n. 



208 



R. PACE 




Figo. 76-80 
Habitus, edeago in visione laterale e ventrale e sesto urotergo libero del 6 . 76-79: Apatelo- 
mìxidota burckhardti gen. n., sp. n.; 80: Apateìomixìdota intermedia gen. n., sp. n. 



ALEOCHARINAE DEL BORNEO 209 

1 Antennomeri 8°, 9° e 10° chiaramente trasversi; elitre poco più larghe 

del pronoto. Lungh. 1,6-1,8 mm 2 

Antennomeri 8°, 9° e 10° più lunghi che larghi; elitre molto più larghe 

del pronoto. Lungh. 2,0 mm. M. Kinabalu A. burckhardti sp. n. 

2 Denti del margine posteriore del sesto urotergo libero del S, esili e 
ricurvi, tranne il mediano che è strettissimo e acutissimo. Lungh. 

1,8 mm. M. Kinabalu A. borneensis sp. n. 

Denti del margine posteriore del sesto urotergo libero del â, corti e 
retti, il mediano a punta arrotondata: Lungh. 1,6 mm. M. Kinabalu 
A. intermedia sp. n. 

Episkilepta gen. n. Figg. 85-91 

Diagnosi. Parti boccali simili a quelle del genere Paralinoglossa Pace, 1982a, 
dell'Australia, ma le paraglosse sono brevemente setolose all'apice (con setole molto 
più lunghe in Paralinoglossa) e la taglia è estremamente ridotta (1,2 mm, invece di 
2,8). Non è noto il S di Paralinoglossa. 

Descrizione. Undici antennomeri; tempie robustamente marginate; palpi 
labiali di due articoli corti; ligula come da fig. 88 (la sua estremità è forse amputata); 
palpi mascellari di 4 articoli, fig. 90; mento trapezoidale, fig. 91; mandibola, fig. 89; 
mesosterno non carenato; mesocoxe lievemente separate tra loro; formula tarsale 4-4- 
5; primo tarsomero posteriore corto. Edeago figg. 86-87. 

Tipo del genere. Episkilepta borneensis sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Piccola e opaca". 

Episkilepta borneensis sp. n. Figg. 85-91 

Holotypus 6, Borneo, Sabah, Mt. Kinabalu N.P., above Poring Hot Springs, 550 m, 
9.V.1987. A. Smetana leg. (MHNG). 

Descrizione. Lungh. 1,2 mm. Capo e pronoto assai opachi, elitre lievemente 
opache, addome lucido. Corpo bruno; antenne brune con i due antennomeri basali e 
l'undicesimo di un giallo paglierino; zampe gialle. La granulosità del capo e del 
pronoto è fittissima, sì da dare un aspetto scabro alla superficie. Le elitre presentano 
sutura saliente e granulosità fittissima. Quest'ultima sull'addome è superficiale. 
Edeago figg. 86-87. 

Tribù DIESTOTINI Mulsant & Rey, 1871 

Anamignusa gen. n. Figg- 92-96 

Diagnosi. Alcuni caratteri (pubescenza del pronoto volto in linea retta 
all' indietro, ligula incisa all'apice, spermateca avvolta a matassa) permettono di 
avvicinare il nuovo genere al genere Chledophila Cameron, 1920. Ma il corpo non è 
a lati paralleli, ma fusiforme e i palpi labiali sono più allungati. 

Descrizione. Undici antennomeri; tempie robustamente marginate; palpi 
labiali di 2 articoli allungati; ligula intera, appena incisa all'apice; paraglosse non 



210 



R. PACE 




FlGG. 81-91 

Edeago in visione laterale e ventrale, spermateca, sesto urotergo libero del 3 , labio con palpo 
labiale, mandibola, maxilla con palpo mascellare e mento. 81-84: Apatelomixidota intermedia 
gen. n., sp. n.; 85-91: Episkilepta borneensis gen. n., sp. n. 

prominenti, fig. 94; mento trapezoidale, con base, minore largamente incavata, fig. 96; 
palpi mascellari di 4 articoli, fig. 95 (l'ultimo articolo è andato perduto, forse in fase 
di raccolta dell'esemplare); mesosterno non carenato; processo mesosternale acuto; 
mesocoxe appena separate fra loro; formula tarsale 4-4-5; primo tarsomero posteriore 
corto; spermateca a matassa e spirale, fig. 93. 



ALEOCHARINAE DEL BORNEO 2 1 1 

Tipo del genere. Atramignusa borneensis sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Colei che è 
mescolata". Esso allude a caratteri di altri generi mescolati insieme a comporre questo 
nuovo genere. 

Comparazioni. Chiave dei generi asiatici della tribù Diestotini, che comprende 
specie con due articoli dei palpi labiali, formula tarsale 4-4-5 e spermateca a matassa 

a spirale. 

1 Pubescenza del pronoto volta all' indietro in linea retta; avancorpo 

finemente e fittamente granuloso 2 

Pubescenza del pronoto volta in avanti obliquamente o all' indietro 
obliquamente; avancorpo per lo più con punteggiatura o granulosità 
robuste o evidenti Diesiota Mulsant & Rey, 1871 

2 Corpo stretto e a lati paralleli; palpi labiali corti; ligula all'apice più 
profondamente incisa; margine anteriore del mento rettilineo 

Chledophila Cameron, 1920 

Corpo largo e fusiforme; palpi labiali allungati; ligula all'apice appena 
incisa; margine anteriore del mento arcuato Anamignusa gen. n. 

Anamignusa borneensis sp. n. Figg. 92-96 

Holotypus ?, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 480 m, 
20.VIII.1988, A. Smetana leg. (MHNG). 

Descrizione. Lungh. 1,7 mm. Corpo lucido e rossiccio, con elitre bruno- 
rossicce; antenne rossicce, con i due antennomeri basali e l'apice dell'undicesimo 
gialli. Non è presente reticolazione sul corpo. La granulosità del capo e delle elitre è 
distinta, quella del pronoto è assai superficiale e quella dell'addome è molto saliente. 
Spermateca fig. 93. 



Tribù BOLITOCHARINI Thomson, 1859 

Antithetusa gen. n. Figg. 97-102 

Diagnosi. Grazie all'eccezionale esilità della ligula e dell'articolo terminale 
dei palpi labiali e mascellari, è possibile avvicinare sistematicamente il nuovo genere 
al genere Plesiosipalia Bernhauer, 1943, della Nuova Zelanda. Ma il corpo è fusi- 
forme (e non simile al genere Geostiba Thomson) e il primo articolo dei palpi labiali è 
molto più lungo del secondo (di lunghezza pari a quella del secondo in Plesiosipalia). 

Descrizione. Undici antennomeri; tempie robustamente marginate; collo 
largo; palpi labiali di 3 articoli, con il primo e il terzo lunghissimi e il secondo assai 
breve; ligula esilissima e intera, fig. 101; palpi mascellari di 4 articoli assai lunghi, 
tranne il primo, fig. 100; mento trapezoidale, con base minore rettilinea, fig. 102; 
processo mesosternale carenato e largo, ad apice tronco; mesocoxe tra loro separate; 
tibie mediane e posteriori con setola isolata esterna; formula tarsale 4-4-5; primo 
tarsomero posteriore appena più corto dei due seguenti riuniti; edeago figg. 98-99. 



212 



R. PACE 




Figg. 92-96 
Habitus, spermateca, labio con palpo labiale, maxilla con palpo mascellare e mento. 92-96: 
Anamignusa borneensis gen. n., sp. n. 



ALEOCHARINAE DEL BORNEO 213 

Tipo del genere. Antithetusa inopinato sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Colei che è 
contrastante", nel senso che ha parti boccali simili a quelle del genere Plesiosipalia, 
ma l'habitus è in contrasto con quello del genere di confronto. 

Comparazioni. Chiave di alcuni generi asiatici e neozelandesi della tribù 
Bolitocharini. 

1 Ligula intera 2 

Ligula divisa . . Bolitochara Mannerheim, 1831, auct.; Pseudatheta Cameron, 
1920; Neoleptusa Cameron, 1939; Omologlusa Pace, 1998; Methistemistiba 
Pace, 1998; Homoiobrachido Pace, 1990; Sulepta Cameron, 1939; Phymatura 
Sahlberg, 1876; Caloderino Ganglbauer, 1895; Tachychara Cameron, 1920 

2 Apice della ligula arrotondato Leptusa Kraatz, 1 856 

Apice della ligula acutissimo 3 

3 Habitus di Oxypoda; primo articolo dei palpi labiali molto più lungo del 
secondo; ligula esile e lunga circa quanto la metà del primo articolo dei 
palpi labiali; margine anteriore del mento rettilineo. Borneo Antithetusa gen. n. 
Habitus di Geostiba; primo articolo dei palpi labiali lungo quanto il 
secondo; ligula robusta e lunga quanto i due articoli basali dei palpi 
labiali; margine anteriore del mento bisinuato. Nuova Zelanda 
Plesiosipalia Bernhauer, 1943 

Antithetusa inopinata sp. n. Figg. 97-102 

Holotypus 6, Borneo, Sabah, Mt. Kinabalu, 1750 m, 27.IV. 1987, Burckhardt & Lobi 
leg. (MHNG). 

Descrizione. Lungh. 1,9 mm. Corpo lucido e giallo-rossiccio, con elitre oscu- 
rate di bruno; antenne rossicce, con i due antennomeri basali gialli; zampe giallo- 
rossicce. La reticolazione del capo e del pronoto è assente, quella delle elitre è distinta 
e quella dell'addome è superficiale. La punteggiatura del capo è svanita, quella delle 
elitre è distinta. Una distinta granulosità fine copre il pronoto, quella dell'addome è 
poco distinta. Edeago figg. 98-99. 

Panbrachyna gen. n. Figg- 103-109 

Diagnosi. Per la presenza di una ligula a base larghissima e divisa all'apice, il 
nuovo genere è distinto da tutti quelli della tribù Bolitocharini, che nella totalità 
presentano ligula a base stretta. 

Descrizione. Undici antennomeri; tempie marginate; palpi labiali di 3 articoli; 
ligula corta e a base larga, fig. 106; palpi mascellari di 4 articoli; apice della galea non 
setoloso, fig. 109; mento a base anteriore rettilinea, fig. 107; mesosterno non 
carenato; processo mesosternale acuto; mesocoxe contigue; formula tarsale 4-4-5; 
edeago figg. 104-105, spermateca fig. 108. 

Tipo del genere. Panbrachyna borneensis sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Piccola tutta 
larga". Si allude alla larghezza della ligula. 



214 



R. PACE 




Figo. 97-102 
Habitus, edeago in visione laterale e ventrale, maxilla con palpo mascellare, labio con palpo 
labiale e mento. 97-102: Antithetusa inopinata gen. n., sp. n. 



Panbrachyna borneensis sp. n. 



Figg. 103-109 



Holotypus S, Borneo. Sabah, Crocker Ra., 1270 m, Km 60 rte. Kota Kinabalu- 
Tambunan, 17.V.1987, Burckhardt & Lobi leg. (MHNG). 

Paratypi: 2 9 9, stessa provenienza; 1 9, Borneo, Sabah, Mt. Kinabalu N.P., Poring 
Hot Springs, 500 m. 6.V.1987. Burckhardt & Lobi leg. 



ALEOCHARINAE DEL BORNEO 



215 




0.05 mm 



Figg. 103-109 
Habitus, edeago in visione laterale e ventrale, labio con palpo labiale, spermateca, mento e 
maxilla con palpo mascellare. 103-109: Panbrachyna borneensis gen. n., sp. n. 

Descrizione. Lungh. 1,2 mm. Corpo lucido e giallo-bruno, con pronoto, base 
ed estremità dell'addome rossicci; antenne brune con i due antennomeri basali giallo- 
rossicci; zampe gialle. La punteggiatura del capo è così fitta da dare l'aspetto di una 
distinta reticolazione. Sul resto del corpo non vi è reticolazione. La granulosità del 
pronoto e delle elitre è distinta, Una vaga scultura squamiforme sta sugli uroterghi 
liberi secondo e terzo. Edeago figg 104-105, spermateca fig. 108. 



Tribù FALAGRIINI Mulsant & Rey, 1873 
Borneopora gen. n. 



Figs. 110-114 



Diagnosi. Taxon vicino al genere Myrmecopora Saulcy, 1864, distinto 
essenzialmente per la presenza dei due rami apicali della ligula assai dilatati (stretti in 
Myrmecopora). 

Descrizione. Undici antennomeri; tempie marginate; collo stretto; palpi labiali 
di tre articoli, di cui il secondo molto ridotto; ligula divisa in due rami larghi, 



216 



R. PACE 




0.1 mm 



Figg. 110-114 
Habitus, spermateca, labio con palpo labiale, mento e maxilla con palpo mascellare. 110-114: 
Borneopora fontis gen. n.. sp. n. 



ALEOCHARINAE DEL BORNEO 2 1 7 

arrotondati all'estremità, fig. 112; palpi mascellari di 4 articoli, di cui l'apicale 
estremamente esile, fig. 114; mento fig. 113; mesosterno non carenato; processo 
mesosternale acuto; mesocoxe lievemente separate tra loro; formula tarsale 4-5-5; 
primo tarsomero posteriore lungo quanto i due seguenti riuniti; spermateca fig. 111. 

Tipo del genere. Borneopora fontis sp. n. 

Etimologia. Il nome femminile del nuovo genere è la fusione dei sostantivi 
Borneo con Myrmecopora e significa "Colei che attraversa il Borneo", seguendo le 
acque dolci. 

Borneopora fontis sp. n. Figg. 110-114 

Holotypus 9, Borneo, Sabah, Mt. Kinabalu N.P., above Poring Hot Springs, 520 m, 
9.V.1987, A. Smetana leg. (MHNG). 

Descrizione. Lungh. 2,7 mm. Corpo lucido, un po' appiattito e bruno, con 
pigidio giallo-rossiccio; antenne con i due antennomeri basali giallo-rossicci; zampe 
giallo-rossicce. Corpo senza, reticolazione. La punteggiatura del capo è netta, ma 
assente sulla linea longitudinale mediana, nel solco e sulla fronte. Vi è una profonda 
fossetta discale del capo. Il pronoto presenta un largo appiattimento mediano, una 
profonda impressione trasversa basale e punteggiatura finissima e fitta, come quella 
delle elitre. Spermateca fig. 111. 



Tribù ATHETINI Casey, 1910 

Planadota gen. n. (per Atheta borneensis Cameron, 1933) Figg- 1 15-120 

Nota. Atheta borneensis Cameron, 1933, è stata attribuita da Sawada (1980) 
al genere Pycnota Mulsant & Rey, 1874. Questa attribuzione generica per Sawada 
aveva carattere provvisorio. Infatti scriveva: In future a new genus may be established 
to accommodate this species. L'attribuzione a Pycnota non è più sostenibile perché 
questo genere è caratterizzato dalla presenza di una ligula intera e la specie borneensis 
presenta ligula divisa all'apice. Questa nuova attribuzione è effettuata a seguito 
dell'esame dell' olotipo 9 dì Atheta borneensis. 

Diagnosi. La forma della ligula permette di avvicinare il nuovo genere ai 
generi Pelioptera Kraatz, 1857 e Aloconota Thomson, 1858. Dal primo si distingue 
per la presenza di mesocoxe contigue, dal secondo per la base della ligula stretta. Da 
entrambi, per la forma della spermateca. L'assenza del S non permette ulteriori 
confronti. 

Descrizione. Undici antennomeri; tempie marginate solo posteriormente; 
palpi labiali di tre articoli; ligula non ristretta alla base e divisa all'estremità, fig. 120; 
palpi mascellari di 4 articoli, fig. 118; mento fig. 119: processo mesosternale acuto; 
mesocoxe contigue fra loro; formula tarsale 4-5-5; primo tarsomero posteriore lungo 
quanto i due seguenti riuniti. Spermateca dell'olotipo fig. 115, spermateca di un 
esemplare fig. 117. 

Tipo del genere. Planadota borneensis (Cameron, 1933) comb. n. 



218 



R. PACE 




Figg. 115-120 
Habitus, spermateca, maxilla con palpo mascellare, mento e labio con palpo labiale. 115: 
Planadota borneensis (Cameron), comb, n., holotypus; 116-120: Planadota borneensìs 
(Cameron) comb. n. 



ALEOCHARINAE DEL BORNEO 219 

Atheta (s. str.) bomeensis Cameron, 1933: 357 

Pycnota bomeensis: Sawada, 1980: 30 

Atheta (s. str.) bomeensis: Hammond, 1984: 208 

Etimologia. Il nome femminile del nuovo genere significa "Dono che induce 
in errore". 

Materiale esaminato (P. bomeensis). Holotypus 9, N. Borneo, Mt. Kina- 
balu, Kenokok, 3300 feet, 23 apr. 1929, Atheta bomeensis Cam. (British Museum). 
1 9, Borneo, Sabah, E Mt. Kinabalu, rte. Ranau-Kota Kinabalu, 1150 m, 24. V. 1987, 
Burckhardt & Lobi leg. 

Paranomusa gen. n. Figg. 121-124 

Diagnosi. Habitus simile a quello di Acrotona Thomson, 1 859, ma la ligula ha 
base larga ed è divisa fino alla base. Per questi caratteri potrebbe essere genere vicino 
a Hydrosmecta Thomson, 1858, ma l'habitus e la spermateca hanno struttura netta- 
mente differente. 

Descrizione. Undici antennomeri; tempie robustamente marginate; palpi 
labiali di 3 articoli, con il secondo cortissimo; ligula a base larga e divisa a metà fino 
alla base, fig. 123; palpi mascellari di 4 articoli; processo mesosternale acuto; meso- 
coxe contigue; formula tarsale 4-5-5; primo tarsomero posteriore lungo quanto i due 
seguenti riuniti; spermateca fig. 122. 

Tipo del genere. Paranomusa kinabaluensis sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Dono che 
trasgredisce alle norme". 

Paranomusa kinabaluensis sp. n. Figg. 121-124 

Holotypus 9, Borneo, Sabah, Mt. Kinabalu N.P., HQ Liwagu Rv. trail, 1520 m, 
11.VIII.1988, A. Smetana leg. (MHNG). 

Paratypus: 1 9, stessa provenienza, ma 1500 m e 4.VIII. 1988. 

Descrizione. Lungh. 1.8 mm. Corpo lucido e giallo-rossiccio chiaro, con elitre 
giallo-brune; antenne brune con i quattro antennomeri basali gialli; zampe gialle. La 
reticolazione del capo, delle elitre e dell'addome è distinta, quella del pronoto è 
assente: La punteggiatura del capo è distinta, quella del pronoto è superficiale. Le 
elitre presentano una granulosità distinta. Spermateca fig. 122. 

Dikraspedellagen.n. Figg. 125-136 

Diagnosi. La presenza di un lunghissima ligula, avente all'estremità due lobi 
ovali, l'apice pubescente della galea molto sviluppato e la forma caratteristica della 
spermateca, sono caratteri non riscontrabili insieme in nessun genere noto della tribù 
Athetini. 

Descrizione. Undici antennomeri; tempie robustamente marginate; palpi 
labiali di 3 articoli; ligula assai lunga, con due lobi apicali, fig. 127; palpi mascellari 
di 4 articoli; parte apicale pubescente della galea molto sviluppata, con alcune setole 
isolate, fig. 128; mento fig. 129; processo mesosternale acuto; mesocoxe tra loro 
contigue; formula tarsale 4-5-5; primo tarsomero posteriore corto; edeago figg. 126, 
133-134, spermateca fig. 130. 



220 



R. PACE 




FlGG. 121-130 

Habitus, spermateca. labio con palpo labiale, mento, edeago in visione laterale e ventrale e 
maxilla con palpo mascellare. 121-124: Paranomusa kinabaluensis gen. n., sp. n.; 125-130: 
Dikraspedella kinabaluensis gen. n., sp. n. 



ALEOCHARINAE DEL BORNEO 221 

Tipo del genere. Dikraspedella kinabaluensis sp. n. 

Etimologia. Il nome femminile della nuova specie significa "Piccola dai due 
lembi". Questi si trovano all'apice della ligula. 

Dikraspedella kinabaluensis sp. n. Figg. 125-131 

Holotypus 9, Borneo, Sabah, Mt. Kinabalu N.P., HQ Liwagu Riv. trail, 1500-1550 m, 
27.IV. 1987. A. Smetana leg. (MHNG). 

Paratypi: 3 3 3 e 1 9, Borneo, Sabah, Mt. Kinabalu N.P., 1580 m, 27.IV.1987, 
Burckhardt & Lobi leg.; 1 9, Borneo, Sabah, Mt. Kinabalu N.P., 1500 m, 30.IV.1987, 
Burckhardt & Lobi leg.; 1 â, Borneo, Sabah, Crocker Ra., 1600 m. Km 51 rte. Kinabalu- 
Tambunan. 18.V.1987, Burckhardt & Lobi leg. 

Descrizione. Lungh. 1,7 mm. Corpo lucido e rossiccio, con elitre e addome 
bruno-rossicci; antenne giallo- brune con i tre antennomeri basali gialli e i tre seguenti 
di un giallo sporco; zampe gialle. La reticolazione dell'avancorpo è distinta. La 
granulosità del capo è confusa nella reticolazione, quella del pronoto e delle elitre è 
distinta. Edeago fig. 126, spermateca fig. 130, sesto urotergo libero del 3 131. 

Dikraspedella smetanai sp. n. Figg- 132-136 

Holotypus 3, Borneo, Sabah, Mt. Kinabalu N.P., HQ Liwagu Rv. tr., 1655 m, 
ll.Vm.1988, A. Smetana leg. (MHNG). 

Paratypi: 1 3 e 1 9, stessa provenienza. 

Descrizione. Lungh. 1,9 mm. Corpo lucido e bruno, con base ed estremità 
addominale giallo-brune; antenne brune con i tre antennomeri basali di un giallo 
sporco; zampe gialle. L'avancorpo è privo di reticolazione, l'addome è distintamente 
reticolato. La punteggiatura del capo e delle elitre è svanita, quella del pronoto è 
assente come la sua granulosità. Edeago figg. 133-134, spermateca fig. 135, sesto 
urotergo libero del 3 fig. 135. 

Nota. La ligula di questa specie è meno lunga di quella del tipo del genere e, 
alla base di ciascun lobo apicale della stessa ligula, è inserita una setola. Inoltre la 
spermateca è piuttosto differente. Pertanto D. smetanai forse dovrebbe appartenere a 
un nuovo sottogenere di Dikraspedella. 

Etimologia. Specie dedicata al suo raccoglitore Dr. A. Smetana di Ottawa, 
noto studioso di Staphylinidae. 

Comparazioni. Chiave delle specie del nuovo genere Dikraspedella. 

1 Capo e pronoto rossicci; reticolazione del pronoto distinta; granulosità 

del pronoto distinta; edeago ventralmente arcuato; spermateca corta. 

Lungh. 1,7 mm. M. Kinabalu D. kinabaluensis sp. n. 

Capo e pronoto bruni; reticolazione del pronoto assente; granulosità del 
pronoto assente; edeago ventralmente bisinuoso; spermateca allungata. 
Lungh. 1,9 mm. M. Kinabalu D. smetanai sp. n. 

Trigonoglossina gen. n. Figg. 137-141 

Diagnosi. Grazie alla singolare forma della ligula e della spermateca, mai 
riscontrate nella tribù Athetini, il nuovo genere non può essere avvicinato sistema- 
ticamente a nessun genere noto, nemmeno a Pelioptera, a cui è simile per i caratteri 
dell'habitus. 



222 



R. PACE 




Figo. 131-139 
Sesto urotergo libero del 6 , habitus, edeago in visione laterale e ventrale, spermateca e labio 
con palpo labiale. 131: Dikraspedella kinabaluensis gen. n., sp. n.; 132-136: Dikraspedella 
smetanai gen. n.. sp. n.; 137-139: Trigonoglossina borneensis gen. n., sp. n. 



ALEOCHARINAE DEL BORNEO 223 

Descrizione. Undici antennomeri; tempie non marginate; palpi labiali di 3 
articoli; ligula intera e triangolare, fig. 139; palpi mascellari di 4 articoli, fig. 140; 
mento a margine anteriore sinuoso, fig. 141; mesosterno carenato; processo meso- 
sternale acuto; mesocoxe contigue tra loro; formula tarsale 4-5-? (zampe posteriori 
perdute); spermateca fig. 138. 

Tipo del genere. Trigonoglossina bomeensis sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Piccola dalla lin- 
gua triangolare". 

Trigonoglossina bomeensis sp. n. Figg. 137-141 

Holotypus 9, Borneo, Sabah, Mt. Kinabalu N.P., HQ at Liwagu Rv., 1500 m, 
30.IV. 1987, A. Smetana leg. (MHNG). 

Descrizione. Lungh. 1,6 mm. Corpo lucido e giallo-rossiccio, con capo ros- 
siccio ed elitre di un giallo sporco; antenne bruno-giallicce con i tre antennomeri 
basali gialli; zampe gialle. La punteggiatura del capo è fitta e superficiale. La 
granulosità del pronoto e delle elitre è assai svanita. Spermateca fig. 138. 

Serikasomina gen. n. Figg- 142-153 

Diagnosi. Habitus e tipo di pubescenza del pronoto (cioè diretta all' indietro in 
linea retta) propri del genere Gastropaga Bernhauer, 1915, ma parti boccali e tipo di 
spermateca differenti, cioè rami apicali della ligula cortissimi (e non lunghi come in 
Gastropaga) e apice pubescente della galea molto sviluppato. 

Descrizione. Undici antennomeri; tempie finemente marginate; palpi labiali 
di 3 articoli; ligula divisa all'estremità in due brevi rami, fig. 147; palpi mascellari di 
4 articoli; galea con estremità apicale pubescente enormemente sviluppata, fig. 143; 
processo mesosternale acuto; mesocoxe contigue fra di loro; formula tarsale 4-5-5; 
primo tarsomero posteriore corto; edeago figg. 145-146, spermateca fig. 144. 

Tipo del genere. Serikasomina smetanai sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Piccolo corpo 
sericeo". 

Serikasomina smetanai sp. n. Figg- 142-148 

Holotypus 3, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 485 m, 
29. Vili. 1988, A. Smetana leg. (MHNG). 

Paratypi: 15 es., stessa provenienza. 

Descrizione. Lungh. 1,3 mm. Corpo lucido e bruno-rossiccio, con pronoto, 
base ed estremità addominale gialli, uriti liberi 3°, 4° e 5° bruni; antenne giallo-brune, 
con i tre antennomeri basali gialli; zampe gialle. L'intero corpo è coperto di gra- 
nulosità finissima e fittissima superficiale. Edeago figg. 145-146, spermateca fig. 144. 

Etimologia. Specie dedicata al suo raccoglitore Dr. A. Smetana di Ottawa, 
noto studioso di Staphylinidae. 

Serikasomina bomeensis sp. n. Figg. 149-151 

Holotypus 6, Borneo, Sabah, Mt. Kinabalu N.P., HQ Liwagu River, 1495 m. 
2 1 . V. 1 987, A. Smetana leg. (MHNG). 



224 



R. PACE 




Figo. 140-148 
Maxilla con palpo mascellare, habitus, mento, spermateca e labio con palpo labiale. 140-141: 
Trigonoglossina borneensis gen. n., sp. n.; 142-148: Serikasomina smetanai gen. n., sp. n. 



ALEOCHARINAE DEL BORNEO 225 

Descrizione. Lungh. 1,8 mm. Corpo lucido. Capo e base delle elitre rossicci, 
pronoto, base ed estremità addominale gialli, resto del corpo bruno; antenne brune 
con i due antennomeri basali e l' undicesimo gialli, antennomeri 3° e 4° di un giallo 
sporco; zampe gialle. La punteggiatura del capo è fitta e netta, quella del pronoto è 
fine e distinta, un debole solco mediano sta sul pronoto. La granulosità delle elitre è 
fitta e poco distinta, quella deir addome è fittissima. Edeago figg. 150-151. 

Serikasomina diversearmata sp. n. Figg- 152-153 

Holotypus S, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 500 m, 
1 I.V. 1987. Burckhardt & Lobi leg. (MHNG). 
Paratypi: 2 S S , stessa provenienza. 

Descrizione. Lungh. 1,7 mm. Corpo lucido e bruno, con pronoto e addome 
giallo-bruni; antenne brune con i due antennomeri basali di un giallo sporco; zampe 
gialle. La reticolazione dell'avancorpo è assente, quella dell'addome è a maglie molto 
trasverse e superficiali. La punteggiatura del capo è superficiale e assente sul disco. 
La granulosità del pronoto e delle elitre è distinta. I tre uroterghi basali presentano una 
granulosità piuttosto fitta, i due successivi granulosità rada. Edeago fig. 153. 

Comparazioni. Chiave delle specie del nuovo genere Serikasomina. 

1 Elitre molto più lunghe del pronoto, fig. 149. Lungh. 1,8 mm. M. Kina- 
balu S. borneensis sp. n. 

Elitre appena più lunghe del pronoto, fig. 142 2 

2 Occhi meno sviluppati; pronoto più trasverso, addome fittamente 
pubescente; edeago più sviluppato, nonostante la taglia corporea sia 

minore. Lungh. 1,3 mm. M. Kinabalu S. smetanai sp. n. 

Occhi più sviluppati; pronoto meno trasverso; addome largamente 
pubescente; edeago meno sviluppato, nonostante la taglia corporea 
maggiore. Lungh. 1,7 mm S. diversearmata sp. n. 

Ektasitrachela gen. n. Figg- 154-158 

Diagnosi. Habitus simile a quello del genere Homoeusa Kraatz, 1856, ma le 
antenne sono molto lunghe, i lati del corpo portano lunghe setole, l'avancorpo è privo 
di punteggiatura e di pubescenza e la formula tarsale è 4-5-5, invece di 5-5-5. 

Descrizione. Undici antennomeri; tempie non marginate, palpi labiali di tre 
articoli; ligula corta, intera e larga, fig. 157; palpi mascellari di 4 articoli, fig. 155; 
mento trapezoidale molto largo e stretto, fig. 158; pronoto più largo delle elitre; 
mesosterno non carenato; processo mesosternale acuto; mesocoxe contigue; formula 
tarsale 4-5-5; primo tarsomero medio e posteriore lungo quanto i tre seguenti riuniti; 
edeago fig. 154. 

Tipo del genere. Ektasitrachela borneensis sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Collo espanso". 
Allude al pronoto più largo delle elitre. 

Ektasitrachela borneensis sp. n. Figg. 154-158 

Holotypus 6, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 500 m, 6.V.1987, 
Burckhardt & Lobi leg. (MHNG). 



226 



R. PACE 




Figo. 149-155 
Habitus, edeago in visione laterale e ventrale, maxilla con palpo mascellare. 149-151: 
Serikasomina borneensis gen. n., sp. n.; 152-153: Serikasomina diversearmata gen. n., sp. n.; 
154-155: Ektasitrachela borneensis gen. n., sp. n. 



ALEOCHARINAE DEL BORNEO 227 

Descrizione. Lungh. 2,0 mm. Corpo lucido e bruno, con pronoto rossiccio; 
antenne di un giallo sporco; zampe giallo-rossicce. L'avancorpo è privo di retico- 
lazione, l'addome mostra una reticolazione molto superficiale e solo sui tre uroterghi 
basali. Il capo, il pronoto e le elitre sono senza punteggiatura e senza granulosità: un 
solo granulo sta sulle elitre. Edeago fig. 154. 

Tribù THAMIARAEINI Fenyes, 1921 

Diabainella gen. n. Figg. 159-166 

Diagnosi. In base alla struttura della spermateca (avvolta a spirale), il nuovo 
genere, nell'ambito della tribù Thamiaraeini, può essere comparato solo con il genere 
Franzidota Pace, 1982b. Ma la ligula nel nuovo genere è lunghissima e non corta e a 
base larga come in Franzidota e il mesosterno è carenato (senza carena in Fran- 
zidota). 

Descrizione. Undici antennomeri; tempie robustamente marginate; palpi 
labiali stiliformi di 2 articoli; ligula lunghissima e divisa all'apice, fig. 163; palpi 
mascellari di 4 articoli; lacinia strettissima, fig. 164; mento fig. 165; mesosterno 
carenato; processo mesosternale acuto; mesocoxe tra loro contigue; formula tarsale 4- 
5-5; primo tarsomero posteriore più lungo dei due successivi riuniti; edeago figg. 160- 
161, spermateca fig. 162, sesto urotergo libero del S fig. 166. 

Tipo del genere. Diabainella borneensis sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Piccola che 
travalica", nel senso che mostra caratteri di altre tribù, così scavalcando le categorie 
tassonomiche stabilite. 

Diabainella borneensis sp. n. Figg. 159-166 

Holotypus S , Borneo, Sabah, Mt. Kinabalu N.P., HQ Liwagu River Trail, 1520 m, 
11. VIII. 1988, A. Smetanaleg. (MHNG). 

Paratypi: 14 es., stessa provenienza; 4 es., Borneo, Sabah, Mt. Kinabalu N.P., HQ 
Liwagu River, 1430 m, 5.VIII.1988 , A. Smetana leg.; 1 S, Borneo, Sabah, Mt. Kinabalu N.P., 
Poring Hot Springs, 550-600 m, 9.V.1987, Burckhardt & Lobi leg.; 1 6, Borneo, Sabah, Mt. 
Kinabalu N.P., Poring Hot Springs, 500 m, 13.V.1987, Burckhardt & Lobi leg.; 4 es., Borneo, 
Sabah, rte. Ranau-Kota Kinabalu, 1150 m, 24.V.1987, Burckhardt & Lobi leg.; 1 $, 1 S, 
Borneo, Sabah, Mt. Kinabalu N.P., 1500 m, 30.IV. 1987, Burckhardt & Lobi leg. 

Descrizione. Lungh. 1,5 mm. Corpo lucido e giallo-rossiccio, con elitre e una 
fascia addominale brune; antenne nero-brune con i tre antennomeri basali gialli e 
apice dell'undicesimo di un giallo sporco; zampe gialle. Sul corpo non è presente 
reticolazione. La punteggiatura del capo è molto superficiale. La granulosità del 
pronoto è molto svanita, quella delle elitre e dell'addome è distinta. Edeago figg. 160- 
161, spermateca fig. 162, sesto urotergo libero del â fig. 166. 

Tribù LOMECHUSINI Fleming, 1821 (=Myrmedoniini) 

Borneozyras gen. n. Figg. 167-172 

Diagnosi. Genere sicuramente appartenente alla tribù Lomechusini, per la 
presenza di setole al margine interno della lacinia (spine in altre tribù). Per la struttura 



228 



R. PACE 




Figo. 156-163 
Habitus, labio con palpo labiale, mento, edeago in visione laterale e ventrale e spermateca. 156- 
158: Ektasitrachela bomeensis gen. n., sp. n.; 159-163: Diabainella borneensis gen. n., sp. n. 



ALEOCHARINAE DEL BORNEO 



229 




Figg. 164-170 
Maxilla con palpo mascellare, mento, sesto urotergo libero del 6 , habitus e labio con palpo 
labiale. 164-166: Diabainella borneensis gen. n., sp. n.; 167-170: Bomeozyras smetanai gen. 
n., sp. n. 



230 R - PACE 

dell'edeago e per l'habitus, è genere simile a Zyras Stephens, 1835, auct., ma la 
ligula è intera (divisa in Zyras) e le antenne sono fortemente ispessite. 

Descrizione. Antenne corte con antennomeri 4° a 10° fortemente trasversi; 
tempie marginate; palpi labiali di 3 articoli; ligula intera con alcune setole apicali, fig. 
170; palpi labiali di 4 articoli, fig. 171; mento fig. 172; processo mesosternale assai 
largo e tronco; mesocoxe tra loro assai separate; formula tarsale 4-5-5; primo tarso- 
mero posteriore corto; edeago figg. 168-169. 

Tipo del genere. Borneozyras smetanai sp. n. 

Etimologia. Il nome maschile del nuovo genere significa " Zyras del Borneo". 

Borneozyras smetanai sp. n. Figg- 167-172 

Holotypus S, Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, Area Eastern 
Ridge tr., 850 m, 28. Vili. 1988, A. Smetana leg. (MHNG). 

Descrizione. Lungh. 4,2 mm. Corpo lucido e rossiccio; antenne rossicce con i 
due antennomeri basali giallo-rossicci; zampe giallo-rossicce. Sul corpo non è pre- 
sente reticolazione. La punteggiatura del capo è ombelicata, molto superficiale e 
assente sulla linea mediana, quella del pronoto è come quella del capo, ma presente 
sulla linea mediana, quella delle elitre è netta e quella dell'addome è fine. Edeago 
figg. 168-169. 

Tribù HOPLANDRIINI Casey, 1910 

Borneusa gen n. Figg- 173-178 

Diagnosi. Genere sicuramente sistematicamente vicino al genere Tacata 
Blackwelder, 1952 {-Atacta Cameron, 1939), ma la ligula è decisamente più larga e 
appena incisa all'apice (incisa fino a metà in Tacata). 

Descrizione. Undici antennomeri; tempie non marginate; mandibole molto 
lunghe e strette, la destra con dente interno, la sinistra senza; palpi labiali di 4 articoli, 
ligula assai larga e appena smarginata all'apice; paraglosse assai prominenti, fig. 174; 
palpi mascellari di 5 articoli, fig. 177; mesosterno non carenato; processo meso- 
sternale acuto; tibie anteriori e medie spinose al lato esterno; formula tarsale 4-5-5; 
primo tarsomero posteriore lungo quanto i due seguenti riuniti; edeago figg. 175-176. 

Tipo del genere. Borneusa insolita sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Colei che è del 
Borneo". 

Borneusa insolita sp. n. Figg. 173-178 

Holotypus o\ Borneo, Sabah, Mt. Kinabalu N.P., Poring Hot Springs, 500 m, 6.V.1987, 
Burckhardt & Lobi leg. (MHNG). 

Descrizione. Lungh. 2,8 mm. Corpo lucido e giallo-bruno, con addome bruno, 
tranne il pigidio che è bruno-rossiccio; antenne brune con i due antennomeri basali e 
la base del terzo gialli; zampe gialle. Il corpo è privo di reticolazione, tranne sulla 
metà posteriore del sesto urotergo libero, dove è robusta. La punteggiatura del capo e 
del pronoto è netta, sul disco del capo, che è ampiamente concavo, è largamente 



ALEOCHARINAE DEL BORNEO 



231 




FlGG. 171-174 

Maxilla con palpo mascellare, mento, habitus, labio con palpo labiale. 171-172: Borneozyras 
smetanai gen. n., sp. n.; 173-174: Borneusa insolita gen. n., sp. n. 



232 R - PACE 

assente. La granulosità delle elitre è assai superficiale, quella sui quattro uroterghi 
basali è distinta. Edeago figg. 175-176. 

Tribù OXYPODINI Thomson, 1859 

Apatelieida gen. n. (per Neosilusa stricticollis Cameron, 1943) Figg- 179-187 

Nota. Per essere attribuita al genere Neosilusa Cameron, 1920, la specie 
stricticollis dovrebbe presentare formula tarsale 4-4-5. Ma essa è 5-5-5 e i palpi 
labiali sono composti di 3 articoli, invece di 2, perciò non può più appartenere alla 
tribù Homalotini. 

Diagnosi. Habitus di Neosilusa Cameron, 1920, ma per la formula tarsale 5-5- 
5, per la forma della spermateca e della ligula, il nuovo genere sembra affine al genere 
Pseudoplandria Fenyes, 1921. Ma i palpi labiali sono composti di 3 articoli, invece di 
4 e i palpi mascellari di 4 articoli, invece di 5. Inoltre non esistono due setole laterali 
della ligula, presenti al contrario in Pseudoplandria e l' edeago mostra una struttura 
nettamente differente e unica. 

Descrizione. Undici antennomeri; tempie marginate; palpi labiali di 3 articoli; 
ligula divisa fino alla sua metà, fig. 186; palpi mascellari di 4 articoli, fig. 177; mento 
fig. 178; mesosterno non carenato; processo mesosternale acuto; mesocoxe appena 
separate tra loro; formula tarsale 5-5-5; primo tarsomero posteriore appena più corto 
dei due seguenti riuniti; edeago figg. 180-181, spermateca figg. 182-183. 

Tipo del genere. Apatelieida stricticollis (Cameron, 1943) comb. n. 

Neosilusa stricticollis Cameron, 1943:40 

Neosilusa stricticollis: Hammond, 1984: 211. 

Etimologia. Il nome femminile del nuovo genere significa "Aspetto 
ingannevole", per il fatto che se non si esaminano in preparato microscopico la 
formula tarsale e le parti boccali, si è indotti, come lo è stato Cameron, ad attribuire la 
specie al genere Neosilusa, data l'impressionante somiglianza dell'habitus. 

Materiale esaminato (A. stricticollis). Holotypus 9, Borneo, Mt. Poi, 5000 
feet, N. stricticollis Cam., type (British Museum). 2 6 S e 4 9 9, Borneo, Sabah, Mt. 
Kinabalu N.P.. Poring Hot Springs. 500 m, 7.V.1987, Burckhardt & Lobi leg. 

Syntemusa gen. n. Figg- 187-198 

Diagnosi. Pur presentando formula tarsale 5-5-5, invece di 4-5-5, il nuovo 
genere si mostra affine al genere Tacata Blackwelder, 1952, per l'habitus e la 
struttura dell' edeago (serie tipica del tipo del genere Tacata, cioè T. floralis 
(Bernhauer, 1915) da me esaminata), ma la ligula è più stretta e ha due setole apicali 
(setole laterali in Tacata), per la lacinia lunga e fortemente ricurva (lacinia corta e 
rettilinea in Tacata) e per il primo tarsomero posteriore corto (lungo quanto i due 
tarsomeri successivi riuniti in Tacata). 

Descrizione. Undici antennomeri; tempie robustamente marginate; palpi 
labiali di tre articoli; ligula a base larga, divisa in due lobi triangolari, con lunga setola 
apicale, fig. 192; palpi labiali di 4 articoli; lacinia fortemente ricurva, fig. 193; mento 
fortemente incavato al lato anteriore, fig. 114; processo mesosternale ad apice acuto, 



ALEOCHARINAE DEL BORNEO 



233 




Figo. 175-183 
Edeago in visione laterale e ventrale, maxilla con palpo mascellare, mento, habitus e sper- 
mateca. 175-178: Borneusa insolita gen. n., sp. n.; 179-182: Apatelieida stricticollis (Cameron) 
comb, n.; 183: Apatelieida stricticollis (Cameron) comb, n., holotypus. 



234 



R. PACE 




Figo. 184-189 
Maxilla con palpo mascellare, mento, mandibola, labio con palpo labiale e spermateca. 184- 
187: Apatelieida stricticollis (Cameron) comb, n.; 188-189: Syntemusa kinabaluensis gen. n., 
sp. n. 



ALEOCHARINAE DEL BORNEO 235 

ma arrotondato; mesocoxe appena separate fra loro; sesto sternite libero del S 
prolungato all' indietro; formula tarsale 5-5-5; primo tarsomero posteriore corto; 
edeago fortemente ricurvo, figg. 190-191 e 195-196, spermateca figg. 188 e 198. 

Tipo del genere. Syntemusa kinabaluensìs sp. n. 

Etimologia. Il nome femminile del nuovo genere significa "Maschio pene- 
trante" e allude alla forma ricurva dell 'edeago, particolarmente adatto alla pene- 
trazione. 

Syntemusa kinabaluensìs sp. n. Figg. 188-194 

Holotypus o\ Borneo, Sabah, Mt. Kinabalu N.P., HQ Bukit Ular trail, 1750 m, 
29.IV. 1987. A. Smetana leg. (MHNG). 

Paratypi: 42 es., stessa provenienza; 150 es., Borneo, Sabah, Mt. Kinabalu N.P., HQ 
Liwagu River, 1500 m, 27.IV. 1987 , A. Smetana leg. 

Descrizione. Lungh. 3,6 mm. Corpo lucido e bruno con pigidio bruno- 
rossiccio; antenne rossicce, con i tre antennomeri basali giallo-rossicci e undicesimo 
bruno-rossiccio; zampe gialle. Il capo e l'addome presentano reticolazione molto 
superficiale, essa è assente sul resto del coipo. La punteggiatura del capo è distinta, 
quella del pronoto è superficiale, quella delle elitre è molto svanita. Spermateca fig. 
188, edeago figg. 190-191. 

Syntemusa smetanai sp. n. Figg. 195-198 

Holotypus 6, Borneo, Sabah. Mt. Kinabalu N.P., HQ at Liwagu Rv., D.E. Bright leg. 
(MHNG). 

Paratypi: 4 S S e 1 9, Borneo, Sabah, Mt. Kinabalu N.P., 1550-1580 m, 27- 
28.IV. 1987, Burckhardt & Lobi leg. 

Descrizione. Lungh. 3,8 mm. Corpo lucido e nero-bruno; antenne brune con i 
cinque antennomeri basali rossicci; zampe giallo-rossicce. La reticolazione del capo e 
del pronoto è assente, quella delle elitre e dell'addome è superficiale. La pun- 
teggiatura del capo è molto svanita. La granulosità del pronoto è saliente solo in 
avanti sulla linea mediana, sul resto della superficie essa è estremamente superficiale, 
come quella delle elitre. Edeago figg. 195-196, spermateca fig. 198. 

Comparazioni. Chiave delle specie del nuovo genere Syntemusa. 

1 Punteggiatura del capo distinta; quarto antennomero più lungo che 

largo; pronoto meno trasverso; edeago non strozzato nella regione 
preapicale, in visione ventrale; spermateca corta. Lungh. 3,6 mm. M. 

Kinabalu S. kinabaluensìs sp. n. 

Punteggiatura del capo superficiale; quarto antennomero trasverso; 
pronoto maggiormente trasverso; edeago strozzato nella regione pre- 
apicale, in visione ventrale; spermateca lunghissima. Lungh. 3,8 mm. 
M. Kinabalu S. smetanai sp n. 



236 



R. PACE 




Figg. 190-193 
Edeago in visione laterale e ventrale, labio con palpo labiale e maxilla con palpo mascellare. 
190-193: Syntemusa kinabaluensis gen. n., sp. n. 



ALEOCHARINAE DEL BORNEOO 



237 




194 




Figg. 194-196 
Mento ed edeago in visione laterale e ventrale. 194: Syntemusa kinabaluensis gen. n., sp. n. 
195-196: Syntemusa smetanai gen. n., sp. n. 



238 



R. PACE 




197 




198 



Figo. 197-198 
Habitus e spermateca. 197-198: Syntemusa smetanai gen. n., sp. n. 

RINGRAZIAMENTI 

Rivolgo i miei più cordiali ringraziamenti a coloro che mi hanno affidato in 
studio il raro materiale oggetto del presente lavoro: il Dr. Ales Smetana di Ottawa, il 
Dr. Ivan Lobi del Museo di Storia Naturale di Ginevra, il Dr. Volker Assing di 
Hannover e il collega Guillaume de Rougemont di Londra. Per il prestito di tipi e per 
l'aiuto nella ricerca bibliografica ringrazio il Dr. Brendell del Museo di Storia 
Naturale di Londra. 



BIBLIOGRAFIA 

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Entomological Society of London 1920: 212-284. 



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Revue suisse de Zoologie 109 (1): 241; mars 2002 



Erratum 

Ng, H. H. & Kottelat, M. 2001. A review of the genus Batasio (Teleostei: Bagridae) 
in Indochina, with the description of B. tigrinus sp. n. from Thailand. Revue suisse de 
Zoologie 108 (3): 495-511. 



Due to the insufficient printing quality of Fig. 1 published in the above mentioned 
paper (p. 498) a new figure is printed here. 




Fig. 1 
Batasio affinis, after photograph of a specimen collected from Tenasserim River (drawing by 
Kelvin K. P. Lim). 



REVUE SUISSE DE ZOOLOGIE 

Tome 109 — Fascicule 1 

Pages 

Benjamin, Suresh P. Smodicinodes schwendingeri sp. n. from Thailand and 
the first male of Smodicinodes Ono, 1993, with notes on the phyloge- 
netic relationships in the tribe Smodicinini (Araneae: Thomisidae) . . . 3-8 

Reutter, Brigitte A., Petit, Eric & Vogel, Peter. Molecular identification 
of an endemic Alpine mammal, Apodemus alpicola, using a PCR-based 
RFLP method 9-16 

Blöchlinger, Beatrice & Lüps, Peter. Proportionsänderungen beim M] im 
Gebiss des mitteleuropäischen Dachses Mêles meles (Mammalia, Car- 
nivora) 17-22 

Juvara-Bals, Ilinca. A revision of the genus Heteroparasitus new status, 
with the description of Heteroparasitus (Medioparasitus) athiasae sub- 
gen, n., sp. n. from Spain and with a key to the genera of Pergamasinae 
(Acari, Gamasida, Parasitidae) 23-46 

Schwendinger, Peter J. & Martens, Jochen. A taxonomic revision of the 
family Oncopodidae III. Further new species of Gnomulus Thorell 
(Opiliones, Laniatores) 47-113 

Lourenço, Wilson R. & Pézier, Adeline. Taxonomic consideration of the 
genus Odontobuthus Vachon (Scorpiones, Buthidae), with description 
of a new species 1 1 5- 1 25 

Lourenço, Wilson R. Nouvelles additions à la faune de scorpions néotropi- 
caux (Arachnida) 127-141 

Zompro, Oliver. A revision of Oreophoetes Rehn, 1904, and description of 
a new genus (Insecta: Phasmatodea: Anareolatae: Diapheromeridae: 
Diapheromerinae: Oreophoetini) 143-153 

Casciotta, Jorge R., Azpelicueta, M. de las Mercedes & Almirón, 
Adriana E. Bryconamericus uporas sp. n. (Characiformes, Characidae), 
a new species from the rio Uruguay basin, in Argentina 155-165 

Deltshew, Christo & Curcic, Bozidar P. M. A contribution to the study of 
the genus Centromerus Dahl (Araneae: Linyphiidae) in caves of the 
Balkan Peninsula 167-176 

Neuhäuser-Happe, Lorenz. Zwei Bythinini neu für Friaul-Julisch Venetien 
(Italien): Gasparobythus kahleni sp. n. und Tychobythinus xambeui 
manfredi ssp. n. (Coleoptera, Staphylinidae, Pselaphinae) 177-187 

Pace, Roberto. Nuovi generi di Aleocharinae del Borneo (Coleoptera, Sta- 
phylinidae) 189-240 

Erratum (Ng, H.H. & Kottelat, M. 2001) 241 



REVUE SUISSE DE ZOOLOGIE 
Volume 109 — Number 1 

Pages 

Benjamin, Suresh P. Smodicinodes schwendingeri sp. n. from Thailand and 
the first male of Smodicinodes Ono, 1993, with notes on the phyloge- 
netic relationships in the tribe Smodicinini (Araneae: Thomisidae) . . . 3-8 

Reutter, Brigitte A., Petit, Eric & Vogel, Peter. Molecular identification 
of an endemic Alpine mammal, Apodemus alpicola, using a PCR-based 
RFLP method 9-16 

Blöchlinger, Beatrice & Lüps, Peter. Changing proportions in Mj in the 

European badger. Mêles meles (Mammalia, Carnivora) 17-22 

Juvara-Bals, Ilinca. A revision of the genus Heteroparasitus new status, 
with the description of Heteroparasitus (Medioparasitus) athiasae sub- 
gen, n., sp. n. from Spain and with a key to the genera of Pergamasinae 
(Acari, Gamasida, Parasitidae) 23-46 

Schwendinger, Peter J. & Martens, Jochen. A taxonomic revision of the 
family Oncopodidae III. Further new species of Gnomulus Thorell 
(Opiliones, Laniatores) 47-113 

Lourenço, Wilson R. & Pézier, Adeline. Taxonomic consideration of the 
genus Odontobuthus Vachon (Scorpiones, Buthidae), with description 
of a new species 115-125 

Lourenço, Wilson R. New records of scorpions for the Neotropics (Arach- 

nida) 127-141 

Zompro, Oliver. A revision of Oreophoetes Rehn, 1904, and description of 
a new genus (Insecta: Phasmatodea: Anareolatae: Diapheromeridae: 
Diapheromerinae: Oreophoetini) 143-153 

Casciotta, Jorge R., Azpelicueta, M. de las Mercedes & Almirón, 
Adriana E. Bryconamericus uporas sp. n. (Characiformes, Characidae), 
a new species from the rio Uruguay basin, in Argentina 155-165 

Deltshew, Christo & Curcic, Bozidar P. M. A contribution to the study of 
the genus Centromerus Dahl (Araneae: Linyphiidae) in caves of the 
Balkan Peninsula 167-176 

Neuhäuser-Happe, Lorenz. Two Bythinini new to Italy: Gasparobythus 
kahleni sp. n. and Tychobythinus xambeui manfredi ssp. n. (Coleoptera, 
Staphylinidae, Pselaphinae) 177-187 

Pace, Roberto. New genera of Aleocharinae from Borneo (Coleoptera, Sta- 
phylinidae) 189-240 

Erratum (Ng, H.H. & Kottelat, M. 2001 ) 241 

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Volume 109 - Number 1 - 2002 

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Penard, E. 1889. Etudes sur quelques Héliozoaires d'eau douce. Archives de Biologie 9: 1-61 . 
Mertens. R. & Wermuth. H. 1960. Die Amphibien und Reptilien Europas. Kramer, Frankfurt am Main, XI + 264 pp. 
Handley. C. O. Jr 1966. Checklist of the mammals of Panama (pp. 753-795). In: Wenzel, R. L. & Tipton, V. J. (eds). 

Ectoparasites of Panama. Field Museum of Natural History, Chicago, XII + 861 pp. 

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Home page RSZ: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 



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