(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Children's Library | Biodiversity Heritage Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Revue suisse de zoologie"

it 



LU 



tv H U-ES 


O 


de la 


O 


SOCIÉTÉ SUISSE DE ZOOLOGIE 


et du 


MBÏ 


MUSÉUM D'HISTOIRE NATURELLE 





de la Ville de Genève 







tome 1 1 4 
fascicule 3 


N 


2007 


LU 




û 




SUJ 




o(/5 




St/5 




O MM 




:d 








zLU 




ID 




:> 




?LU 


Jel GENEVE SEPTEMBRE 2007 ISSN 0035 - 418 X 


*tó 



REVUE SUISSE DE ZOOLOGIE 



TOME 114— FASCICULE 3 

Publication subventionnée par: 

Académie suisse des Sciences naturelles (SCNAT) 

Ville de Genève 

Société suisse de Zoologie 



DANIELLE DECROUEZ 

Directrice du Muséum d'histoire naturelle de Genève 

ALICE CIBOIS, PETER SCHUCHERT 

Chargés de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

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: taxonomie, 

systématique, faunistique, phylogénie, évolution, morphologie et anatomie comparée. 

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 



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



Prix de l'abonnement: 

SUISSE Fr. 225.— UNION POSTALE Fr. 250.- 

(en francs suisses) 

Les demandes d'abonnement doivent être adressées 

à la rédaction de la Revue suisse de Zoologie, 

Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse 



ANNALES 

de la 

SOCIÉTÉ SUISSE DE ZOOLOGIE 

et du 

MUSÉUM D'HISTOIRE NATURELLE 

de la Ville de Genève 



tome 1 14 
fascicule 3 
2007 




LU 
O 

o 



o 
o 

N 

LU 

û 



>- 
o 



LU 
oÇ/} 
SC/} 

zLU 

-> 

(A) ~ 



il * t^ 

M GENEVE SEPTEMBRE 2007 ISSN 0035 -418 X C^ ■■■■ 



REVUE SUISSE DE ZOOLOGIE 



TOME 114 — FASCICULE 3 

Publication subventionnée par: 

Académie suisse des Sciences naturelles (SCNAT) 

Ville de Genève 

Société suisse de Zoologie 



DANIELLE DECROUEZ 

Directrice du Muséum d'histoire naturelle de Genève 

ALICE CIBOIS, PETER SCHUCHERT 

Chargés de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

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: taxonomie, 

systématique, faunistique, phylogénie, évolution, morphologie et anatomie comparée. 

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 



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



Prix de l'abonnement: 

SUISSE Fr. 225.— UNION POSTALE Fr. 250.- 

(en francs suisses) 

Les demandes d'abonnement doivent être adressées 

à la rédaction de la Revue suisse de Zoologie, 

Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse 



Revue suisse de Zoologie 114 (3): 441-469; septembre 2007 



Description of a new African genus and a new tribe of 
Speleketorinae (Psocodea: 'Psocoptera': Prionoglarididae) 

Charles LIENHARD 

Muséum d'histoire naturelle, c. p. 6434, CH-1211 Genève 6, Switzerland. 

E-mail : Charles .lienhard@ ville-ge .eh 



Description of a new African genus and a new tribe of Speleketorinae 
(Psocodea: 'Psocoptera': Prionoglarididae). - Priorioglaridids are pro- 
bably the most basal family of extant psocids and may be considered as 
living fossils. The genus Afrotrogla gen. n. is described for three new 
species from southern Africa, two of them only known from caves: A. oryx 
sp. n. (South Africa, in cave, type species), A. maraisi sp. n. (Namibia) and 
A. f abella sp. n. (Namibia, in cave). The male of the second African genus 
of the subfamily, Sensitibilla Lienhard, is described for the first time and 
two new species of Sensitibilla are described, one of them only known from 
a cave: S. brandbergensis sp. n. (Namibia) and S. roessingensis sp. n. 
(Namibia, in cave). Some structures of the type species S. strinata Lienhard 
are also illustrated. A comparison of these two genera with the third known 
genus of the subfamily, the North American Speleketor Gurney, shows that 
the African genera are characterized by some striking synapomorphies in 
male and female genitalia. Thus a subdivision of the subfamily 
Speleketorinae in two tribes is proposed: Speleketorini for Speleketor and 
Sensitibillini trib. n. for Sensitibilla and Afrotrogla. Among other features, 
Sensitibillini are characterized by the presence of a trichobothrium on the 
hindtarsus. Tarsal trichobothria are not known elsewhere in insects. Male 
and female terminalia of Speleketor irwini Mockford are also illustrated. 

Keywords: Trogiomorpha - new species - cave fauna - living fossils - 
trichobothria - Namibia - South Africa - North America. 



INTRODUCTION 

Within the order Psocodea (sensu Yoshizawa & Johnson, 2006) the 'Psocoptera' 
family Prionoglarididae forms one of the most basal clades of the basal suborder 
Trogiomorpha and has recently been classified in an infraorder of its own, the 
Prionoglaridetae (see Yoshizawa et al., 2006). The family has been subdivided into two 
subfamilies by Lienhard (2004), Prionoglaridinae and Speleketorinae. Each of the sub- 
families has been shown to be monophyletic by both morphological and molecular 
analyses (Lienhard, 2004; Yoshizawa et al., 2006). The nominate subfamily contains 
the Palaearctic genus Prionoglaris Enderlein (3 species, see Lienhard & Smithers, 



Manuscript accepted 07.03.2007 



442 C. LIENHARD 

2002) and the Oriental genus Siamo glaris Lienhard (monotypic, see Lienhard, 2004); 
the subfamily Speleketorinae contains the Nearctic genus Speleketor Gumey (3 
species, see Lienhard & Smithers, 2002) and the Aethiopian genus Sensitibilla 
Lienhard (up to now monotypic, see Lienhard, 2000). Most of the few known priono- 
glaridid species live in caves or similar habitats and are considered as very rare. 

The family is characterized among extant members of the Psocoptera by its 
unique and absolutely diagnostic fore wing venation, in particular by the well-de- 
veloped and strongly arched basal section of Sc, joining Rl near base of pterostigma, 
and by the presence of a long cross-vein between base of pterostigma and distal section 
of Rs. However, very similar wing venations have been observed in several of the 
oldest known fossil Trogiomorpha, recently described from Cretaceous amber and 
assigned to different families (see Baz & Ortufio, 2000, 2001; Perrichot et al., 2003; 
Azar & Nel, 2004). Therefore the characters of wing morphology have to be consi- 
dered as symplesiomorphic in extant Prionoglarididae for which the term "living 
fossils" may be appropriate. Thus, Yoshizawa et al. (2006) have tentatively postulated 
that they are Pangaean relicts, in view of their extremely disjunct distribution, their 
cavernicolous biology and in agreement with the results of the most recent phylo- 
genetic and palaeontological analyses of Psocodea. 

Contrary to the monophyly of the subfamilies, the monophyly of the family 
Prionoglarididae was only weakly supported by the available molecular data 
(Yoshizawa et al., 2006) and its morphological definition is only based on the tentative 
autapomorphy of the phallosome structure and the possibly autapomorphic simplifi- 
cation or reduction of the lacinia in adults (Mockford, 1984; Lienhard, 2004). 
However, the latter may also be interpreted as an adaptive homoplasy related to the 
particular biology of these generally cavernicolous psocids. 

In this paper five new African species of the subfamily Speleketorinae, 
belonging to the genus Sensitibilla and to a new genus, Afrotrogla gen. n., are 
described on the basis of material from Namibia and South Africa deposited in the 
National Museum of Namibia, Windhoek. This material allows the description of the 
formerly unknown male of Sensitibilla and of both sexes of the new genus, enabling 
us for the first time to compare also male genital structures between the African and 
American members of the subfamily. The African genera have some striking synapo- 
morphies in male and female genitalia rendering these structures completely different 
from the genitalia of Speleketor. This fundamental difference in genital morphology, 
together with some other characters, justifies the establishment of a new tribe, 
Sensitibillini trib. n., for the African genera. The remaining genus of the subfamily, the 
North American Speleketor, constitutes the nominate tribe Speleketorini. 

From this study of new African material the presence of tibial and tarsal tri- 
chobothria in these psocids, as described for Sensitibilla strinata Lienhard (see 
Lienhard, 2000), can be confirmed. The first SEM micrographs of such trichobothria, 
which are unique in Psocoptera, are presented together with a brief discussion on leg 
trichobothria in insects. 

The following abbreviations are used in the descriptions: BL = body length (in 
alcohol); F = hindfemur (length); fl, f2, etc. = antennal flagellomeres (length); FW = 
forewing (length); HW = hindwing (length); IO/D= shortest distance between 



NEW AFRICAN SPELEKETORINAE 443 

compound eyes divided by anteroposterior diameter of compound eye in dorsal view 
of head; P1-P4 = articles of maxillary palp; T = hindtibia (length); tl, t2, t3 = tarso- 
meres of hindtarsus (length, measured from condyle to condyle). Abbreviations of 
wing veins and cells are used according to Yoshizawa (2005). - The material examined 
has been deposited in the following institutions: MHNG Muséum d'histoire naturelle, 
Geneva, Switzerland; NMN National Museum of Namibia, Windhoek. 

TAXONOMIC TREATMENT 

Key to the tribes and genera of Speleketorinae and to the species of 
Sensitibillini 

Note. A key to the subfamilies of Prionoglarididae (Prionoglaridinae and 
Speleketorinae) and to the genera of Prionoglaridinae (Priono'glaris Enderlein and 
Siamoglaris Lienhard) has been given by Lienhard (2004). For figures of Speleketor 
spp. see also Gurney (1943), Mockford (1984, 1993) and Lienhard (2000); for figures 
of Sensitibilla strinata see also Lienhard (2000). 

1 Hindwing with vein Rs 2-branched (Fig. 3i). Forefemur with a longitu- 

dinal row of articulated spines on anterior face. Some long and fine tri- 
chobothria present on femora and on some trochanters, no trichobothria 
on other segments of legs. P4 with 7 thin-walled conical sensilla, two of 
them situated in basal half (Fig. 3c). Female genitalia (Fig. lc): Ovi- 
positor consisting of a pair of very broad and simple external valvulae, 
peripherically setose, laterally articulated to clunium but not fused to 
subgenital plate ventrobasally (a reduced triangular and bare dorsal 
valvula is also present, completely covered by the external valvula); 
spermathecal duct short and relatively wide. Male genitalia (Fig. 2cd): 
Phallosome with a posterolateral pair of pore-bearing processes, scle- 
rites of phallosome anteriorly closed, posteriorly open (Speleketorini) 

Speleketor Gurney, 1943 

Three Americal species known (keyed by Mockford, 1993). Type species: 
S.flocki Gurney. 

1 ' Hindwing with vein Rs simple (Fig. 4b). Forefemur with only a longitu- 
dinal row of normal setae on anterior face. Some long and fine tri- 
chobothria (see PI. 1) present on tibiae and hindtarsus, no trichobothria 
on other segments of legs. P4 with 2-5 thin- walled conical sensilla situ- 
ated in apical half. Female genitalia (Figs 4g, 8c): Ovipositor consisting 
only of a pair of external valvulae, laterally articulated to clunium and 
ventrobasally fused to subgenital plate, each valvula bearing a distal 
process with a claw-like articulated spine at its tip; spermathecal duct 
long and thin. Male genitalia (Figs 5c, 9df): Phallosome lacking pore- 
bearing processes, sclerites of phallosome anteriorly open, posteriorly 
convergent or forming a closed aedeagal arch (Sensitibillini trib. n.) 2 

2(1') Hindwing with vein M 2-branched (Fig. 4b). P4 with 5 thin- walled 
conical sensilla (Fig. 3e). Female genitalia (Fig. 4g): Posterior part of 
subgenital plate sclerotized and clearly visible medially between ovi- 



444 C. LIENHARD 

positor valvulae; spermapore situated near posterior end of a mem- 
branous sac bearing a complex scaffolding of sclerotized struts. Male 
genitalia (Fig. 5c): Phallosome with a narrow sclerite forming a simple 
aedeagal arch (Afrotrogla gen. n.) 3 

2' Hindwing with vein M simple (Fig. 9b). P4 with 2 thin- walled conical 
sensilla (Fig. 3g). Female genitalia (Fig. 8c): Posterior part of subgenital 
plate membranous and almost completely covered ventrally by the 
ovipositor valvulae; spermapore situated at the distal end of a small cap- 
like structure bearing a simple needle-like sclerite (the spermathecal 
duct running through the eye of the needle). Male genitalia (Fig. 9df): 
Phallosome with a pair of relatively broad sclerites, each bearing a 
narrow internal branch, these branches posteriorly convergent or fused 
to form a median aedeagal arch {Sensitibilla Lienhard, 2000) 5 

3(2) Head with striking dark brown colour pattern (Fig. 5a) . . Afrotrogla oryx sp. n. 

3' Head pattern different or head uniformly medium brown 4 

4(3') Head pattern as in Fig. 6a. Relatively small species: Body length and 
forewing length about 3 mm, hindtibia length 1.2 mm. Compound eyes 
relatively large: IO/D 2.6 Afrotrogla maraisi sp. n. 

4' Head uniformly medium brown. Large species: Body length and 
forewing length about 4 mm, hindtibia length 2.2 mm. Compound eyes 
relatively small: IO/D 3.4 Afrotrogla fab ella sp. n. 

5(2') Male (unknown in S. strinata) 6 

5' Female (unknown in S. roessingensis) 7 

6(5) Small species: Hindtibia length 1.0 mm. Abdomen white, lacking 
hypodermal pigmentation. Phallosome as in Fig. 9d 
Sensitibilla brandbergensis sp. n. 

6' Slightly larger species: Hindtibia length 1.3 mm. Abdomen with some 
brown hypodermal pigmentation. Phallosome as in Fig. 9f 
Sensitibilla roessingensis sp. n. 

7(5') Small species: Hindtibia length 1 .0 mm. Abdomen white, lacking hypo- 
dermal pigmentation. Female genitalia (Fig. 8ac): Subgenital plate 
entirely membranous (except for sclerotized rims in zone of fusion with 
ovipositor valvulae), its posterior part triangular, with bluntly pointed 
apex; ovipositor valvula laterally articulated near an tero ventral angle of 
clunium, anterior margin of clunium prolonged into a broad ventral fold 
on the valvula; distal process of ovipositor valvula with only one spine- 
like seta in apical half apart from the spine on its tip; sclerotized plate on 
spermathecal wall large (greatest width about 500 pirn) 

Sensitibilla brandbergensis sp. n. 

Larger species: Hindtibia length 1.5 mm. Abdomen with some brown 
hypodermal pigmentation. Female genitalia: Anterior part of subgenital 
plate with a bilaterally symmetrical sclerified area connected to 
anteroventral angle of clunium; membranous posterior lobe of sub- 
genital plate apically rounded; ovipositor valvula laterally articulated at 
posteroventral angle of clunium, anterior margin of clunium not 



NEW AFRICAN SPELEKETORINAE 445 

prolonged onto the valvula; distal process of ovipositor valvula with 
3-4 spine-like setae in apical half apart from the spine on its tip; 
sclerotized plate on spermathecal wall much smaller (greatest width 
about 250 pm) Sensitibilla strinata Lienhard, 2000 

Sensitibillini trib. n. 

Diagnosis: Belonging to the subfamily Speleketorinae of the Prionoglarididae 
as defined by Lienhard (2004). Habitus similar to Speleketor (Fig. la). Hindwing (Figs 
4b, 9b) with Rs simple and M 2-branched or simple. Forefemur lacking a longitudinal 
row of short articulated spines on anterior face. Pretarsal claws with distinct preapical 
tooth (Fig. 4d). Trichobothrial pattern on legs (see Lienhard, 2000: figs 17-21): 
Foretibia and midtibia with two external trichobothria in nymphs, and adults; hindtibia 
with two such trichobothria in nymphs, usually the proximal one not differentiated in 
adults; hindtarsus with one trichobothrium on second article in nymphs and adults 
(PI. 1); coxa, trochanter and femur of all legs and tarsus of foreleg and midleg lacking 
trichobothria in nymphs and adults. P2 with a subbasal sensory spur, P4 with 2-5 thin- 
walled conical sensilla in apical half (Fig. 3e). Tines of lacinial tip well-developed in 
nymphs (Fig. 5f), strongly reduced in adults (Fig. 4ef). Fore wing with a row of acu- 
minate denticles on distal section of vein Sc delimiting the pterostigma basally (see 
Lienhard, 2000: Fig. 2). Paraprocts in both sexes dorsally with a group of several 
relatively short trichobothria inserted in simple pit-like sockets, and with one normal 
seta between them (Fig. 5d). Female genitalia (Fig. 4g, 8c): Ovipositor consisting only 
of a pair of external valvulae, laterally articulated to clunium and ventrobasally fused 
to subgenital plate, each valvula bearing a distal process with a claw-like articulated 
spine at its tip; spermathecal duct long and thin. Male genitalia (Figs 7bc, 9df): 
Phallosome lacking pore-bearing processes, sclerites of phallosome anteriorly open, 
posteriorly convergent or forming a closed aedeagal arch. 

Type genus: Sensitibilla Lienhard. 

Additional genus: Afrotrogla gen. n. 

Discussion: The particular trichobothrial pattern on legs, especially the 
presence of a tarsal trichobothrium, is probably an autapomorpy of this tribe (see also 
General Discussion, below). An even more impressive autapomorphy is the very par- 
ticular structure of external ovipositor valvulae and subgenital plate, basally fused to 
form a functional unit, which is unknown elsewhere in Psocoptera. Two additional 
autapomorphies of this tribe are the presence of a row of acuminate denticles on the 
distal section of Se on the forewing (no such denticles in Speleketor and the 
Prionoglaridinae) and the reduction of the Rs branching in the hindwing (Rs bifurcate 
in all other Prionoglarididae, see Fig. 3hi), while M initially remains 2-branched in the 
hindwing (simple in Sensitibilla), with Ml and M2 originating separately from Rs-M 
fusion, as in Speleketor (Fig. 3i), or from M stem and Rs-M fusion, respectively (M bi- 
furcate in Prionoglaridinae, Fig. 3h). The absence of the pair of posterolateral pore- 
bearing processes of the phallosome can also be interpreted as an autapomorphy of this 
tribe. Such processes are present in Speleketor and the Prionoglaridinae (see Lienhard. 
2004). They have been considered as homologous with the pore-bearing external para- 



446 C. LIENHARD 

meres of other Psocoptera by Mockford (1984). If this homology is correct, then these 
processes belong to the groundplan of the trogiomorphan phallosome and their 
presence in Speleketor and the Prionoglaridinae is a symplesiomorphy. 

Afrotrogla gen. n. 

Diagnosis: Habitus very similar to Sensitibilla and Speleketor (see Fig. la and 
Gurney, 1943: fig. 3). General morphology of female and nymph as described for 
Sensitibilla (see Lienhard, 2000), with the following differences. P4 with 5 thin- walled 
conical sensilla in apical half (Fig. 3def). Hindwing (Fig. 4b) with Rs simple and M 2- 
branched (Ml and M2 originating separately from Rs-M fusion or from M stem and 
Rs-M fusion, respectively). Third article of hindtarsus in females preapically with a 
dorsal pair of long and slender curved hairs (Fig. 4d); the corresponding hairs much 
shorter in males and on foretarsus and midtarsus of both sexes (cf. Fig. 5e). Females 
with some long backwards-directed sternal setae near midline at about basal one third 
of abdomen (Fig. 4c, 6g); these setae absent in males. Ventral half of female paraproct 
with numerous short setae (Fig. 6b), lacking the circular field of long dense pilosity 
present in Sensitibilla (cf. Lienhard, 2000: Fig. 29). Female genitalia (Fig. 4g): 
Posterior part of subgenital plate sclerotized and clearly visible medially between 
ovipositor valvulae; spermapore situated near posterior end of a membranous sac 
bearing a complex scaffolding of sclerotized struts; spermathecal duct long and thin; 
spermatheca thin- walled, its wall with small pores and, near origin of duct, with some 
sclerotized wrinkles (Figs 4g, 6c). General morphology of male as in female (for 
exceptions, see above). Male terminalia: Epiproct and paraproct simple (Fig. 7d), or 
epiproct with prominent ventral lobe (Fig. 5d); hyandrium simple (Fig. 5b); phallo- 
some with a narrow sclerite forming a simple aedeagal arch (Fig. 7bc). 

Type species: Afrotrogla oryx sp. n. 

Etymology: The name is feminine in gender and refers to the African distri- 
bution of the genus and to the cave dwelling habits of its type species (from Greek 
"trogle" = hole, cave). 

Discussion: See General Discussion, below. 

Afrotrogla oryx sp. n. Figs 4, 5 

Type material: NMN. holotype 2 (4 microscopical slides), South Africa, Richtersveld, 
Wondergat (in cave), SE 2816 Bd, 24.LX.1994, leg. E. Marais. - NMN, paratypes: 1 6 allotype 
(1 microscopical slide, rest in alcohol) and 1 nymph (2 microscopical slides), same data as for 
holotype. 

Description: Male and female: See diagnosis of tribe and genus, with the 
following additions. Head yellowish, with striking cuticular colour pattern (Fig. 5a) 
consisting of a dark brown postclypeus and a broad, dark brown transversal band 
running from gena over frons and parts of vertex; labrum medium brown; compound 
eyes black; antenna brown. Prothorax yellowish, except for dark brown pronotum. 
Forelegs entirely yellowish to light brown, midlegs and hindlegs ditto, but with dark 
brown coxae. Pterothorax entirely dark brown. Wings transparent (pterostigma very 
slightly opaque), veins brown. Abdomen dorsally and laterally with much dark brown 



NEW AFRICAN SPELEKETORINAE 



447 





Fig. 1 
Speleketor irwini Mockford, female from Palm Springs, California: (a) Habitus, lateral view, left 
appendages not figured, scale 1 mm. (b) Epiproct and right paraproct. (c) Genitalia, ventral view: 
Ovipositor valvulae with their articulation to clunium, subgenital plate, spermapore, sperma- 
thecal duct and beginning of spermathecal sac (dorsal valvulae and spermafheca represented by 
interrupted lines). 



448 



C.LIENHARD 




Fig. 2 
Speleketor irwini Mockford, male from Palm Springs, California: (a) Hypandrium. (b) Epiproct 
and right paraproct. (c) Phallosome. lateral view, de = ductus ejaculatorius. (d) Ditto, dorsal 
view. 



NEW AFRICAN SPELEKETORINAE 



449 




Fig. 3 
Thin-walled conical sensilla of P4, other pilosity not figured (a-g): (a) Prionoglaris stygia 
Enderlein, female from type locality, (b) Siamoglaris zebrina Lienhard, male holotype. (c) 
Speleketor flocki Gurney, female paratype. (d) Afrotrogla maraisi sp. n., female holotype, apical 
half of both P4. (e) Ditto, left maxillary palp, spine-like sensillum of P2 also figured, (f) 
Afrotrogla f abella sp. n., male holotype, apical half of both P4. (g) Sensitibilla roessingensis 
sp. n., male holotype, with detail of proximal sensillum. - Hindwing (h-i): (h) Prionoglaris 
stygia Enderlein, female from type locality, (i) Speleketor flocki Gurney, female paratype. 



450 C. LIENHARD 

hypodermal pigment, pattern not clearly defined (examined after 11 years in alcohol), 
but essentially consisting of two broad transversal bands; terminalia brown. 

Head capsule with a slightly concave semicircular groove between lateral 
ocelli, passing by the bifurcation point of the epicranial suture; frontal suture weakly 
developed; ocelli arranged in a flat triangle (Fig. 5a); compound eyes small, approxi- 
mately of same size in both sexes (IO/D: 9 2.8, 6 2.9). Antenna thin and apparently 
very long (extrapolated from lengths of basal flagellomeres, see measurements; all 
antennae damaged in the material available), the basal three flagellomeres together 
about equal to body length (the very similar values for the antennae of the holotype, 
both broken after f4, show that probably no asymmetrical regenerative length growth 
occurred, see remark in Lienhard, 2004: 870). Mouthparts generally as described for 
Sensitibilla strinata by Lienhard (2000). All maxillary palps broken in the material 
available. Lacinia with reduced apical tines, outer part of its tip usually slightly 
indented (Fig. 4ef). Venation of forewing typical for the family (Fig. 4a). Hindwing: 
Rl always present; Ml originating from Rs-M fusion (Fig. 4b) or from M stem, 
slightly basally to Rs-M fusion; M2 originating from Rs-M fusion. Spine-like setae on 
tibiae and tarsi with more or less rounded tips. Pearman-organ of hindcoxa not 
differentiated, only a mirror-like membranous area present basally on inner side of 
hindcoxa, both midcoxae with a distinctly prominent hyaline tubercle on inner side. 
Third article of hindtarsus of female subapically with a pair of slender dorsal hairs, 
slightly curved ventrally and longer than length of claw (Fig. 4d). These specialized 
hairs not differentiated in male, there the corresponding setae shorter than the claw 
(cf. Fig. 5e). Proximal trichobothrium of hindtibia usually not differentiated in adults, 
corresponding to the general pattern of leg trichobothria in Sensitibillini, but this 
trichobothrium on one hindtibia of the allotype male also well-developed. Female with 
two tufts of long backwards-directed sternal setae medially at about basal one third of 
abdomen, very close to midline and practically fused to one single tuft (Fig. 4c). No 
such sternal setae in male. 

Female terminalia (Fig. 4g): Epiproct and paraproct simple (cf. Fig. 6b). 
Ovipositor valvula with a long and almost straight distal process bearing a short claw- 
like apical spine and numerous short stout hairs, most of them situated on dorsal side 
of process; basal part of valvula with a transversal row of subapical dorsal setae and 
one long ventral seta on each side of base of distal process; basal part of valvula ex- 
ternally articulated at postero ventral angle of clunium, its internal margin curved and 
well-sclerotized, running parallel to hind margin of subgenital plate, posterointernal 
angles of valvulae almost touching each other medially. Posterior part of subgenital 
plate well-sclerotized but otherwise simple and bare, its anterior margin concave and 
anterolaterally prolonged into a long sclerotized rim fused to inner base of the 
corresponding ovipositor valvula; anterior part of subgenital plate a weakly sclero- 
tized, almost circular plate bearing some sparse setae; transition zone between posteri- 
or and anterior parts of subgenital plate membranous; sclerotized anterior margin of 
posterior part with a group of about a dozen small hairs situated on the slightly hollow 
median zone of this margin (thus their insertion points not visible in ventral view), no 
porus present between these hairs. Spermapore situated near posterior end of an elon- 
gate membranous sac (interrupted line in Fig. 4g), with a small circular sclerite, in situ 



NEW AFRICAN SPELEKETORINAE 



451 




Fig. 4 
Afrotrogla oryx sp. n., female holotype: (a) Forewing, scale 1 mm. (b) Hindwing, same scale. 

(c) Tuft of long backwards-directed sternal setae medially at about basal one third of abdomen. 

(d) Apex of third article of hindtarsus, with dorsal pair of long curved hairs, only one claw 
illustrated, (e) Lacinial tip, left, (f) Ditto, right, (g) Genitalia, ventral view, scale 0.2 mm: 
Ovipositor valvulae with ventrolateral parts of clunium, subgenital plate and, observed in situ 
through ventral membranes, spermapore and spermathecal duct with accessory structures and 
sclerotized wrinkles on spermathecal wall. 

clearly visible through membranous zone just anteriorly to concave margin of posterior 
part of subgenital plate. Spermathecal duct long and thin, somewhat spirally curled, its 
posterior half situated in the above mentioned membranous sac, the latter equipped 



452 C. LIENHARD 

with a complex scaffolding of sclerotized struts. The median paired struts situated 
ventrally on the sac, originating as a simple rod at the spermapore and, towards their 
anterior ends, successively bifurcating into a circular strut forming the anterior border 
of the sac, and into a pair of long lateral struts. The latter backwards-directed, slightly 
curved and with free posterior ends (probably movable, cf. Figs 4g and 6d). Sperma- 
thecal wall, near origin of duct, with some bilaterally symmetrical, sclerotized 
wrinkles. One spermatophore (sperm packet) clearly visible in the spermatheca of the 
holotype, simple, slightly pear shaped, but much more elongate than the sperma- 
tophores of Sensitibilla strinata illustrated by Lienhard (2000: fig. 25). 

Male terminalia: Epiproct simple, paraproct with a ventrally prominent pos- 
terior lobe (Fig. 5d). Hypandrium (Fig. 5b) with a characteristic sclerotized pattern. 
Phallosome developed as a membranous sac (cf. Fig. 7c), dorsally with a relatively 
long but simple, arched and posteriorly truncate aedeagal sclerite (Fig. 5c). 

Nymph: One nymph (probably male) examined, late instar (see Discussion, 
below). See diagnosis of tribe and genus, with the following additions. Body colour 
entirely yellowish white (adult head pattern not yet recognizable), except for dark 
pigment in compound eyes and ocellar dots, and two large brown transversal bands on 
abdominal tergites. General morphology as desribed for the nymph of Sensitibilla 
strinata by Lienhard (2000). Both maxillary palps broken. Lacinia with three well- 
developed apical tines (Fig. 5f) (one lacinia damaged). Terminal article of hindtarsus 
subapically without a pair of long curved hairs (Fig. 5e) (see Discussion, below). 

Measurements: Female holotype. BL = 3.1 mm; FW = 4.0 mm; HW = 1 .9 mm; 
F = 1030 //m; T = 1750 //m; tl= 630 //m; t2 = 140 //m; t3 = 175 pm; flagellomeres 
(left/right): fl = 900/870 ^m; f2 = 1040/960 pm; f3 = 1220/1200 pm. - Male allotype. 
BL=3.6mm;FW = 3.7mm;HW = 1.8 mm; F = 1020 //m; T= 1150pm; tl= 610/im; 
t2= HOpm; t3 = 165 pm. 

Etymology: The specific epithet is a noun in apposition alluding to the South 
African Oryx antelope (Oryx gazella gazella) which has a somewhat similar dark 
brown colour pattern on its head. 

Discussion: The type locality of Afrotrogla oryx, the Wondergat cave, is located 
in the arid northwestern corner of South Africa, north of Springbok and close to the 
Namibian border, quite far away from the type localities of the other known species of 
Sensitibillini in western, central or northeastern Namibia. This is the only species of 
the genus of which both sexes are known. Males of A. oryx and A.fabella are very 
similar in their phallosome morphology, but these species can easily be distinguished 
by paraproct shape, body size and especially by the colour pattern of the head. The very 
complex morphology of female genitalia allows a more detailed comparison of A. oryx 
with A. marnisi, the latter only known from the female holotype collected in central 
Namibia (Windhoek). The striking differences in the structure of the subgenital plate 
indicate that these species are only distantly related (see Discussion of A. marnisi). 

The absence of the dorsal pair of long curved preapical hairs in the male, 
present on female hindtarsus, corresponds to the situation observed in the nymph. 
Therefore the latter is believed to be a male nymph, because it seems likely that these 
specialized hairs are present in female nymphs, rather than being generally absent in 
nymphs and restricted to adult females. 



NEW AFRICAN SPELEKETORINAE 



453 




Fig. 5 
Afrotrogla oryx sp. n., male allotype (a-d): (a) Head, frontal view, (b) Hypandrium, with left 
attachment to clunium. (e) Sclerite of phallosome, scale 0.2 mm. (d) Epiproct and left paraproct. 
- Afrotrogla oryx sp. n., nymph (e-f): (e) Hindtarsus, t2 with trichobothrium and apical part of 
tl, only one claw illustrated, (f) Lacinial tip. 



Afrotrogla maraisi sp. n. Figs 3 de, 6 

Type material: NMN, holotype 9 (3 microscopical slides, rest in alcohol), Namibia, 
Windhoek, 10.X.1995, leg. E. Marais. 

Description: Female (male unknown): See diagnosis of tribe and genus, with 
the following additions. Head generally light brown, the following areas with reddish 



454 C. LIENHARD 

brown hypodermal pigment (Fig. 6a): Gena and base of mandible; a triangular arrow- 
like patch dorsally between lateral ocellus and compound eye pointing towards anterior 
end of vertical suture; a small longitudinal patch near anterior end of vertical suture. 
Compound eyes black, antenna and maxillary palp light brown. Prothorax light brown, 
pterothorax medium brown, legs yellowish to light brown. Wings transparent (ptero- 
stigma very slightly opaque), veins medium brown. Abdomen yellowish, with much 
dark brown hypodermal pigment, pattern not clearly defined (examined after 10 years 
in alcohol), but apparently consisting of some segmental patches and transversal bands; 
terminalia light to medium brown. 

Shape of head capsule (Fig. 6a) similar to that of A. oryx, frontal suture reduced, 
ocelli arranged in a flat triangle (Fig. 6a), compound eyes small (IO/D 2.6). Antenna 
thin and distinctly longer than forewing. One antenna of the holotype apparently 
complete (i.e. last flagellomere slightly tapering to a regularly rounded tip, bearing a 
conspicuous terminal sensillum), 11 -segmented, the other one broken after f4 (the 
considerably higher values for lengths of f 1 to f3 for the latter may indicate some 
regenerative length growth, see remark in Lienhard, 2004: 870). Mouthparts generally 
as described for Sensitibilla strinata by Lienhard (2000). Maxillary palp long and 
slender, P4 with five thin-walled conical sensilla in apical half, their arrangement 
somewhat variable (Fig. 3de). Lacinia and wing venation very similar to that of A. 
oryx. Hindwing: Rl present, Ml and M2 originating separately from Rs-M fusion. 
Spine-like setae on tibiae and tarsi with more or less rounded tips. Pearman-organ of 
hindcoxa not differentiated, only a mirror-like membranous area present basally on 
inner side of hindcoxa, both midcoxae with a distinctly prominent hyaline tubercle on 
inner side. Trichobothrial pattern on legs typical for Sensitibillini. The dorsal pair of 
long curved hairs subapically on third article of hindtarsus well-differentiated, longer 
than claw. Two well-separated tufts of long backwards-directed sternal setae present 
near midline at about basal one third of abdomen (Fig. 6g). 

Female terminalia: Epiproct and paraproct simple (Fig. 6b). Ovipositor valvula 
(Fig. 6e) with a long, relatively broad and slightly curved distal process bearing a short 
claw-like apical spine and numerous spine-like setae, most of them situated on dorsal 
side of the process and three on its external margin; basal part of valvula with a trans- 
versal row of subapical dorsal setae and one long ventral seta on each side of base of 
distal process; basal part of valvula externally articulated at postero ventral angle of 
clunium, its internal margin only slightly curved and not sclerotized in apical part, 
running parallel to lateral hindmargin of subgenital plate, but the two valvulae not 
touching each other medially. Posterior part of subgenital plate (Fig. 6e) well-sclero- 
tized and bearing a few short hairs, laterobasally fused to inner base of valvulae but not 
prolonged into a long sclerotized rim; anterior part of subgenital plate membranous 
except for a narrow sclerotized longitudinal area just anteriorly to the middle of the 
convex anterior margin of its posterior part. This margin laterally prolonged into a pair 
of short curved stylets, ventrally prominent and backwards-directed, the right stylet 
bearing a short fine hair on its rounded tip, the left one bare. Sclerotized part of sub- 
genital plate perforated by a conspicuous porus in the middle of its anterior margin, 
between the bulged bases of the stylets; this porus opening to a sclerotized, digitiform, 
backwards-directed invagination of the subgenital plate (interrupted line in Fig. 6f); 



NEW AFRICAN SPELEKETORINAE 



455 




Fig. 6 
Afrotrogla maraisi sp. n., female holotype: (a) Head, frontal view, (b) Epiproct and right para- 
proct. (c) Origin of spermathecal duct and sclerotized wrinkle on spermathecal wall, (d) 
Spermapore and posterior part of spermathecal duct, with accessory structures, ventral view, 
scale 0.2 mm. (e) Subgenital plate, ovipositor valvulae and ventrolateral parts of clunium, ventral 
view, same scale as Fig. 6d. (f) Detail of anteromedian porus on subgenital plate, compare with 
Fig. 6e. (g) Pair of tufts of long backwards-directed sternal setae at about basal one third of 
abdomen, middle of sternite shown by an interrupted line. 



456 C. LIENHARD 

poms surrounded by heavy sclerotization and laterally flanked by some short stout 
setae (Fig. 6f), some very fine hyaline hairs (length 3-10 pirn) also visible inside the 
lumen of the porus (only observable at high magnification). Spermapore with a longi- 
tudinally oval annular sclerite (Fig. 6d), in situ situated just dorsally of the digitiform 
invagination of the subgenital plate, near the posterior end of a rounded membranous 
sac; spermathecal duct of about the same relative length as in A. oryx (see Fig. 4g), 
somewhat spirally curled, its posterior part situated in the above mentioned membra- 
nous sac, the latter equipped with a complex scaffolding of sclerotized struts showing 
the same elements as in A. oryx (cf. Figs 6d and 4g). Median paired ventral struts 
bifurcate posteriorly and not in contact with spermapore sclerite, circular strut 
(interrupted line in Fig. 6d) dorsomedially prolonged by a backwards directed process 
(NB. Only a small rudiment of such a process visible in A. oryx, see Fig. 4g), lateral 
struts straight, only slightly backwards-directed, probably movable. Spermathecal 
wall, near origin of duct, with a slightly undulated sclerotized wrinkle (Fig. 6c). 
Several spermatophores (sperm packets) visible inside the spermatheca of the holo- 
type, pear-shaped, very similar to the spermatophores of Sensitibilla strinata illustrated 
by Lienhard (2000: fig. 25). 

Measurements: Female holotype. BL = 2.9 mm; FW = 3.0 mm; HW = 1 .6 mm; 
F = 730 //m; T = 1230 pirn; tl= 430 //m; t2 = 113 //m; t3 = 145 pirn; flagellomeres 
(left/right): fl = 650/740 pirn; Î2 = 660/800 //m; f3 = 610/750 pm (left antenna 
complete, 11-segmented, total length 3.9 mm). 

Etymology: The species is dedicated to Dr Eugene Marais, curator of the 
entomological collections of the National Museum Namibia and collector of all 
currently known specimens of the genus Afrotrogla. 

Discussion: The type of Afrotrogla maraisi was collected in Windhoek (central 
Namibia), but unfortunately no further information on its habitat is available 
(collecting by trap?). The species can easily be distinguished from A. oryx by its head 
pattern and by the structure of its genitalia: Different shape and pilosity of ovipositor 
valvula; different shapes of spermapore sclerite, of membranous sac around posterior 
part of spermathecal duct and of its accessory sclerites; strikingly different structure of 
subgenital plate. The presence, on the subgenital plate of A. maraisi, of a basal porus 
with a digitiform invagination and a pair of lateral stylets (compared with the simple 
subgenital plate of A. oryx), indicates that these species are only distantly related to 
each other and could even belong to different genera. At present their generic 
separation would of course be premature in view of the poorly known species diversity 
in Sensitibillini. For some remarks on functional morphology see General Discussion 
(below). 

It seems highly unlikely that A. f abella (male described below, female 
unknown) could be the unknown male of A. maraisi. As shown by A. oryx and by 
Sensitibilla brandbergensis , there exists no significant sexual dimorphism concerning 
general body size, wing length, IO/D index or colouration of head in Sensitibillini. 
A. maraisi (female) and A. f abella (male) clearly differ in all these characters (see key, 
above). 



NEW AFRICAN SPELEKETORINAE 457 

Afrotrogla f abella sp. n. Figs 3f , 7 

Type material: NMN, holotype 3 (4 microscopical slides), Namibia, Grootfontein 
District, Märchenhöhle (in cave), SE 1917 Cb, 30.VIII.1990, leg. E. Marais. 

Description: Male (female unknown): See diagnosis of tribe and genus, with 
the following additions. Head uniformly medium brown, no hypodermal pigment 
pattern visible (examined after 15 years in alcohol, but abdominal hypodermal pigment 
perfectly preserved). Compound eyes black, antenna medium brown, maxillary palp 
light brown. Prothorax light brown, pterothorax dorsally dark brown, legs light to 
medium brown. Wings transparent (pterostigma very slightly opaque), veins dark 
brown. Abdomen yellowish, with characteristic dark brown hypodermal pigment 
pattern, especially well-developed dorsally in posterior two thirds of abdomen 
(Fig. 7a), terminalia dark brown. 

Head capsule slightly concave between lateral ocelli, frontal suture weakly 
developed, ocelli arranged in a flat triangle, almost in a line. Compound eyes very 
small (IO /D 3.4). Antenna thin and apparently very long (extrapolated from lengths of 
basal flagellomeres, see measurements; both antennae damaged in the holotype, one of 
them broken after f3, the other after f6), the basal three flagellomeres together about 
equal to body length (the very similar values for both antennae of the holotype show 
that probably no asymmetrical regenerative length growth occurred, see remark in 
Lienhard, 2004: 870). Mouthparts generally as described for Sensitibilla strinata by 
Lienhard (2000). Maxillary palp long and slender, P4 with five thin- walled conical 
sensilla in apical half, their arrangement somewhat variable (Fig. 3f). Lacinia and wing 
venation very similar to that of A. oryx. Hindwing: Rl present. Ml and M2 originating 
separately from Rs-M fusion. Spine-like setae on tibiae and tarsi with more or less 
rounded tips. Pearman-organ of hindcoxa not differentiated, only a mirror-like 
membranous area present basally on inner side of hindcoxa, both midcoxae with a 
distinctly prominent hyaline tubercle on inner side. Trichobothrial pattern on legs 
typical for Sensitibillini. Third article of hindtarsus broken on both hindlegs. 
Abdominal sternites with transversal rows of few short hairs, no tuft(s) of long sternal 
setae present in basal half of abdomen. 

Male terminalia: Epiproct simple, paraproct with some sclerotized wrinkles 
near group of trichobothria, without prominent posteroventral lobe (Fig. 7d). 
Hypandrium simple, similar in shape to that of A. oryx, but entirely brown (cuticular 
pigmentation) except for narrow hyaline hindmargin. Phallosome a membranous sac 
(Fig. 7c), dorsally with a simple, arched and posteriorly rounded aedeagal sclerite 
(Fig. 7b) (NB. The extent of the anterior opening of the aedeagal arch depends on the 
degree of squashing of the slide-mounted phallosome). 

Measurements: Male holotype. BL = 4.2 mm; FW = 4.1 mm; HW = 2.2 mm; 
F = 1250 piva; T = 2220 /*m; tl = 850 /<m; t2 = 165 /<m; t3 broken on both hindlegs; 
flagellomeres (left/right): fl = 1420/1 370 //m; f2 = 1530/1410 /^m; f3 = 15 10/1420 //m. 

Etymology: The specific epithet refers to the type locality, a cave with the 
German name Märchenhöhle ("fairy-tale cave"); it is a noun in apposition (from Latin 
"f abella" = Märchen, fairy-tale). 



458 



C. LIENHARD 




Fig. 7 
Afrotrogla fabella sp. n., male holotype: (a) Colour pattern of abdomen, dorsal view, (b) Sclerite 
of phallosome, scale 0.2 mm. (c) Phallosome. with membranous parts asymmetrically deformed 
by slide mounting, dorsal view, de = ductus ejaculatorius. (d) Epiproct and right paraproct. 



Discussion: The type locality of Afrotrogla fabella is situated in northeastern 
Namibia (19°32'S 17°14'E), far away from Windhoek and the South African 
Wondergat cave, from where the two other species of the genus are known. For 
distinction from these species see the corresponding discussions and the key (above). 

Sensitibilla Lienhard 
Lienhard, 2000: 872. 

Diagnosis: See Lienhard (2000) with the following additions. Hindwing with 
veins Rs and M simple, Rl usually present, rarely absent (see Fig. 9b and Lienhard, 
2000: Figs 3, 4). No long curved dorsal hairs subapically on third article of hindtarsus 
(Fig. 8b). P4 with 2 thin-walled conical sensilla in apical half (Fig. 3g). Both sexes 
without particularly long backwards-directed sternal setae near midline in basal half of 
abdomen. Female genitalia (Fig. 8c): Spermapore situated at the posterior end of a 
small cap-like structure, bearing a simple needle-like accessory sclerite (the sperma- 
thecal duct running through the eye of the needle). General morphology of male as in 
female. Male terminalia: Epiproct and paraproct simple (Fig. 9e); hypandrium simple 
(Fig. 9c); phallosome lacking pore-bearing processes, with a pair of anteriorly 
divergent and relatively broad sclerites, each bearing a narrow posterointernal branch, 
these branches convergent (Fig. 9f) or fused to form a median aedeagal arch (Fig. 9d). 

Type species: Sensitibilla strinata Lienhard. 

Discussion: See General Discussion, below. 



NEW AFRICAN SPELEKETORINAE 459 

Sensitibilla strinatii Lienhard Plate 1 

Lienhard, 2000: 874. 

Material Examined: MHNG, 9 holotype, 29 and 7 nymphs paratypes, Namibia, 
Arnhem Cave ( 1 24 km SE of Windhoek) , in cave , 2 1 .X . 1 999 , leg . P. Strinati . 

Description: Female (male unknown). See original genus and species descrip- 
tion, key (above) and extended generic diagnosis (above). 

Discussion: See Discussion of S. brandbergensis and S. roessingensis (below). 

Sensitibilla brandbergensis sp. n. Figs 8, 9a-d 

Type material: NMN, holotype S (2 microscopical slides), Namibia, Brandberg, 
Wasserfallfläche, 1980m, 21°13.5'S 14°31.1'E, 10-12.XI.1998, Malaise trap, river bed, leg. 
A. H. Kirk-Spriggs. - NMN, paratypes: 19(1 microscopical slide, rest- in alcohol), Namibia, 
Brandberg, Wasserfallfläche, 1960m, 21°10.77'S 14°32.87 , E, 7-10.IV.1999, Malaise trap, well- 
vegetated valley below waterfall, bushy Karoo-Namib shrubland (NA99-M05), leg. S. van Noort 
& S. G. Compton. 19 (alcohol), Namibia, Brandberg, Messum Valley, 700m, 21°13.29'S 
14°30.98'E, 5-17.IV.1999, Malaise trap, bushy Karoo-Namib shrubland (NA99-M12), leg. S. 
van Noort & S. G. Compton. - MHNG, paratypes: 9 allotype (2 microscopical slides), Namibia, 
Brandberg, plateau, 1960m, 21°10'50"S 14°32'50"E, 19-21.10.1998, yellow pan trap, leg. R. 
Butlin & J. Altringham. 1 9 (2 microscopical slides), Namibia, Brandberg, Hungorob Valley, 
1180m, 21°1140'S 14°31.69'E, 5-16.IV.1999, Malaise trap, bushy Karoo-Namib shrubland 
(NA99-M10), leg. S. van Noort & S. G. Compton. 

Description: Male and female: See diagnosis of tribe and genus, with the 
following additions. General colouration yellowish white to very light brown. 
Compound eyes black. Wings transparent, veins light brown. Abdomen lacking hypo- 
dermal pigment (examined after 6-7 years in alcohol); terminalia yellowish to very 
light brown. 

General morphology as described for S. strinatii (see Lienhard, 2000). Head 
capsule slightly concave between lateral ocelli, compound eyes small (IO/D: â 3.0, 9 
2.6-2.9). Antenna thin and apparently very long (extrapolated from lengths of basal 
flagellomeres, see measurements; all antennae damaged in the material available), the 
basal three flagellomeres together somewhat shorter than body length (very similar 
values for the antennae of the allotype, broken after f6 and f4, show that probably no 
asymmetrical regenerative length growth occurred, see remark in Lienhard, 2004: 
870). P4 with two simple, thin- walled conical sensilla in apical half. Wing venation as 
in Fig. 9ab, Rl present in hindwing. Spine-like setae on tibiae and tarsi with distinctly 
rounded tips. Pearman-organ of hindcoxa not differentiated, both midcoxae with a dis- 
tinctly prominent hyaline tubercle on inner side. Trichobothrial pattern on legs typical 
for Sensitibillini. 

Female terminalia (Fig. 8ac): Epiproct and paraproct as in S. strinatii 
(Lienhard, 2000: Fig. 29), the circular field of dense and long pilosity on paraproct 
well-developed. Subgenital plate entirely membranous, except for sclerotized rims in 
zone of fusion with ovipositor valvulae; posterior part of subgenital plate triangular, 
with subacute apex, almost completely covered by basal parts of ovipositor valvulae, 
the latter touching each other medially. Ovipositor valvula laterally articulated near 
anteroventral angle of clunium, ventromedially fused with subgenital plate in a sclero- 
tized rim; anterior margin of clunium prolonged into a broad membranous ventral fold 



460 



C. LIENHARD 




Plate 1 
Sensi tibi Ila strinata Lienhard, female paratype, hindleg. (A) Second article of hindtarsus with 
trichobothrium situated dorsally on slightly thickened central zone (dorsolateral view, base of 
article near bottom of photo); the trichobothrium is actually an erect filiform hair (cf. Fig. 5e), 
its artificially curved appearance on the photo results from the treatment of the leg for SEM 
examination. (B) Base of trichobothrium of hindtarsus with adjacent setae and campaniform 
sensillum, same view as A. (C) Ditto, dorsal view. (D) Socket of tibial trichobothrium, sensillum 
broken but insertion point visible. SEM micrographs made by J. Wuest (MHNG). 



on the valvula. Distal process of ovipositor valvula slightly curved, with a long claw- 
like apical spine, one external spine-like seta in distal half and several such setae in 
basal half, two or three of them on ventral side of the process. Basal part of valvula 
with a transversal row of subapical dorsal setae and some ventral setae, a particularly 
long one near rounded posterointernal margin. Spermapore situated in a small mem- 
branous posteriorly rounded cap, spermathecal duct running through the eye of a 
simple needle-like acessory sclerite, duct relatively long but not distinctly spirally 
curled. Spermathecal wall, near origin of duct, with a large kidney- shaped, weakly 



NEW AFRICAN SPELEKETORINAE 



461 




Fig. 8 
Sensitibilla brandbergensis sp. n., female: (a) Spermatheca containing two spermatophores, with 
kidney-shaped sclerotized area of spermathecal wall, spermathecal duct not illustrated, (b) Apex 
of third article of hindtarsus, only one claw illustrated, (c) Genitalia, ventral view, scale 0.2 mm: 
Ovipositor valvulae with ventrolateral parts of clunium, subgenital plate and, observed in situ 
through ventral membranes, spermapore and spermathecal duct with accessory structures and 
kidney-shaped sclerotized area of spermathecal wall (compare with Fig. 8a). 



sclerotized plate (its greatest width about 500 ptm) sparsely covered by very fine 
denticles and bearing a pair of heavily sclerotized teeth directed to the lumen of the 
spermatheca and connected to each other by a narrow somewhat sclerotized band (in 



462 



C. LIENHARD 




Fig. 9 
Sensi tibi I la brandbergensis sp. n., male holotype (a-d): (a) Forewing, scale 1 mm. (b) Hindwing, 
same scale, (c) Hypandrium. (d) Phallosome, ventral view, scale 0.2 mm. - Sensitibilla roessin- 
gensis sp. n., male holotype (e-f): (e) Epiproct and left paraproct. (f) Phallosome, ventral view, 
same scale as Fig. 9d. 



one paratype female one of these teeth not developed). Spermatophores elongately 
pear-shaped (Fig. 8a). 

Male terminalia: Epiproct and paraproct simple (cf. Fig. 9e). Hypandrium 
simple and almost colourless (Fig. 9c). Phallosome (Fig. 9d) with a pair of foliaceous 
scleri tes, anteriorly divergent, much enlarged in middle, posteriorly each with a well- 
developed lateral lobe and a narrow internal branch; these branches medially fused and 
forming a narrow aedeagal arch. 



NEW AFRICAN SPELEKETORINAE 463 

Measurements: Male holotype. BL = 1 .4 mm (strongly shrunk); FW = 2.6 mm; 
HW= 1.4mm;F = 600^m;T= 1000 //m;tl = 350/<m;t2 = 100/im;t3= 125/im.- 
Female allotype. BL = 2.3 mm (somewhat expanded); FW = 2.7 mm; HW = 1.5 mm; 
F = 630 pm; T = 1040 pm; tl = 350 pm; t2 = 105 pm; t3 = 125 pm; flagellomeres 
(left/right): fl = 490/480 pm; f2 = 510/535 pirn; f3 = 525/550 /im. 

Etymology: The specific epithet refers to the type locality, Brandberg, the 
highest mountain massif of Namibia. 

Discussion: S. brandbergensis is easy to distinguish from S. strinata and 
S. roessingensis by its smaller size, by the almost complete absence of any pigmenta- 
tion and, above all, by the very characteristic genitalia of both sexes (see also key, 
above). The type locality of S. brandbergensis, the Brandberg massif, is situated in 
western Namibia, north of the town Swakopmund. S. brandbergensis has not been col- 
lected in caves, as were most of the other Sensitibillini treated in this study, but in semi- 
desertic shrubland, in Malaise traps and once in a yellow pan trap. Nothing is known 
on its biology and it is not unlikely that these minute insects live in soil crevices, from 
where they regularly come out during the night. The almost complete absence of any 
pigmentation also points towards such an edaphico-nocturnal mode of life, which is not 
fundamentally different from life in caves. 

Sensitibilla roessingensis sp. n. Figs 3g, 9ef 

Type material: NMN, holotype 6 (3 microscopical slides), Namibia, Swakopmund 
District, Rössing Wes Cave (in cave), SE 2214 Db, 10. Vili. 1995, leg. E. Marais. 

Description: Male (female unknown): See diagnosis of tribe and genus, with 
the following additions. General colouration yellowish to light brown. Compound eyes 
black. Wings transparent, veins light brown. Abdomen with some brown hypodermal 
pigment, pattern not clearly visible (examined after 10 years in alcohol), but essentially 
consisting of some segmental transverse bands; terminalia yellowish to light brown. 

General morphology as described for S. strinata (see Lienhard, 2000). 
Compound eyes small (IO/D 2.9). Antennae broken after fl and f2. P4 with two simple 
thin-walled conical sensilla in apical half (Fig. 3g). Both forewings damaged, Rl 
present in both hindwings. Hindwing relatively long (1 .3x greatest width of forewing). 
Spine-like setae on tibiae and tarsi with distinctly rounded tips. Pearman-organ of hind- 
coxa not differentiated, both midcoxae with a distinctly prominent hyaline tubercle on 
inner side. Trichobothrial pattern on legs typical for Sensitibillini. 

Male terminalia: Epiproct and paraproct simple (Fig. 9e). Hyandrium simple 
and almost colourless (cf. Fig. 9c). Phallosome (Fig. 9f) with a pair of somewhat 
spindle-shaped sclerites, anteriorly divergent, posteriorly each with a weakly 
developed rounded lobe and a narrow internal branch; these branches curved towards 
midline of phallosome apex but not fused into a continuous aedeagal arch, medioapical 
lobe of phallosome deeply notched. 

Measurements: Male holotype. BL = 2.2 mm; HW = 1 .6 mm; F = 720 pm; T = 
1260 pm; tl = 470 pm; t2 = 125 pm; t3 = 150 pm; flagellomeres (left/right): fl = 
540/530 pm; il = - /540 pm. 



464 C. LffiNHARD 

Etymology: The specific epithet refers to the type locality, Rössing Wes Cave. 

Discussion: The type locality of Sensitibilla roessingensis is situated in western 
Namibia, on the Namib Desert plains, about 30 km from the town of Swakopmund. It 
seems rather unlikely that S. roessingensis could be the unknown male of S. strinata, 
described from Arnhem Cave in central Namibia. The specific diversity observed in 
Sensitibillini suggests that these isolated and widely separated populations belong to 
different species, even if they are closely related in view of their general morphology. 
S. roessingensis is apparently slightly smaller than S. strinata (T = 1 .5 mm in the latter) 
but has relatively long hindwings (in S. strinata hindwing length only very slightly 
exceeding greatest width of forewing). The forewing length of these species could not 
be compared, because both forewings are damaged in the holotype of S. roessingensis. 
In the holotype of S. strinata the proximal P4 sensillum is slightly lobate (Lienhard, 
2000: Fig. 8), in the holotype of S. roessingensis it is simple (detail in Fig. 3g), but this 
difference is of doubtful taxonomic value. An additional argument not to assign tenta- 
tively the male from Rössing cave to S. strinata is the fact that the latter is the type 
species of the genus. The definition of the genus could become problematic due to such 
an unclear concept of the type species. 

GENERAL DISCUSSION 
Phylogenetics 

Due to the symplesiomorphic presence of an archaic wing venation, the extant 
Prionoglarididae may be considered as "living fossils" (see Introduction). However, 
the different clades of this family are characterized by striking autapomorphies and 
surprising specializations unknown elsewhere in Psocoptera. Characters of Priono- 
glaridinae have been treated by Lienhard (1988, 1998, 2004). In the following some 
characters of Speleketorinae are discussed with regard to the phylogeny of this sub- 
family. 

The presence of trichobothria on legs, unique in Psocoptera (see also below), 
has been considered as an important synapomorphy of Speleketorinae (see Lienhard, 
2000, 2004), but it may also be interpreted as an adaptive homoplasy related to the 
particular biology of these generally cavernicolous psocids. However, the monophyly 
of Speleketorinae is also supported by molecular data (see Yoshizawa et al., 2006) and 
by the slight but almost identical extent of metamorphosis of their mouthparts, in 
particular the lacinia, during adult moult (see Lienhard, 2004). Therefore the presence 
of leg trichobothria is here also considered as an autapomorphy of this subfamily. Each 
tribe is characterized by a particular arrangement of trichobothria on the leg segments 
(see key above and Lienhard, 2000: figs 17-24). Most elements of the characteristic 
trichobothrial pattern in adults are already present in nymphs (at least in later instars, 
first instars not examined) so that these can easily be assigned to a tribe, in spite of the 
very similar habitus of all prionoglaridid nymphs. It is difficult to decide what kind of 
trichobothrial pattern should be considered as plesiomorphic within the subfamily. The 
presence of a tarsal trichobothrium in Sensitibillini, unique in insects (see below), may 
be an indication that the Sensitibillini pattern has to be considered as apomorphic 
compared to the Speleketorini pattern. However, each pattern could be an autapo- 



NEW AFRICAN SPELEKETORINAE 465 

morphy of the respective tribe with regard to an unknown initial pattern of their 
common ancestor. It is interesting to see that in both clades, Speleketorini and 
Sensitibillini, 2-3 trichobothria per leg are generally present; in Sensitibillini they have 
"migrated" to the distal segments (tibiae and hindtarsus), whereas in Speleketorini they 
can be found on the more proximal segments (trochanters and femora). In bugs 
(Hemiptera: Heteroptera), where leg trichobothria have almost certainly evolved 
several times independently, they are present on coxae, trochanters and femora (Schuh, 
1975) (occasionally trichobothria-like setae also occur on tibiae in some semiaquatic 
bugs, see Moller Andersen, 1982). According to Schuh (1975) the Heteroptera, like the 
Arachnida, seem to be disposed to the multiple evolution of these structures, whereas 
leg trichobothria are unknown elsewhere in insects. The example of the Speleketorinae 
shows that leg trichobothria have also evolved in some basal Psocodea, the sister-group 
of the Condylognatha (i.e. Thysanoptera and Hemiptera) within the Paraneoptera (see 
Yoshizawa & Saigusa, 2001). See also discussion on Funcional Morphology (below). 

The Speleketorinae (or perhaps the Prionoglarididae in general) could also be 
characterized by a reduced number of antennal segments. The normal trogiomorphan 
antenna has more than 20 segments (i. e. scapus, pedicellus and at least 18 flagello- 
meres); however, a slightly reduced number (19 segments) has been mentioned by 
Yoshizawa et al. (2006) for an undescribed species of Psyllipsocidae. In 
Prionoglarididae the antennae are always very thin and long and therefore broken in 
most of the material available. I am not aware of any published number of antennal 
segments of a complete antenna in Prionoglaris, and I could not find any complete 
antenna in the material of this genus at my disposal (MHNG). In Siamoglaris the 
antenna is more than 10-segmented (Lienhard, 2004: 870) but the exact number of 
segments remains unknown. Mockford (1984: 170) observed three complete antennae 
with a variable number of segments (7, 10, 15) in Speleketor. Lienhard (2000: 874) 
reported one complete 13-segmented antenna in a female of Sensitibilla strinata (i.e. 
last flagellomere slightly tapering to a regularly rounded tip, bearing a conspicuous 
terminal sensillum). In Afrotrogla maraisi one complete antenna with 11 segments has 
been mentioned in the above description. Maybe the reduced and apparently variable 
number of antennal segments could also constitute an autapomorphy of the subfamily 
Speleketorinae, or even of the whole family Prionoglarididae. However, more flagel- 
lomere counts of undoubtedly complete antennae are needed to confirm the taxonomic 
value of this character. 

The Sensitibillini are characterized by the synapomorphic structure of their 
female ovipositor, i.e. external valvulae with a strong distal process and ventrobasal- 
ly fused to subgenital plate to form a functional unit (see also below). The genus 
Speleketor has relatively plesiomorphic female genitalia, as described by Yoshizawa et 
al. (2006) for Trogiomorpha. However, this genus is characterized by the autapo- 
morphic structure of the phallosome, having its basal sclerites anteriorly fused (phal- 
losome anteriorly open in Sensitibillini, as usual in the suborder Trogiomorpha), and 
by the autapomorphic presence of a row of small spines on anterior face of the fore- 
femur. Another possible autapomorphy of the Sensitibillini is the absence of the pore- 
bearing processes of the phallosome, which probably belong to the groundplan of the 
Trogiomorpha (see Discussion under Sensitibillini above, where two additional wing 
characters of minor importance are also discussed). 



466 C. LEENHARD 

Within Sensitibillini, the genus Afrotrogla is characterized by highly complex 
structures associated with the spermapore and by an extremely simplified phallosome 
(see also below). In addition, females of this genus have a dorsal pair of long curved 
hairs subapically on the third article of the hindtarsus and some long backwards- 
directed sternal setae near the midline in the basal half of the abdomen. All of these 
characters are probably autapomorphies of Afrotrogla. 

The genus Sensitibilla is characterized by the particularly tight basal fusion of 
the ovipositor valvulae with the subgenital plate, so that the posterior part of the latter 
becomes entirely membranous, being completely covered by the valvulae. Two other 
autapomorphies of this genus are the absence of M2 in the hindwing and the reduction 
of the number of thin-walled conical sensilla on P4 (2 in Sensitibilla vs 5-7 in other 
prionoglaridids, see Fig. 3). 

The high species diversity of Sensitibillini (i. e. six species, each from a 
different locality, all collected by non-specialists) in the relatively sparse material 
known from southern Africa indicates that these psocids are probably highly diver- 
sified in this region, especially in caves, but possibly also outside caves (see collecting 
data of Sensitibilla brandbergensis) . I am sure that further sampling in southern Africa, 
and especially in Namibia, will significantly increase our knowledge of this interesting 
group of Psocoptera. 

Functional morphology 

Genitalia: The possible biological significance (digging organ?) of the parti- 
cular ovipositor valvulae and of their partial fusion with the subgenital plate was 
briefly discussed by Lienhard (2000) for Sensitibilla. The fusion, unique in Psocoptera, 
is also present but less developed in Afrotrogla. The conspicuous porus (probably a 
glandular opening) on the subgenital plate of Afrotrogla maraisi is also a highly 
specialized structure. Its function is unknown, but it possibly forms a functional unit 
with the pair of lateral stylets present on the subgenital plate of this species (see Fig. 
6ef). The porus opens into a digitiform invagination of the subgenital plate and is sur- 
rounded by a series of stout setae. Some very fine hyaline hairs (length 3-10/im) could 
also be observed in the lumen of the porus. This pilosity has probably a protective 
function against intrusion of dust etc. into the porus. Structures similar to this porus and 
to the stylets have never been observed in Psocoptera. They may play a role in ovi- 
position by producing a protective secretion and distributing it on the surface of newly 
laid eggs. However, the absence of the porus and of the stylets in Afrotrogla oryx and 
in Sensitibilla shows that these structures are not essential parts of the functional unit 
"ovipositor valvulae - subgenital plate", present in all Sensitibillini. The unusual 
tuft(s) of long sternal setae basally on the abdomen of females of Afrotrogla and the 
pair of long curved subapical hairs on their hindtarsus (both absent in males) probably 
also play a role in reproductive biology. 

The membranous vesicle associated with the spermapore and its complex 
accessory structures, present in females of Afrotrogla, are also unique in Psocoptera. 
These structures are counterbalanced by an extreme simplification of phallosome 
sclerites in this genus. In A. oryx the phallosome sclerite and the accessory structures 
to spermapore are of very similar width (cf. scales to Figs 4g and 5c). It seems likely 



NEW AFRICAN SPELEKETORINAE 467 

that the otherwise relatively complex trogiomorphan phallosome structures are here 
functionally replaced by the complex female structures associated with the sperma- 
pore. According to Klier (1956) the pincer-like pair of parameres in male Trogio- 
morpha clasps the evaginated spermapore (opening of spermathecal duct) of females 
during the long-lasting copulation (often more than two hours) and is responsible for 
the close grip of male and female genitalia during sperm transmission. In Afrotrogla 
this role may have been transferred, at least partially, to the complex set of sclerites 
present near the opening of the spermathecal duct in females, in view of the reduction 
of the phallosome sclerites to a simple and rigid aedeagal arch. In this context it is 
interesting to note that in Sensitibilla the much more complex phallosome, bearing a 
pair of movable sclerites, is paralleled by a very simple spermapore sclerite. 

Trichobothria on legs: These trichobothria are very fine, but relatively rigid, 
long and erect filiform hairs inserted in special sockets. They have briefly been 
discussed by Lienhard (2000), who mentioned that they are probably vibroreceptors 
sensitive to air currents and low frequency air vibrations (sound) and may be inter- 
preted as adaptations to cave life. Some SEM micrographs are here presented for the 
first time (PL 1); they show the shape of the deep pit-like trichobothrial socket and the 
differences between the latter and the socket of a normal tactile hair. The particular 
trichobothrial pattern of Sensitibillini has been described in the diagnosis of the tribe. 
The presence of a trichobothrium on the second article of the hindtarsus (PI. 1ABC) in 
nymphs and adults of all Sensitibillini is of some general interest, because tarsal 
trichobothria seem to be unknown elsewhere in insects (Schuh, 1975 and Schuh in 
litt). Trichobothria are known from different body parts in insects, but on legs they 
have only been found in some Heteroptera families: In Miridae, Gerridae and Veliidae 
trichobothria regularly occur on coxae, trochanters and femora (Schuh, 1975), but 
never on tarsi (occasionally trichobothria-like setae are seen on tibiae in some semi- 
aquatic Heteroptera, see Moller Andersen, 1982). 

The trichobothrium on the second article of the hindtarsus of Sensitibillini is 
situated dorsally on the thickened central part of the article, in a zone of granular 
cuticle, and it is always accompanied by a campaniform sensillum (PI. 1BC). Such an 
auxiliary sensillum could not be observed near the tibial trichobothria. Similar 
campaniform sensilla, which are sensitive to slight deformations in the cuticle, have for 
instance been observed near cereal trichobothria of the cricket Gryllus bimaculatus 
(see Barth, 1986: fig. lie). No histological examination of these tarsal trichobothria 
was made, but the central thickening of the second article of the hindtarsus (PL 1A) 
indicates that the neurological equipment of this specialized sensillum is particularly 
space consuming. All tarsal articles without trichobothria, of all pairs of legs, are of 
normal cylindrical shape in Sensitibillini, as is the case in Speleketor and in Priono- 
glaridinae. 

ACKNOWLEDGEMENTS 

I am very grateful to Eugene Marais (NMN) for the loan of all unidentified 
psocids in his care, to Patrick Schmitz (MHNG) for transporting this material from 
Windhoek to Geneva, and to Kevin Johnson (Illinois Natural History Survey, 



468 C. LIENHARD 

Champaign) and Edward Mockford (Illinois State University, Normal) for placing at 
my disposal some specimens of Speleketor irwini and S. flocki, respectively. Many 
thanks also go to Edward Mockford and Peter Schwendinger (MHNG) for reading the 
manuscript and making valuable suggestions, and to Randall T. Schuh (American 
Museum of Natural History, New York) for providing general information on tricho- 
bothria in insects. Jean Wuest and Florence Marteau (MHNG) are acknowledged for 
producing the SEM micrographs and for assembling the plate. 

REFERENCES 

Azar, D. & Nel, A. 2004. Four new Psocoptera from Lebanese amber (Insecta: Psocomorpha 

[sic!]: Trogiomorpha). Annales de la Société entomologique de France (Nouvelle Série) 

40(2): 185-192. 
Barth, F. G. 1986. Zur Organisation sensorischer Systeme: die cuticularen Mechanoreceptoren 

der Arthropoden. Verhandlungen der Deutschen Zoologischen Gesellschaft 79: 69-90. 
Baz, A. & Ortuno, V. M. 2000. Archaeatropidae, a new family of Psocoptera from the 

Cretaceous amber of Alava, Northern Spain. Annals of the Entomological Society of 

America 93(3): 367-373. 
Baz, A. & Ortuno, V. M. 2001. New genera and species of empheriids (Psocoptera: 

Empheriidae) from the Cretaceous amber of Alava, northern Spain. Cretaceous Research 

22: 575-584. 
Gurney, A. B. 1943. A synopsis of the psocids of the tribe Psyllipsocini, including the 

description of an unusual new genus from Arizona (Corrodentia: Empheriidae: 

Empheriinae). Annals of the entomological Society of America 36: 195-220. 
Klier, E. 1956. Zur Konstruktionsmorphologie des männlichen Geschlechtsapparates der 

Psocopteren. Zoologische Jahrbücher (Abteilung Anatomie) 75: 207-286. 
Lienhard, C. 1988. Vorarbeiten zu einer Psocopteren-Fauna der Westpaläarktis. IV. Die Gattung 

Prionoglaris Enderlein (Psocoptera: Prionoglarididae). Mitteilungen der 

Schweizerischen Entomologischen Gesellschaft 61: 89-108. 
Lienhard, C. 1998. Psocoptères euro-méditerranéens. Faune de France 83: XX+517 pp. 
Lienhard, C. 2000. A new genus of Prionoglarididae from a Namibian cave (Insecta: 

Psocoptera). Revue suisse de Zoologie 107(4): 871-882. 
Lienhard, C. 2004. Siamoglaris zebrina gen. n., sp. n., the first representative of Priono- 
glarididae from the Oriental Region (Insecta: Psocoptera). Revue suisse de Zoologie 

111(4): 865-875. 
Lienhard, C. & Smithers, C. N. 2002. Psocoptera (Insecta): World Catalogue and Bibliography. 

Instrumenta Biodiversitatis 5: xli+745 pp. Muséum d'histoire naturelle, Genève. 
Mockford, E. L. 1984. Two new species of Speleketor from southern California with comments 

on the taxonomic position of the genus (Psocoptera: Prionoglaridae). Southwestern 

Naturalist 29(2): 169-179. 
Mockford, E. L. 1993. North American Psocoptera (Insecta). Flora and Fauna Handbook 10: 

XVIII+455 pp. Sandhill Crane Press, Gainesville, Florida. 
Moller Andersen, N. 1982. The semiaquatic bugs (Hemiptera, Gerromorpha): Phylogeny, 

adaptations, biogeography and classification. Entomonograph 3: 455 pp. 
Perrichot, V, Azar, D., Néraudeau, D. & Nel, A. 2003. New Psocoptera in the Early Creta- 
ceous amber of SW France and Lebanon (Insecta: Psocoptera: Trogiomorpha). 

Geological Magazine 140(6): 669-683. 
Schuh, R. T. 1975. The structure, distribution and taxonomic importance of trichobothria in the 

Miridae (Hemiptera). American Museum Novitates 2585: 1-26. 
Yoshizawa, K. 2005. Morphology of Psocomorpha (Psocodea: 'Psocoptera'). Insecta 

Matsumurana, New Series 62: 1-44. 



NEW AFRICAN SPELEKETORINAE 469 



Yoshizawa, K. & Johnson, K. P. 2006. Morphology of male genitalia in lice and their relatives 
and phylogenetic implications. Systematic Entomology 31: 350-361. 

Yoshizawa, K. & Saigusa, T. 2001. Phylogenetic analysis of paraneopteran orders (Insecta: 
Neoptera) based on forewing base structure, with comments on monophyly of 
Auchenorrhyncha (Hemiptera). Systematic Entomology 26: 1-13. 

Yoshizawa, K., Lienhard, C. & Johnson, K. P. 2006. Molecular systematics of the suborder 
Trogiomorpha (Insecta: Psocodea: 'Psocoptera'). Zoological Journal of the Linnean 
Society 146: 287-299. 



Revue suisse de Zoologie 114 (3): 471-505; septembre 2007 



A review of the catfish genus H ara, with the description of four 
new species (Silur if or mes: Erethistidae) 

Heok Hee NG 1 & Maurice KOTTELAT 2 

1 Fish Division, Museum of Zoology, University of Michigan, 1 109 Geddes Avenue, 
Ann Arbor, Michigan 48109-1079, USA. 

Current address: Raffles Museum of Biodiversity Research, Department of 
Biological Sciences, National University of Singapore, 6 Science Drive 2, #03-01, 
Singapore 117546. Email: heokheen@umich.edu 

2 Case postale 57, Cornol, CH-2952, Switzerland. Email: mkottelat@dplanet.ch 

A review of the catfish genus Hara, with the description of four new 
species (Siluriformes: Erethistidae). - Species of the erethistid catfish 
genus Hara are reviewed in this study. Eight species are recognized: Hara 
filamentosa, H. hara, H. horai, H. jerdoni, H. longissima, H. mesembrina, 
H. minuscula, and H. spinulus, of which the latter four are described as new 
herein. Erethistes maesotensis is a junior synonym of Hara filamentosa, and 
H. saharsai and H. serrata junior synonyms of H. hara. A neotype is 
designated for H. filamentosa Blyth, 1860 and a lectotype is designated for 
Hara horai Misra, 1976. 

Keywords: Sisoroidea - Bangladesh - India - Myanmar - Thailand. 

INTRODUCTION 

The Erethistidae are a small family of sisoroid catfishes inhabiting hillstreams 
in the subhimalayan region and western Indochina. The erethistid catfish genus Hara 
consists of small to medium sized catfishes with a brown cryptic coloration and heavily 
tuberculate skin. Species of Hara are diagnosed from other members of the family in 
having an anteroventral flange on the ventral surface of the complex centrum and the 
parapophysis of the fifth vertebra with a vertical lamina (de Pinna, 1996). The genus 
has been recorded from the Ganges-Brahmaputra river drainages eastwards to the 
Salween River drainage in western Thailand (Hora, 1950; Kottelat, 1989; 
Ukkatawewat & Vidthayanon, 1998). 

Recently, the authors obtained material from southern Thailand that consider- 
ably extends the southernmost extent of the distribution of Hara. Our attempts to iden- 
tify this material prompted us to investigate the identities of specimens collected from 
India, Myanmar and Thailand. The following study presents the results of this investi- 
gation, which reviews the species of Hara. We report here eight species of Hara, of 
which four are described as new. 

We received Thomson & Page's (2006) list of generic and species names in the 
families Erethistidae and Sisoridae after the present work was completed. A compila- 



Manuscript accepted 07.03.2007 



472 H. H. NG & M. KOTTELAT 

tion of such a nature would make sense as the completion of a revision of the various 
genera and species (but not before). Thomson & Page treat Hara as a junior synonym 
of Erethistes. In the discussion below, we mention the results of a phylogenetic ana- 
lysis based on both morphological and molecular data that make it premature to 
formally synonymise these genera. This is discussed in greater detail in HHN's 
doctoral dissertation (Ng, 2006). Thomson & Page's list of species and synonymies of 
Hara are apparently compilations, and do not affect our conclusions below (based on 
our studies of a considerable number of specimens). 

MATERIALS AND METHODS 

Measurements were made point to point with dial callipers and data recorded to 
0.1 mm. Counts and measurements were made on the left side of specimens whenever 
possible. Subunits of the head are presented as proportions of head length (HL). Head 
length itself and measurements of body parts are given as proportions of standard 
length (SL). Measurements and counts were made following Ng & Kottelat (1998) 
with the following addition: the length of the posterior process on coracoid is measured 
from the anterior edge of the coracoid where the process is situated to the posterior tip 
of the process. 

Fin rays were counted under a binocular dissecting microscope using trans- 
mitted light. Vertebrae were counted from radiographs obtained with a Hewlett 
Packard Faxitron 43855B cabinet x-ray machine. Vertebral counts and terminology 
follow those of Roberts (1994). Material for this study is deposited in the following 
collections: Natural History Museum, London (BMNH), California Academy of 
Sciences, San Francisco (CAS), collection of the second author, Cornol (CMK), 
Muséum d'histoire naturelle Genève (MHNG), Naturhistoriska Riksmuseet, 
Stockholm (NRM), Museum of Zoology, University of Michigan, Ann Arbor 
(UMMZ), and the Zoological Reference Collection of the Raffles Museum of 
Biodiversity Research, Singapore (ZRC). 

RESULTS 

Hara filamentosa Blyth, 1860 Fig. 1 

Hara filamentosa Blyth, 1860: 152 (type locality: Tenasserim) 
Erethistes conta (non Hamilton): Vinciguerra, 1890: 250. 

Erethistes maesotensis Kottelat, 1983: 71, Fig. 1 (type locality: Mae Nam Moei, 5 km west of 
Mae Sot. Thailand) 

Material examined: CMK 11961 (2), 48.9-53.1 mm SL; CMK 14689 (9), 42.1-44.4 
mm SL; Thailand: Tak province, Mae Nam Moei at Ban Wa Le. CMK 12175 (2), 44.6-47.0 mm 
SL; ZRC 42223 (1), 46.3 mm SL; Thailand: Tak province, stream at km 57 on road from Mae 
Sot to Wa Le (5 km before Wa Rei). ZRC 41269 (1), 44.3 mm SL; Thailand: Tak province, Mae 
Nam Moei at 16 17'47.1"N 98°42'44.9"E. MHNG 2150.5 (1 paratype of Erethistes maesoten- 
sis), 19.7 mm SL; Thailand: Tak province, Mae Nam Moei, 5 km W of Mae Sot. ZRC 50647 (1), 
43.1 mm SL; Thailand: Tak province, Mae Nam Moei at Mae Sot (border with Myanmar), 
16°41'13.2"N 98°30'54.0"E. CMK 14640 (11), 32.1-48.6 mm SL; MHNG 2680.076 (3), 40.2- 
48.1 mm SL; CMK 15102 (17), 20.6-30.1 mm SL; MHNG 2680.075 (5), 24.1-27.2 mm SL; 
Thailand: Tak province. Huai Mae Lamao, about 2 km N of road 105 on road from Ban Mae 
Lamao to Ban Mae Ramat. CMK 5529 (1), 48.3 mm SL; Thailand: Tak province. Huai Mae 
Lamao at Ban Mae Lamao. UMMZ 197583 (1), 47.4 mm SL; Thailand: Tak province, Mae 



A REVIEW OF THE CATFISH GENUS H ARA 



473 




Fig. 1 
Hara filamentosa, neotype, MHNG 2687.038, 43.7 mm SL; Myanmar: Megathat Chaung. Scale 
bar: 25 mm. 



474 H - H - NG & M - KOTTELAT 

Lamao, tributary of Salween River. CMK 14769 (1), 55.2 mm SL; Thailand: Mae Hong Son 
province, Huai Hai, about 5 km from Mae Hong Son to Pai road, on road turning off north at km 
44. CMK 16186 (5), 41.4-52.4 mm SL; CMK 17951 (2), 44.8-62.7 mm SL; Thailand: Mae Hong 
Song province, Salween River at Mae Sam Leap. CMK 16198 (1), 45.3 mm SL; Thailand: Mae 
Hong Son province, Nam Mae Kong Ka, 4-8 km upstrean of confluence with Salween River. 
CMK 16227 (1), 46.9 mm SL; Thailand: Mae Hong Son province, Salween River about 1 km 
upstream of Ban Mae Sam Leap. BMNH 1893.2.16.12-14 (3), 49.4-59.1 mm SL. - Myanmar: 
Meetan (=Mitan Chaung, a rivulet flowing S from the summit of Mulayet Taung, 16°H'N 
98°32'E, in the Salween basin). MHNG 2687.038, neotype, 43.7 mm SL; CMK 17786 (4), 46.8- 
59.7 mm SL; CMK 17951 (2), 45.1-61 .2 mm SL; Myanmar: Kayin state, Ataran River drainage, 
stream "Chon Son" between Kyondaw and Phadaw, about 20 km NW of Payathouzu (at border 
with Thailand), 15°25'N 98°15'E. UMMZ 243692 (6), 46.4-50.0 mm SL; Myanmar: from 
aquarium trade. UMMZ 245962 (1), 53.7 mm SL; Myanmar: Bago division, Pyu township, Pyu 
stream (tributary of Sittang River) ca. 229 km from Yangon, 18°29'N 96°26'E. UMMZ 245964 
(9), 56.7-67.1 mm SL; Myanmar: Kayin state, hillstreams in Ataran River drainage in the vicin- 
ity of Payathonzu. 

Diagnosis: Hara filamentosa can be distinguished from congeners (except 
H. mesembrina and H. longissima) in sometimes having (vs. always lacking) a fila- 
mentous extension to the first principal ray of the upper caudal-fin lobe; it is also dis- 
tinguished from all congeners except H. hara and H. longissima in having a short pos- 
terior process on coracoid (reaching to midway of distance between bases of pectoral 
spine and first pelvic-fin ray vs. two thirds or beyond; length 19.9-22.8% SL vs. 23.5- 
38.3). It differs from H. longissima in having a shorter (length 14.9-17.8% SL vs. 18.8- 
21.3) and deeper (depth 6.1-7.2% SL vs. 5.0-6.1) caudal peduncle (depth 2.1-2.9 times 
in its length for H. filamentosa vs. 3.1-3.9 times in H. longissima) and deeper body 
(depth 13.9-19.2% SL vs. 12.5-14.5). Hara filamentosa can be further distinguished 
from H. hara in having a longer adipose-fin base (15.7-21.7% SL vs. 11.8-15.6), 
deeper caudal peduncle (depth 6.1-7.2% SL vs. 5.0-5.8) and lacking (vs. presence) of 
serrations on the anterior edge of the dorsal spine, from H. horai in having a deeper 
caudal peduncle (depth 6.1-7.2% SL vs. 5.2-6.2) and a shorter adipose-fin base (length 
15.7-21.7% SLvs. 10.0-13.7) and head (length 26.5-29.1% SLvs. 30.3-34.2), from//. 
jerdoni in having a shorter pectoral spine (length 23.3-29.9% SL vs. 42.2-51.7), more 
soft pectoral-fin rays (total number of soft fin rays modally 7 vs. 5), more vertebrae 
(31-34 vs. 27-29), the supraoccipital spine not reaching (vs. reaching) the anterior 
nuchal plate and a deeply forked (vs. emarginate) caudal fin. It differs from H. mesem- 
brina in having a shorter caudal peduncle (length 14.9-17.8% SL vs. 17.2-18.9), from 
H. minuscula in having more soft pectoral-fin rays (total number of soft fin rays 
modally 7 vs. 5) and the supraoccipital spine not reaching (vs. reaching) the anterior 
nuchal plate, and from H. spinulus in having a longer dorsal spine (length 20.4-27.6% 
SL vs. 14.7-19.8), adipose-fin base (length 15.7-21.7% SL vs. 12.9-16.1) and caudal 
peduncle (length 14.9-17.8 % SLvs. 11.8-15.0). 

Description: Morphometric data in Table 1. Head and body moderately 
compressed. Dorsal profile curved from tip of snout to level of nares, then evenly 
sloping to origin of dorsal fin; evenly sloping ventrally to origin of adipose fin and 
gently concave from origin of adipose fin to end of caudal peduncle. Ventral profile flat 
to pelvic-fin base; sloping gently dorsally to end of anal-fin base and gently concave 
from end of anal-fin base to end of caudal peduncle. Anus and urogenital openings 



A REVIEW OF THE CATFISH GENUS H ARA 



475 



Table 1. Morphometric data for Hara filamentosa (n=25: 43.7-67.1 mm SL). 



Range 



Mean±SD 



%SL 

Predorsal length 

Preanal length 

Prepelvic length 

Prepectoral length 

Length of dorsal fin base 

Dorsal spine length 

Anal fin length 

Pelvic fin length 

Pectoral fin length 

Pectoral spine length 

Caudal fin length 

Length of adipose fin base 

Dorsal to adipose distance 

Post-adipose distance 

Caudal peduncle length 

Caudal peduncle depth 

Body depth at anus 

Head length 

Head width 

Head depth 

Length of posterior process on coracoid 

%HL 

Snout length 
Interorbital distance 
Eye diameter 
Nasal barbel length 
Maxillary barbel length 
Inner mandibular barbel length 
Outer mandibular barbel length 



42.1-46.7 


44.3±1.44 


69.1-72.9 


70.4±1.14 


49.0-52.3 


51.0+1.04 


21.3-27.8 


24.0±1.66 


17.2-20.7 


1 8 .9± 1.16 


20.4-27.6 


23.6±2.48 


13.0-17.2 


14.8+1.10 


16.0-19.6 


17.2+1.20 


25.6-32.4 


28.8±2.16 


23.3-29.9 


27. 0± 1.99 


20.9-30.3 


26.7±2.80 


15.7-21.7 


18.7+1.99 


5.1-11.0 


7.2±1.96 


14.1-18.3 


16.5±1.50 


14.9-17.8 


16.7±0.88 


6.1-7.2 


6.8±0.37 


13.9-19.2 


16.9+1.65 


26.5-29.1 


28.0±0.91 


21.0-27.0 


24 .7± 1.75 


19.1-24.3 


21.5+1.54 


19.9-22.8 


21.3+1.04 


50.0-60.7 


54.9+3.60 


32.2-41.4 


37.3±3.04 


12.5-15.5 


13.7+1.21 


12.6-22.1 


17.0±3.59 


57.5-94.4 


85.0±10.66 


31.0-45.4 


38.1±4.47 


37.4-61.3 


52.5±8.00 



located at vertical through posterior three quarters of adpressed pelvic fin. Skin tuber- 
culate, tubercles distributed evenly throughout head and body. Lateral line complete 
and midlateral. Vertebrae 14+17=31 (1), 14+18=32 (2), 15+17=32 (2), 14+19=33 (1), 
15+18=33 (5), 16+17=33 (4), 15+19=34 (2) or 16+18=34 (7). 

Snout narrow; profile dorsally rounded when viewed laterally and acutely 
triangular when viewed from above. Snout with pronounced steepening of curvature 
anteriorly at level of posterior nares when viewed laterally. Anterior and posterior nares 
large and separated by flap of skin comprising base of nasal barbel. Gill openings 
narrow, extending from immediately ventral to posttemporal to immediately lateral to 
point immediately ventral to base of pectoral spine. Bony elements of dorsal surface of 
head covered with thin, tuberculate skin and very prominent. Supraoccipital spine not 
reaching anterior nuchal plate. Weberian lamina well developed, approximately same 
length as supraoccipital spine and extending parallel to either side of spine. Eye ovoid, 
horizontal axis longest; located entirely in dorsal half of head, with faint dorsal ridge 
on frontal bone dorsal to eye. Orbit with free margin. 



476 H. H. NG & M. KOTTELAT 

Barbels in four pairs. Maxillary barbel long and slender, extending to middle of 
pectoral-spine base. Nasal barbel short, subtended by flap of skin at base and extend- 
ing to midway between posterior edge of posterior nares and anterior margin of orbit. 
Inner mandibular-barbel origin close to midline, extending to middle of distance 
between its base and that of pectoral spine. Outer mandibular barbel originating 
posterolateral of inner mandibular barbel, extending to about three quarters of distance 
between its base and that of pectoral spine. 

Mouth inferior, premaxillary tooth band not exposed when mouth is closed. 
Upper lip with indistinct plicae. Oral teeth small and villiform, in irregular rows on all 
tooth-bearing surfaces. Premaxillary teeth in single crescentic band across midline. 
Palate edentulous. Dentary teeth in two narrow crescentic bands separated at midline. 

Dorsal fin located above anterior third of body, with 1,5 ,i (25) rays; dorso- 
posteri or fin margin straight. Dorsal spine moderately long and gently curved; with 
smooth anterior margin and posterior margin with 4-7 serrae. Adipose fin with anterior 
margin straight and posterior margin angular. Caudal fin strongly forked, with i,7,7,i 
(1), i,7,8,i (23) or i,8,7,i (1) principal rays; upper lobe slightly longer than lower. First 
principal ray of upper caudal-fin lobe produced into a filament extending posteriorly 
for length of upper caudal lobe proper. Procurrent rays symmetrical and extending only 
slightly anterior to fin base. Anal-fin base approximately in line with adipose-fin base. 
Anal fin with convex margin anteriorly and straight margin posteriorly, with iii,6,i (3), 
iv,6 (2) iv,6,i (8), iii,7 (2), iii,7,i (5), v,6 (1), iv,7 (1) or iv,7,i (3) rays. Pelvic-fin origin 
anterior to vertical through posterior end of dorsal-fin base. Pelvic fin with pointed tip 
and straight posterior margin, with i,5 (25) rays; tip of adpressed fin just reaching anal- 
fin origin. Pectoral fin with 1,5 ,i (3) or 1,6 ,i (22) rays; fin margin gently convex 
posteriorly; anterior spine margin with 14-23 small distally directed serrations, 
posterior margin with 9-13 large medially directed serrations. Coracoid with well 
developed posterior processes, extending to midway between base of posteriormost 
pectoral-fin ray and pelvic-fin origin. 

Colour: Head and body dark chestnut brown to light chocolate brown, with a 
mottling caused by numerous smaller, irregular darker brown patches. Belly and ven- 
tral surfaces of head light brown. Two thin, irregular light brown bars on body: first 
chevron-shaped and located between dorsal- and adipose-fin bases, second thicker than 
first and located on caudal peduncle. Dorsal fin dark mottled brown, except for thin 
hyaline distal margin. Pectoral fin dark brown, with thin hyaline distal margin and 
either a small ovoid hyaline spot or a crescentic median hyaline band on middle third 
of fin. Pelvic and anal fins hyaline, with dark brown bases and subdistal stripes. 
Adipose fin brown as in body, with lighter coloured distal margin. Caudal fin hyaline, 
with irregular W-shaped band subdistally and with small dark brown flecks randomly 
distributed throughout fin. Two small light brown patches present at bases of caudal- 
fin lobes in some individuals. Maxillary and mandibular barbels light brown, with dark 
brown annuii. 

Distribution. Hara filamentosa is known from the Ataran, Salween and Sittang 
river drainages (Fig. 2). In the Salween, it has been collected in the Salween proper and 
the Mae Nam Moei. in Tak and Mae Hong Son Provinces of Thailand. 



A REVIEW OF THE CATFISH GENUS HARA 



All 




•O Hara filamentosa 
+* Hara hara 
** Hara horai 
♦ Hara jerdoni 
D Hara longissima 
V Hara mesembrina 
AA Hara minuscula 
X» Hara spinulus 



5 a 

80 



1000 



500 

kilometers 

Fig. 2 
Map showing collecting localities of Hara examined. Outlined symbols are type localities. 



Hara hara (Hamilton 1 822) 



Fig. 3 



Pimelodus hara Hamilton, 1822: 190 (type locality: Nathpur, India) 

Pimelodus asperus M'Clelland, 1844: 404, pi. 24 fig. 2 (type locality: Hooghly River S of 
Ranaghat) 



478 



H. H. NG & M. KOTTELAT 




A REVIEW OF THE CATFISH GENUS HARA 479 



Hara saharsai Datta Munshi & Srivastava, 1988: 265, fig. 5 (type locality: Kosi Belt, northern 

Bihar, India) 
Hara serratus Vishwanath & Kosygin, 2000: 143, figs 2-5 (type locality: Jiri River at Jiribam, 

Manipur, India) 

Material examined: UMMZ 244697 (neotype), 47.2 mm SL; India: W Bengal, 
Hooghly River S of Ranaghat. UMMZ 208748 (44), 33.0-50.0 mm SL; Bangladesh: Gowain 
River at Gowainghat. 

Diagnosis: Hara hara can be distinguished from congeners in possessing (vs. 
lacking) serrations on the anterior edge of the pectoral spine. It further differs from 
H. filamentosa, H. mesembrina and H. longissima in always lacking (vs. sometimes 
having) a filamentous extension to the first principal ray of the upper caudal-fin lobe; 
it is also distinguished from all congeners except H. filamentosa and H. longissima in 
having a short posterior process on coracoid (reaching to midway of distance between 
bases of pectoral spine and first pelvic-fin ray vs. two thirds or beyond; length 19.9- 
24.8% SL vs. 23.5-38.3). Hara hara further differs from H . filamentosa in having a 
shorter adipose-fin base (11.8-15.6% SL vs. 15.7-21.7) and slenderer caudal peduncle 
(depth 5.0-5.8% SL vs. 6.1-7.2), from H. horai in having a longer caudal fin (25.8- 
28.0% SL vs. 20.5-23.5), slenderer body (11.5-14.3% SL vs. 15.1-17.9) and a shorter 
head (length 27.2-30.9% SL vs. 30.3-34.2), from H.jerdoni in having a shorter pec- 
toral spine (length 26.9-33.3% SL vs. 42.2-51.7), more soft pectoral-fin rays (total 
number of soft fin rays modally 7 vs. 5), more vertebrae (3 1-34 vs. 27-29), the supraoc- 
cipital spine not reaching (vs. reaching) the anterior nuchal plate, and a deeply forked 
(vs. emarginate) caudal fin. It further differs from H. mesembrina in having a slenderer 
caudal peduncle (depth 5.0-5.8% SL vs. 6.0-7.4), from H. minuscula in having a 
narrower head (width 22.4-24.3% SL vs. 24.5-27.2), more soft pectoral-fin rays (total 
number of soft fin rays modally 7 vs. 5) and the supraoccipital spine not reaching (vs. 
reaching) the anterior nuchal plate, and from H. spinulus in having a longer dorsal 
spine (22.7-27.4% SL vs. 14.7-19.8), narrower head (width 22.4-24.3% SL vs. 25.9- 
30.0) and slenderer caudal peduncle (depth 5.0-5.8% SL vs. 6.7-8.5). 

Description: Morphometric data in Table 2. Exactly as in Hara filamentosa, 
except for the following. Vertebrae 14+18=32 (7), 15+17=32 (4), 14+19=33 (1), 
15+18=33 (9) or 15+19=34 (3). Dorsal fin with 1,5 ,i (24) rays. Dorsal spine moderately 
long and gently curved; with serrated anterior margin and posterior margin with 8-13 
serrae. Caudal fin with i,7,6,i (24) principal rays. First principal ray of upper caudal-fin 
lobe not produced into a filament. Anal fin with iv,6,i (24) rays. Pelvic fin with i,5 (24) 
rays. Pectoral fin with 1,6 ,i (24) rays; anterior spine margin with 17-24 small distally 
directed serrations, posterior margin with 12-17 large medially directed serrations. 
Coracoid with well developed posterior processes, extending to midway between base 
of posteriormost pectoral-fin ray and pelvic-fin origin. 

Colour: Head and body chestnut brown or light brown, with a mottling caused 
by numerous smaller, irregular darker brown patches. Belly and ventral surfaces of 
head light brown. Two thin, irregular light brown bars on body: first chevron-shaped 
and located between dorsal- and adipose-fin bases, second thicker than first and located 

Fig. 3 
Hara hara, neotype, UMMZ 244697, 47.2 mm SL; India: Hooghly River at Ranaghat. Scale bar: 
25 mm. 



480 



H. H. NG & M. KOTTELAT 



Table 2. Morphometric data for Hara hara (n=24: 33.0-50.0 mm SL). 

Range 



Mean±SD 



%SL 

Predorsal length 

Preanal length 

Prepelvic length 

Prepectoral length 

Length of dorsal fin base 

Dorsal spine length 

Anal fin length 

Pelvic fin length 

Pectoral fin length 

Pectoral spine length 

Caudal fin length 

Length of adipose fin base 

Dorsal to adipose distance 

Post-adipose distance 

Caudal peduncle length 

Caudal peduncle depth 

Body depth at anus 

Head length 

Head width 

Head depth 

Length of posterior process on coracoid 

%HL 

Snout length 
Interorbital distance 
Eye diameter 
Nasal barbel length 
Maxillary barbel length 
Inner mandibular barbel length 
Outer mandibular barbel length 



44.0-45.5 


44.6±0.64 


66.9-70.9 


68.7+1.54 


47.6-53.6 


51.1+2.28 


21.2-27.3 


24.4±2.22 


16.7-21.0 


18.6±1.60 


22.7-27.4 


24.5±1.88 


14.6-16.8 


15.6±0.82 


15.8-18.2 


16.8±0.91 


30.0-36.2 


32.5+2.40 


26.9-33.3 


30.0±2.32 


25.8-28.0 


27.0±1.01 


11.8-15.6 


14.0±1.46 


10.8-12.7 


12.0±0.71 


15.7-18.6 


17.2+1.27 


14.6-19.4 


17.7+1.85 


5.0-5.8 


5.4±0.34 


11.5-14.3 


12.8+1.28 


27.2-30.9 


29.6±1.41 


22.4-24.3 


23.5±0.91 


18.6-20.1 


19.3+0.61 


19.9-24.8 


22.0±1.79 


50.0-54.7 


51.8+1.87 


31.5-38.0 


35.4±2.53 


9.4-10.9 


10.1±0.65 


14.8-21.9 


18.0±3.07 


80.8-108.2 


93.4±9.78 


34.2-49.6 


43.6±6.13 


52.1-67.2 


60.8±6.13 



on caudal peduncle. Dorsal fin dark mottled brown, except for thin hyaline distal 
margin. Pectoral fin dark brown, with thin hyaline distal margin and either a small 
ovoid hyaline spot or a crescentic median hyaline band on middle third of fin. Pelvic 
and anal fins hyaline, with dark brown bases and subdistal stripes. Adipose fin brown 
as in body, with lighter coloured distal margin. Caudal fin hyaline, with irregular W- 
shaped band subdistally and with small dark brown flecks randomly distributed 
throughout fin. Two small light brown patches present at bases of caudal-fin lobes in 
some individuals. Maxillary and mandibular barbels light brown, with dark brown 
annuii. 

Distribution: Hara hara is known from the Brahmaputra and Ganges river 
drainages (Fig. 2). 



Hara horai Misra, 1976 



Fig. 4 



Hara horai Misra, 1976: 245, PL 9 figs 1-3 (type locality: Terai and Duars, northern Bengal, 
India) 

Material examined: ZSI FF955 (lectotype), 58.8 mm SL; India: West Bengal, Terai and 
Duars. UMMZ 245633 (2), 56.2-73.1 mm SL; ZRC 50578 (2), 41 .8-62.7 mm SL; India: Assam, 



A REVIEW OF THE CATFISH GENUS HARA 481 

Tinsukia, Dibru River at Guijan. ZRC 50576 (3), 44.8-50.4 mm SL; India: Assam, Guijan. ZRC 
38914 (2), 83.4-85.4 mm SL; ZRC 41364 (1), 77.8 mm SL; India: Assam, from aquarium trade. 

Diagnosis: Hara horai can be distinguished from H. filamentosa in always 
lacking (vs. sometimes having) a filamentous extension to the first principal ray of the 
upper caudal-fin lobe and having a slenderer caudal peduncle (depth 5.2-6.2% SL vs. 
6.1-7.2), shorter adipose-fin base (length 10.0-13.7% SL vs. 15.7-21.7), longer head 
(length 30.3-34.2% SL vs. 26.5-29.1) and posterior process on coracoid (reaching to 
two-thirds distance between bases of pectoral spine and first pelvic-fin ray vs. midway; 
length 23.5-26.2% SL vs. 19.9-22.8), from H. hara in having a shorter caudal fin (20.5- 
23.5% SL vs. 25.8-28.0), deeper body (15.1-17.9% SL vs. 11.5-14.3), and a longer 
head (length 30.3-34.2% SL vs. 27.2-30.9), and from H.jerdoni in having a shorter 
pectoral spine (length 28.0-33.4% SL vs. 42.2-51.7), more soft pectoral-fin rays (total 
number of soft fin rays modally 7 vs. 5), more vertebrae (31-34 vs. 27-29), the supra- 
occipital spine not reaching (vs. reaching) the anterior nuchal plate and a deeply forked 
(vs. emarginate) caudal fin. It differs from H. longissima in always lacking (vs. some- 
times having) a filamentous extension to the first principal ray of the upper caudal-fin 
lobe and having a shorter adipose-fin base (length 10.0-13.7% SL vs. 13.3-16.5) and 
caudal peduncle (length 15.5-17.9% SL vs. 18.8-21.3), a deeper body (depth 15.1- 
17.9% SL vs. 12.5-14.5), a longer head (length 30.3-34.2% SL vs. 26.0-29.1) and a 
smaller eye (diameter 8.0-9.8% HL vs. 11.3-13.8), from H. mesembrina in always 
lacking (vs. sometimes having) a filamentous extension to the first principal ray of the 
upper caudal-fin lobe and having a shorter adipose-fin base (length 10.0-13.7% SL vs. 
13.5-17.0), slenderer caudal peduncle (depth 5.2-6.2% SL vs. 6.0-7.4), longer head 
(length 30.3-34.2% SL vs. 22.1-27.3) and smaller eye (diameter 8.0-9.8% HL vs. 13.1- 
17.9), from H. minuscula in having a longer head (30.3-34.2% SL vs. 24.8-28.4), 
shorter posterior process on coracoid (reaching to two thirds of distance between bases 
of pectoral spine and first pelvic-fin ray vs. three quarters; length 23.5-26.2% SL vs. 
29.5-32.9), more soft pectoral-fin rays (total number of soft fin rays modally 7 vs. 5) 
and the supraoccipital spine not reaching (vs. reaching) the anterior nuchal plate, and 
from//, spinulus in having a longer dorsal spine (length 21.0-28.4% SLvs. 14.7-19.8), 
shorter adipose-fin base (length 10.0-13.7% SL vs. 12.9-16.1), longer (length 15.5- 
17.9% SL vs. 11.8-15.0) and slenderer (depth 5.2-6.2% SL vs. 6.7-8.5) caudal 
peduncle, longer head (length 30.3-34.2% SL vs. 26.9-28.9) and smaller eye (diameter 
8.0-9.8% SLvs. 11.2-14.4). 

Description: Morphometric data in Table 3. Exactly as in Hara filamentosa, 
except for the following. Vertebrae 14+17=31 (1), 15+16=31 (1), 14+18=32 (6), 
15+17=32 (1) or 14+19=33 (1). Dorsal fin with 1,5 ,i (10) rays. Dorsal spine moderate- 
ly long and gently curved; with smooth anterior margin and posterior margin with 8-1 1 
serrae. Caudal fin with i,7,6,i (10) principal rays. First principal ray of upper caudal-fin 
lobe not produced into a filament. Anal fin with iv,6,i (10) rays. Pelvic fin with i,5 (10) 
rays. Pectoral fin with 1,6 ,i (10) rays; anterior spine margin with 19-25 small distally 
directed serrations, posterior margin with 9-17 large medially directed serrations. 
Coracoid with well developed posterior processes, extending to two-thirds distance 
between base of posteri ormost pectoral-fin ray and pelvic-fin origin. 



482 



H. H. NG & M. KOTTELAT 




A REVIEW OF THE CATFISH GENUS HARA 



483 



Table 3. Morphometric data for Hara horai (n=10: 41.8-85.4 mm SL). 



Range 



Mean±SD 



%SL 

Predorsal length 

Preanal length 

Prepelvic length 

Prepectoral length 

Length of dorsal fin base 

Dorsal spine length 

Anal fin length 

Pelvic fin length 

Pectoral fin length 

Pectoral spine length 

Caudal fin length 

Length of adipose fin base 

Dorsal to adipose distance 

Post-adipose distance 

Caudal peduncle length 

Caudal peduncle depth 

Body depth at anus 

Head length 

Head width 

Head depth 

Length of posterior process on coracoid 

%HL 

Snout length 

Interorbital distance 

Eye diameter 

Nasal barbel length 

Maxillary barbel length 

Inner mandibular barbel length 

Outer mandibular barbel length 



42.1-49.2 


46.4±2.35 


68.6-70.9 


69.9±0.86 


50.7-53.2 


52.0+0.88 


23.0-29.4 


25.8±2.22 


19.1-23.3 


21.0±1.82 


21.0-28.4 


24.0±2.60 


11.2-17.7 


14.6±2.25 


14.8-18.7 


17.2+1.73 


30.9-36.5 


32.6±3.01 


28.0-33.4 


30.4±2.07 


20.5-23.5 • 


23 .5± 1.72 


10.0-13.7 


12.2±1.40 


7.7-12.5 


10.8+2.00 


14.0-16.7 


15.6±1.08 


15.5-17.9 


16.6±0.89 


5.2-6.2 


5.7+0.39 


15.1-17.9 


167+1.01 


30.3-34.2 


31.6+1.61 


23.1-29.6 


27.3±2.27 


19.6-21.9 


21.1+0.83 


23.5-26.2 


24 .7± 1.63 


48.0-53.3 


50.0±2.06 


30.8-36.5 


33.7±2.10 


8.0-9.8 


8.8+0.82 


15.8-26.0 


21.2±4.90 


71.5-84.6 


76.2±5.08 


32.4-40.0 


36.9±3.37 


40.4-52.6 


45.9±4.48 



Colour: Head and body dark chestnut dark chocolate brown, without mottling. 
Belly and ventral surfaces of head light brown. Two thin, irregular light brown bars on 
body: first chevron-shaped and located between dorsal- and adipose-fin bases, second 
thicker than first and located on caudal peduncle. Dorsal fin dark brown, except for thin 
hyaline distal margin. Pectoral fin dark brown, with thin hyaline distal margin and 
either a small ovoid hyaline spot or a crescentic median hyaline band on middle third 
of fin. Pelvic and anal fins hyaline, with dark brown bases and subdistal stripes. 
Adipose fin brown as in body, with lighter coloured distal margin. Caudal fin hyaline, 
with irregular W-shaped band subdistally and with small dark brown flecks randomly 
distributed throughout fin. Maxillary and mandibular barbels light brown, with dark 
brown annuii. 



Distribution: Hara horai is known from the Brahmaputra River drainage 



(Fig. 2). 



Fig. 4 
Hara horai, ZRC 38914, 83.4 mm SL; India: Assam. Scale bar 50 mm. 



484 H. H. NG & M. KOTTELAT 

Harajerdoni Day, 1870 Fig. 5 

Hara jerdoni Day, 1870: 39, PI. 4 (type locality: Sylhet district, Bangladesh) 

Material examined: UMMZ 208401 (12), 15.6-20.6 mm SL; Bangladesh: Dakatia 
River at Chandpur. UMMZ 244581 (1), 19.2 mm SL; India: West Bengal, Dolong River (a trib- 
utary of the Mansai River) in the vicinity of Shildanga, 26?23'13"N 89°14'31"E. 

Diagnosis: Hara jerdoni can be distinguished from all congeners in having a 
longer pectoral spine (length 42.2-51.7% SLvs. 23 .3-34.7), fewer vertebrae (27-29 vs. 
30-35) an emarginate (vs. deeply forked) caudal fin and a longer posterior process on 
coracoid (reaching to four fifths of distance between bases of pectoral spine and first 
pelvic-fin ray vs. three quarters or less; length 34.4-38.3% SL vs. 19.9-32.9). It further 
differs from all congeners except H. minuscula in having fewer soft pectoral-fin rays 
(total number of soft fin rays modally 5 vs. 6-7) and the supraoccipital spine reaching 
(vs. not reaching) the anterior nuchal plate. Hara jerdoni is also the smallest species of 
Hara, not reaching ca. 25 mm SL in maximum size (vs. ca. 30 mm SL in H. minuscu- 
la and 60 mm SL in other congeners; Table 4). 

Description: Morphometric data in Table 5 . Exactly as in Hara filamentosa, ex- 
cept for the following. Vertebrae 14+13=27 (1), 14+14=28 (3), 15+13=28 (6) or 
15+14=29 (2). Supraoccipital spine reaching anterior nuchal plate. Dorsal fin with 1,5 ,i 
(12) rays. Dorsal spine moderately long and gently curved; with smooth anterior mar- 
gin and posterior margin with 4-5 serrae. Caudal fin emarginate, with i,5,5,i (12) prin- 
cipal rays; without filamentous extension to first principal ray of upper lobe. Anal fin 
with iii,5,i (11) or iii,6,i (1) rays. Pelvic fin with i,5 (12) rays. Pectoral fin with I,4,i (12) 
rays; anterior spine margin with 18-25 small distally directed serrations, posterior mar- 
gin with 10-12 large medially directed serrations. Coracoid with well developed pos- 
terior process, extending to four-fifths distance between base of posteriormost pectoral- 
fin ray and pelvic-fin origin. 

Colour: Head and body dark chestnut brown to light chocolate brown, with a 
mottling caused by numerous smaller, irregular darker brown patches. Belly and ven- 
tral surfaces of head light brown. Two thin, irregular light brown bars on body: first 
chevron-shaped and located between dorsal- and adipose-fin bases, second thicker than 
first and located on caudal peduncle. Dorsal fin dark mottled brown, except for thin 
hyaline distal margin. Pectoral fin dark brown, with thin hyaline distal margin and ei- 
ther a small ovoid hyaline spot or a crescentic median hyaline band on middle third of 
fin. Pelvic and anal fins hyaline, with dark brown bases and subdistal stripes. Adipose 
fin brown as in body, with lighter coloured distal margin. Caudal fin hyaline, with ir- 
regular W-shaped band subdistally and with small dark brown flecks randomly distrib- 
uted throughout fin. Two small light brown patches present at bases of caudal-fin lobes 
in some individuals. Maxillary and mandibular barbels light brown, with dark brown 
annuii. 

Distribution: Hara jerdoni is known from the Brahmaputra River drainage 
(Fig. 2), and has also been recorded from the Ganges River drainage (Husain & Tilak, 
1978). 

Fig. 5 
Harajerdoni, UMMZ 208401, 19.2 mm SL; Bangladesh: Dakatia River. Scale bar: 10 mm. 



A REVIEW OF THE CATFISH GENUS H ARA 



485 




486 



H. H. NG & M. KOTTELAT 



Table 4. Maximum known sizes of Hara species 



Species 



Maximum size (SL) 



Hara filamentosa 
Hara hara 
Hara horai 
Hara jerdoni 
Hara longissima 
Hara mesembrina 
Hara minuscula 
Hara spinulus 



67 mm 
50 mm 
85 mm 
21 mm 
55 mm 
55 mm 
30 mm 
50 mm 



Table 5. Morphometric data for Hara jerdoni (n=ll: 15.6-20.6 mm SL). 

Range 



Mean±SD 



%SL 

Predorsal length 

Preanal length 

Prepelvic length 

Prepectoral length 

Length of dorsal fin base 

Dorsal spine length 

Anal fin length 

Pelvic fin length 

Pectoral fin length 

Pectoral spine length 

Caudal fin length 

Length of adipose fin base 

Dorsal to adipose distance 

Post-adipose distance 

Caudal peduncle length 

Caudal peduncle depth 

Body depth at anus 

Head length 

Head width 

Head depth 

Length of posterior process on coracoid 

%HL 

Snout length 
Interorbital distance 
Eye diameter 
Nasal barbel length 
Maxillary barbel length 
Inner mandibular barbel length 
Outer mandibular barbel length 



38.5-43.3 
67.3-70.3 
53.1-54.3 
18.0-19.7 
19.1-19.7 
19.1-21.4 
14.4-16.3 
14.4-15.5 
46.6-46.9 
42.2-51.7 
24.5-27.9 
9.9-12.9 
10.8-12.0 
15.5-18.2 
14.9-15.4 
4.7-5.3 
15.5-16.1 
27.6-29.4 
25.5-31.3 
22.6-23.7 
34.4-38.3 

41.6-47.2 
37.7-41.6 
12.3-17.0 
16.7-28.1 
52.8-105.3 
37.7-51.7 
47.2-84.2 



41.5±2.62 
68.9+1.51 
53.8±0.64 
18.6±0.93 
19.4+0.31 
19.9+1.30 
15.4±0.96 
14.8±0.59 
46.8+0.21 
48.0±4.69 
26.6+1.81 
11.0+1.68 
11.4±0.60 
16.5+1.46 
15.1+0.25 
5.1±0.32 
15.8+0.31 
28.6±0.93 
28.2±2.92 
23.2±0.57 
36.4±1.40 

44.2+2.81 
39.9+2.00 
14.2+2.48 
21.2±6.05 
11.1 ±26 35 
46.2±7.45 
66.0+18.51 



Hara longissima sp. no v. 



Fig. 6 



Holotype: UMMZ 245902, 51.2 mm SL; Myanmar: Kachin state, Myitkyina district, 
hillstreams approximately 2 hours by foot on road from Mogaung to Kamaing; Tin Win & Kyaw 
Zin, 1 August 2004. 

Paratypes: UMMZ 245945 (10), 49.0-55.3 mm SL; data as for holotype. 



A REVIEW OF THE CATFISH GENUS H ARA 



487 




Fig. 6 
Hara longissima, holotype, UMMZ 245902, 51.2 mm SL; Myanmar: hillstreams approximately 
2 hours by foot on road from Mogaung to Kamaing. Scale bar: 25 mm. 



488 H. H. NG & M. KOTTELAT 

Diagnosis: Hara longissima can be distinguished from all congeners in having 
a longer caudal peduncle (18.8-21 .3% SL vs. 11 .7-19.4) and from congeners except H. 
filamentosa and H. mesembrina in having (vs. lacking) a filamentous extension of the 
first principal ray of the upper caudal-fin lobe. It further differs from H . filamentosa in 
having a slenderer (depth 5.0-6.1% SL vs. 6.1-7.2) caudal peduncle and body (depth 
12.5-14.5% SL vs. 13.9-19.2), from H. hara in lacking (vs. having) serrations on the 
anterior edge of the dorsal spine, and from H. horai in having a longer adipose-fin base 
(length 13.3-16.5% SLvs. 10.0-13.7), a slenderer body (depth 12.5-14.5% SL vs. 15.1- 
17.9), a shorter head (length 26.0-29.1% SL vs. 30.3-34.2) and a larger eye (diameter 
11.3-13.8% HL vs. 8.0-9.8). Hara longissima can be further distinguished from both 
H. jerdoni and H. minuscula in having a shorter posterior process on coracoid 
(reaching to midway of distance between bases of pectoral spine and first pelvic-fin ray 
vs. at least three quarters; its length 20.7-23.9% SL vs. 29.5-38.3), more soft pectoral- 
fin rays (total number of soft fin rays modally 7 vs. 5) and vertebrae (33-35 vs. 27-32), 
and the supraoccipital spine not reaching (vs. reaching) the anterior nuchal plate, and 
from H. jerdoni in having a shorter pectoral spine (length 25.9-31.8% SL vs. 42.2- 
51.7). It further differs from H. mesembrina in having a smaller eye (diameter 11.3- 
13.8% HL vs. 13.1-17.9) and shorter posterior process on coracoid (reaching to 
midway of distance between bases of pectoral spine and first pelvic-fin ray vs. two 
thirds; its length 20.7-23.9% SL vs. 23.6-27.0), and from H. spinulus in having a longer 
dorsal spine (length 22.6-26.7% SL vs. 14.7-19.8), slenderer body (depth 12.5-14.5% 
SL vs. 17.2-20.9), narrower head (width 23.7-25.2% SL vs. 25.9-30.0), shorter 
posterior process on coracoid (reaching to midway of distance between bases of pec- 
toral spine and first pelvic-fin ray vs. two thirds; length 20.7-23.9% SL vs. 24.9-28.0) 
and more vertebrae (33-35 vs. 30-32). 

Description: Morphometric data in Table 6. Exactly as in Hara filamentosa, 
except for the following. Vertebrae 15+18=33 (2), 16+17=33 (1), 15+19=34 (2), 
16+18=34 (3) or 16+19=35 (3). Dorsal fin with 1,5 ,i (11) rays. Dorsal spine moderately 
long and gently curved; with smooth anterior margin and posterior margin with 6-8 
serrae. Caudal fin with i,7,6,i (11) principal rays. First principal ray of upper caudal-fin 
lobe produced into a filament extending posteriorly for half length of upper caudal lobe 
proper. Anal fin with iv,6,i (10) or v,6,i (1) rays. Pelvic fin with i,5 (11) rays. Pectoral 
fin with I,6,i (11) rays; anterior spine margin with 18-19 small distally directed 
serrations, posterior margin with 13-14 large medially directed serrations. Coracoid 
with well developed posterior processes, extending to midway between base of 
posteriormost pectoral-fin ray and pelvic-fin origin. 

Colour: Head and body light chocolate brown, with a mottling caused by 
numerous smaller, irregular darker brown patches. Belly and ventral surfaces of head 
light brown. Two thin, irregular light brown bars on body: first chevron-shaped and 
located between dorsal- and adipose-fin bases, second thicker than first and located on 
caudal peduncle. Dorsal fin dark mottled brown, except for thin hyaline distal margin. 
Pectoral fin dark brown, with thin hyaline distal margin and either a small ovoid 
hyaline spot or a crescentic median hyaline band on middle third of fin. Pelvic and anal 
fins hyaline, with dark brown bases and subdistal stripes. Adipose fin brown as in body, 



A REVIEW OF THE CATFISH GENUS HARA 



489 



Table 6. Morphometric data for Hara longissima (n=ll: 49.0-55.3 mm SL). 





Holotype 


Range 


Mean±SD 


%SL 








Predorsal length 


46.1 


42.7-46.5 


44.8±1.52 


Preanal length 


67.8 


63.2-69.4 


68.1+1.73 


Prepelvic length 


50.4 


48.1-51.2 


49.6±0.96 


Prepectoral length 


23.0 


20.1-24.9 


22.1+1.29 


Length of dorsal fin base 


19.3 


16.5-19.9 


17.7+1.09 


Dorsal spine length 


25.8 


22.6-26.7 


24 .7± 1.37 


Anal fin length 


14.5 


12.6-15.6 


14.0±1.00 


Pelvic fin length 


16.6 


16.4-21.8 


17.9+1.67 


Pectoral fin length 


32.0 


28.0-33.5 


31.4+1.80 


Pectoral spine length 


29.5 


25.9-31.8 


29.2+1.80 


Caudal fin length 


20.7 


18.8-27.4 


23.6±2.72 


Length of adipose fin base 


15.4 


13.3-16.5 


14.8±0.93 


Dorsal to adipose distance 


11.1 


9.0-12.5 


10.9+1.28 


Post-adipose distance 


16.8 


15.7-18.8 


17.4±1.02 


Caudal peduncle length 


19.9 


18.8-21.3 


19.9±0.85 


Caudal peduncle depth 


5.9 


5.0-6.1 


5.7±0.38 


Body depth at anus 


13.7 


12.5-14.5 


13.4±0.60 


Head length 


28.3 


26.0-29.1 


27.9±0.89 


Head width 


25.2 


23.7-25.2 


24.6±0.47 


Head depth 


21.9 


19.7-22.0 


20.7±0.89 


Length of posterior process on coracoid 


22.9 


20.7-23.9 


22.3±0.97 


%HL 








Snout length 


53.1 


48.4-60.3 


54.9±3.52 


Interorbital distance 


36.6 


32.9-38.3 


36.0+1.34 


Eye diameter 


13.8 


11.3-13.8 


12.8±0.74 


Nasal barbel length 


24.8 


13.8-30.7 


21.0+5.56 


Maxillary barbel length 


84.1 


77.4-99.3 


90.5±7.37 


Inner mandibular barbel length 


41.4 


40.7-50.3 


44.5±3.10 


Outer mandibular barbel length 


57.9 


53.5-72.3 


59.8±6.04 



with lighter coloured distal margin. Caudal fin hyaline, with irregular W-shaped band 
subdistally and with small dark brown flecks randomly distributed throughout fin. Two 
small light brown patches present at bases of caudal-fin lobes in some individuals. 
Maxillary and mandibular barbels light brown, with dark brown annuii. 

Distribution: Hara longissima is known from the Irrawaddy River drainage in 
northern Myanmar (Fig. 2). 

Etymology: The name comes from the Latin longissimus , meaning longest, 
and refers to the relatively long caudal peduncle of this species. An adjective. 



Hara mesembrina sp. nov. 



Fig. 7 



Holotype: UMMZ 245903, 47.3 mm SL; Thailand: Ranong province, Baan Na district, 
hillstreams flowing from Langkatuek, Klong Naka; K. Udomritthiruj , April 2005. 

Paratypes: UMMZ 245602 (9), 31.4-55.3 mm SL; UMMZ 245963 (6). 42.4-55.8 mm 
SL; data as for holotype. CMK 12204 (3), 34.7-40.2 mm SL; Thailand: Ranong, stream N of 
Khura Bhuri, 100 km S of Ranong, 9°17'48"N 98°23 , 50"E; K. Kubota, March 1996. 



490 



H. H. NG & M. KOTTELAT 




FlG. 7 
Hara mesembrina, holotype, UMMZ 245903, 47.3 mm SL; Thailand: hillstreams flowing from 
Langkatuek. Scale bar: 25 mm. 



A REVIEW OF THE CATFISH GENUS HARA 491 

Diagnosis: Hara mesembrina can be distinguished from congeners except 
H. filamentosa and H. longissima in having (vs. lacking) a filamentous extension to the 
first principal ray of the upper caudal-fin lobe. It differs from H. filamentosa in having 
a longer posterior process on coracoid (reaching to two thirds of distance between 
bases of pectoral spine and first pelvic-fin ray vs. midway; length 23.6-27.0% SL vs. 
19.9-22.8) and a longer caudal peduncle (length 17.2-18.9% SL vs. 14.9-17.8), and 
from H. longissima in having a shorter caudal peduncle (length 17.2-18.9% SL vs. 
18.8-21.3), larger eye (diameter 13.1-17.9% HL vs. 11.3-13.8) and longer posterior 
process on coracoid (reaching to two thirds of distance between bases of pectoral spine 
and first pelvic-fin ray vs. midway; length 23.6-27.0% SL vs. 20.7-23.9). Hara mesem- 
brina can be further distinguished from H. hara in having a deeper caudal peduncle 
(depth 6.0-7.4% SL vs. 5.0-5.8) and longer posterior process on coracoid (reaching to 
two thirds of distance between bases of pectoral spine and first pelvic-fin ray vs. mid- 
way; length 23.6-27.0% SL vs. 19.9-24.8), from H. horai in having a deeper caudal 
peduncle (depth 6.0-7.4% SL vs. 5.2-6.2), shorter head (length 22.1-27.3% SL vs. 30.3- 
34.2), longer adipose-fin base (length 13.5-17.0% SL vs. 10.0-13.7) and larger eye 
(diameter 13.1-17.9% HL vs. 8.0-9.8), and from both H.jerdoni and H. minuscula in 
having a shorter posterior process on coracoid (length 23.6-27.0% SL vs. 29.5-38.3), 
more soft pectoral-fin rays (total number of soft fin rays modally 6 vs. 5) and vertebrae 
(32-35 vs. 27-32) and the supraoccipital spine not reaching (vs. reaching) the anterior 
nuchal plate. It further differs from H.jerdoni in having a shorter pectoral spine (length 
26.4-30.1% SL vs. 42.2-51.7), and from H. spinulus in having a longer dorsal spine 
(length 20.2-25.7% SL vs. 14.7-19.8) and caudal peduncle (length 17.2-18.9% SL vs. 
11.8-15.0) and slenderer body (depth 13.1-16.7% SL vs. 17.2-20.9). 

Description: Morphometric data in Table 7. Exactly as in Hara filamentosa, 
except for the following. Vertebrae 16+16=32 (1), 14+19=33 (1), 15+18=33 (7), 
16+17=33 (2), 15+19=34 (3), 16+18=34 (4) or 16+19=35 (1). Dorsal fin with 1,5 ,i (19) 
rays. Dorsal spine moderately long and gently curved; with smooth anterior margin and 
posterior margin with 4-9 serrae. Caudal fin with i,7,7,i (19) principal rays. First prin- 
cipal ray of upper caudal-fin lobe produced into a filament extending posteriorly for 
half length of upper caudal lobe proper. Anal fin with iii,6,i (5), iv,6 (3), iv,6,i (6), iii,7,i 
(2), iv,7 (2) or iv,7,i (1) rays. Pelvic fin with i,5 (19) rays. Pectoral fin with I,6,i (19) 
rays; anterior spine margin with 10-20 small distally directed serrations, posterior 
margin with 8-13 large medially directed serrations. Coracoid with well developed 
posterior process, extending to two thirds distance between base of posteriormost 
pectoral-fin ray and pelvic-fin origin. 

Colour: Head and body chestnut brown to light chocolate brown, with a 
mottling caused by numerous smaller, irregular darker brown patches. Belly and 
ventral surfaces of head light brown. Two thin, irregular light brown bars on body: first 
chevron-shaped and located between dorsal- and adipose-fin bases, second thicker than 
first and located on caudal peduncle. Dorsal fin dark mottled brown, except for thin 
hyaline distal margin. Pectoral fin dark brown, with thin hyaline distal margin and 
either a small ovoid hyaline spot or a crescentic median hyaline band on middle third 
of fin. Pelvic and anal fins hyaline, with dark brown bases and subdistal stripes. 



492 



H. H. NG & M. KOTTELAT 



Table 7. Morphometric data for Hara mesembrina (n=19: 31.4-55.8 mm SL). 





Holotype 


Range 


Mean±SD 


%SL 








Predorsal length 


44.8 


42.0-46.5 


44 .0± 1.32 


Preanal length 


68.8 


64.8-69.3 


67 .3± 1.56 


Prepelvic length 


50.6 


46.4-53.4 


49.3±2.20 


Prepectoral length 


24.5 


19.4-25.0 


22.8±1.79 


Length of dorsal fin base 


19 


17.4-20.2 


18.6±0.99 


Dorsal spine length 


25.7 


20.2-25.7 


22.5±1.53 


Anal fin length 


15.3 


14.0-17.2 


15.3+1.19 


Pelvic fin length 


18.8 


15.2-19.7 


17.8+1.43 


Pectoral fin length 


32.6 


28.4-33.3 


31.1±1.56 


Pectoral spine length 


29.9 


26.4-30.1 


28.2+1.25 


Caudal fin length 


27.0 


21.0-28.6 


25.7±2.10 


Length of adipose fin base 


15.7 


13.5-17.0 


15.0+1.10 


Dorsal to adipose distance 


7.3 


7.1-12.4 


9.5+1.80 


Post-adipose distance 


16.7 


15.0-19.3 


16.8+1.32 


Caudal peduncle length 


18.8 


17.2-18.9 


18.3±0.56 


Caudal peduncle depth 


6.3 


6.0-7.4 


6.6±0.48 


Body depth at anus 


14.4 


13.1-16.7 


14.8+1.08 


Head length 


27.8 


25.8-29.5 


27.8±0.87 


Head width 


25.3 


22.1-27.3 


24.3±1.67 


Head depth 


20.7 


18.7-23.5 


21.6+1.41 


Length of posterior process on coracoid 


23.8 


23.6-27.0 


25.2+1.19 


%HL 








Snout length 


50.4 


39.6-59.6 


50.7+5.74 


Interorbital distance 


35.3 


32.5-36.4 


34.3±1.25 


Eye diameter 


13.5 


13.1-17.9 


14.6+1.48 


Nasal barbel length 


19.5 


8.8-27.2 


17.2±5.03 


Maxillary barbel length 


88.7 


76.3-99.3 


85.0+7.08 


Inner mandibular barbel length 


41.4 


22.9-45.4 


40.9±6.29 


Outer mandibular barbel length 


48.9 


42.9-61.5 


52.8±5.40 



Adipose fin brown as in body, with lighter coloured distal margin. Caudal fin hyaline, 
with irregular W-shaped band subdistally and with small dark brown flecks randomly 
distributed throughout fin. Two small light brown patches present at bases of caudal- 
fin lobes in some individuals. Maxillary and mandibular barbels light brown, with dark 
brown annuii. 

Distribution: Hara mesembrina is known from the small rivers draining the 
southern extremity of the Tenasserim Range (in the Isthmus of Kra in southern 
Thailand) that debouch into the Andaman Sea (Fig. 2). 

Etymology: The name comes from the Greek \iEor\\ifipivoL, (mesembrinos), 
meaning southern, referring to the distribution of this species, which is the southern- 
most of all Hata species. An adjective. 



Hara minuscula sp. nov. 



Fig. 8 



Holotype: CAS 223734, 28.4 mm SL; Myanmar: Bago, Dayame Chaung, 1 .6 km N of 
Daik-U; T. R. Roberts, 9 March 1985. 

Paratypes: CAS 61339 (45), 25.8-30.0 mm SL; data as for holotype. NRM 39883 (23), 
25.0-31.0 mm SL; Myanmar: Bago. roadside stream about 64 km on road from Taungoo to 



A REVIEW OF THE CATFISH GENUS HARA 493 

Nyaunglaybin, 18°19'5"N 96°30'7"E; F. Fang & A. Roos, 19 March 1997. UMMZ 245671 (1), 

30.2 mm SL; Myanmar: from aquarium trade, collector and date unknown. ZRC 43576 (8), 23.3- 

29.3 mm SL; Myanmar: Bago; H. H. Tan, 28 February 1999. 

Diagnosis: Hara minuscula can be distinguished from congeners except H.jer- 
doni in having a longer posterior process on coracoid (reaching to three quarters of dis- 
tance between bases of pectoral spine and first pelvic-fin ray vs. two thirds or less; 
length 29.5-32.9% SL vs. 19.9-28.0), fewer soft pectoral-fin rays (total number of soft 
fin rays modally 5 vs. 6-7) and the supraoccipital spine reaching (vs. not reaching) the 
anterior nuchal plate. It can be distinguished from H. jerdoni in having a shorter 
pectoral spine (length 28.6-34.7% SL vs. 42.2-51.7) and posterior process on coracoid 
(length 29.5-32.9% SL vs. 34.4-38.3) and more vertebrae (30-32 vs. 27-29). Hara 
minuscula can be further distinguished from H . filamentosa in having fewer vertebrae 
(30-32 vs 31-34), a longer pectoral spine (length 28.6-34.7% SL vs. 23.3-29.9) and 
always lacking (vs. sometimes with) a filamentous extension of the first principal ray 
of the upper caudal-fin lobe, from H. hara in having a wider head (width 24.5-27.2% 
SL vs. 22.4-24.3) and lacking (vs. having) serrations on the anterior edge of the pec- 
toral spine, and from H. fiorai in having a shorter head (length 24.8-28.4% SL vs. 30.3- 
34.2). It further differs from H. longissima in having a shorter caudal peduncle (length 
11.7-14.5% SL vs. 18. 8-2 1.3), deeper body (depth 16.0-18.0% SL vs. 12.5-14.5), fewer 
vertebrae (30-32 vs. 33-34) and lacking (vs. having) a filamentous extension of the first 
principal ray of the upper caudal-fin lobe, from H. mesembrina in having a shorter 
caudal peduncle (length 1 1 .7-14.5% SL vs. 17.2-18.9), deeper body (depth 16.0-18.0% 
SL vs. 13.1-16.7) and lacking (vs. having) a filamentous extension of the first princi- 
pal ray of the upper caudal-fin lobe, and from H. spinulus in having a slenderer caudal 
peduncle (depth 4.1-7.0% SL vs. 6.7-8.5). Hara minuscula is also the smallest species 
of Indochinese Hara, with a maximum recorded size of ca. 30 mm SL (vs. ca. 55-67 
mm SL in other Indochinese congeners); compared to Indian congeners, only H. jer- 
doni is smaller (Table 4). 

Description: Morphometric data in Table 8. Exactly as in Hara filamentosa, 
except for the following. Vertebrae 15+15=30 (10), 16+14=30 (1), 15+16=31 (16), 
16+15=31 (7) or 16+16=32 (2). Supraoccipital spine reaching anterior nuchal plate. 
Dorsal fin with 1,5 ,i (36) rays. Dorsal spine moderately long and gently curved; with 
smooth anterior margin and posterior margin with 5-6 serrae. Caudal fin with i,7,74 
(36) principal rays; without filamentous extension to first principal ray of upper lobe. 
Anal fin with iii,6,i (14), iv,5,i (9) or iv,6,i (13) rays. Pelvic fin with with i,5 (36) rays. 
Pectoral fin with 1,44 (17) or 1,5 (19) rays; anterior spine margin with 13-17 small 
distally directed serrations, posterior margin with 9-11 large medially directed 
serrations. Coracoid with well developed posterior process, extending to three quarters 
distance between base of posteriormost pectoral-fin ray and pelvic-fin origin. 

Colour: Head and body dark chestnut brown to light chocolate brown, with a 
mottling caused by numerous smaller, irregular darker brown patches. Belly and 
ventral surfaces of head light brown. Two thin, irregular light brown bars on body: first 
chevron-shaped and located between dorsal- and adipose-fin bases, second thicker than 
first and located on caudal peduncle. Dorsal fin dark mottled brown, except for thin 
hyaline distal margin. Pectoral fin dark brown, with thin hyaline distal margin and 



494 



H. H. NG & M. KOTTELAT 




A REVIEW OF THE CATFISH GENUS HARA 



495 



Table 8. Morphometric data for Hara minuscula (n=36: 25.8-31 .0 mm SL). 





Holotype 


Range 


Mean±SD 


%SL 








Predorsal length 


44.9 


42.9-46.7 


44.7±1.26 


Preanal length 


71.6 


68.8-75.1 


72.3+1.74 


Prepelvic length 


56.1 


52.2-56.1 


54. 2± 1.30 


Prepectoral length 


20.0 


17.9-22.2 


20. 4± 1.32 


Length of dorsal fin base 


20.7 


13.9-21.7 


18.5±2.47 


Dorsal spine length 


20.0 


18.7-22.9 


21.2+1.29 


Anal fin length 


14.7 


13.9-16.7 


15.3±0.93 


Pelvic fin length 


17.5 


13.8-19.7 


17.4+1.63 


Pectoral fin length 


33.3 


30.3-36.5 


33.1+1.92 


Pectoral spine length 


32.6 


28.6-34.7 


31.3+1.68 


Caudal fin length 


34.7 


27.0-35.9 


32.2±2.87 


Length of adipose fin base 


15.4 


12.9-18.4 


15.5+1.48 


Dorsal to adipose distance 


8.4 


8.4-11.6 


9.7±1.07 


Post-adipose distance 


15.1 


12.3-16.2 


14.3±0.99 


Caudal peduncle length 


13.7 


11.7-14.5 


13.5+0.83 


Caudal peduncle depth 


6.7 


4.1-7.0 


5.5±0.92 


Body depth at anus 


16.5 


16.0-18.0 


17.0+0.66 


Head length 


28.1 


24.8-28.4 


26. 8± 1.02 


Head width 


26.3 


24.5-27.2 


25.6±0.79 


Head depth 


24.6 


22.6-26.8 


24.1 + 1.24 


Length of posterior process on coracoid 


32.6 


29.5-32.9 


31.3+1.13 


%HL 








Snout length 


50.0 


45.5-54.5 


50.9±2.40 


Interorbital distance 


41.3 


37.0-42.9 


40.0+1.80 


Eye diameter 


13.8 


10.7-15.6 


12.9±1.42 


Nasal barbel length 


21.3 


17.3-32.5 


22.8±4.71 


Maxillary barbel length 


92.5 


74.0-100.0 


86.5±7.38 


Inner mandibular barbel length 


47.5 


40.0-57.1 


50.1±5.41 


Outer mandibular barbel length 


73.8 


58.2-73.8 


65.0±4.43 



either a small ovoid hyaline spot or a crescentic median hyaline band on middle third 
of fin. Pelvic and anal fins hyaline, with dark brown bases and subdistal stripes. 
Adipose fin brown as in body, with lighter coloured distal margin. Caudal fin hyaline, 
with irregular W-shaped band subdistally and with small dark brown flecks randomly 
distributed throughout fin. Two small light brown patches present at bases of caudal- 
fin lobes in some individuals. Maxillary and mandibular barbels light brown, with dark 
brown annuii. 

Distribution: Hara minuscula is known from the Sittang river drainage 
(Fig. 2). 

Etymology: The name comes from the Latin minusculus, meaning rather small, 
referring to the small size of this species. An adjective. 



Fig. 8 
Hara minuscula, CAS 223734, holotype, 28.4 mm SL; Myanmar: Dayame Chaung. Scale bai" 
10 mm. 



496 H. H. NG & M. KOTTELAT 

Hara spinulus sp. nov. Fig. 9 

Holotype: NRM 52556, 50.3 mm SL; Myanmar: Rakhine state, Baw Di Chaung at Baw 
Di bridge, 32 km from Gwa on road to Ngathaingchaung, 17°34'15"N 94°43'47"E; S. O. 
Kullander et al., 18 March 1998. 

Paratypes: NRM 40750 (3), 34.7-48.5 mm SL; data as for holotype. UMMZ 246173 
(1 ), 49.8 mm SL; Myanmar: Rakhine state, Thandwe Chaung; Than Kyaw Toe, November 2005. 

Diagnosis: Hara spinulus can be distinguished from congeners (except H. mi- 
nuscula) in having a shorter dorsal spine (14.7-19.8% SL vs. 20.2-27.6). It can be 
further distinguished from H. filamentosa in having a shorter adipose-fin base (length 
12.9-16.1% SL vs. 15.7-21.7) and caudal peduncle (length 11.8-15.0% SL vs. 14.9- 
17.8) and always lacking (vs. sometimes with) a filamentous extension of the first prin- 
cipal ray of the upper caudal-fin lobe, from H. hara in having a wider head (width 25.9- 
30.0% SL vs. 22.4-24.3), deeper caudal peduncle (depth 6.7-8.5% SL vs. 5.0-5.8) and 
lacking (vs. having) serrations on the anterior edge of the dorsal spine, and from H. 
horai in having a longer adipose-fin base (length 12.9-16.1% SL vs. 10.0-13.7), short- 
er (length 11.8-15.0% SL vs. 15.5-17.9) and deeper (depth 6.7-8.5% SL vs. 5.2-6.2) 
caudal peduncle, shorter head (length 26.9-28.9% SL vs. 30.3-34.2) and larger eye 
(diameter 1 1 .2-14.4% SL vs. 8.0-9.8). Hara spinulus can be further distinguished from 
both H.jerdoni and H. minuscula in having a deeper caudal peduncle (depth 6.7-8.5% 
SL vs. 4.1-7.0), shorter posterior process on coracoid (reaching to two thirds of 
distance between bases of pectoral spine and first pelvic-fin ray vs. at least three 
quarters; length 24.9-28.0% SL vs. 29.5-38.3), more soft pectoral-fin rays (total 
number of soft fin rays modally 7 vs. 5) and the supraoccipital spine not reaching (vs. 
reaching) the anterior nuchal plate, from H.jerdoni in having a shorter pectoral spine 
(length 27.5-30.3% SL vs. 42.2-53.8) and caudal peduncle (length 11.8-15.0% SL vs. 
15.1-18.4), more vertebrae (30-32 vs. 27-29), from H. mesembrina in having a shorter 
caudal peduncle (length 11.8-15.0% SL vs. 17.2-18.9), a deeper body (depth 17.2- 
20.9% SL vs. 13.1-16.7) and lacking (vs. having) a filamentous extension of the first 
principal ray of the upper caudal-fin lobe, and from H. longissima in having a deeper 
body (depth 17.2-20.9% SL vs. 12.5-14.5), wider head (width 25.9-30.0% SL vs. 23.7- 
25.2), longer posterior process on coracoid (reaching to two thirds of distance between 
bases of pectoral spine and first pelvic-fin ray vs. midway; length 24.9-28.0% SL vs. 
20.7-23.9), fewer vertebrae (30-32 vs. 33-35) and lacking (vs. having) a filamentous 
extension of the first principal ray of the upper caudal-fin lobe. 

Description: Morphometric data in Table 9. Exactly as in Hara filamentosa, 
except for the following. Vertebrae 15+15=30 (1), 15+16=31 (1), 15+17=32 (2) or 
16+16=32 (1). Dorsal fin with 1,54 (1), 1,6 (3) or 1,64 (1) rays. Dorsal spine modera- 
tely short and gently curved; with smooth anterior margin and posterior margin with 3- 
5 serrae. Caudal fin strongly forked, with i,7,7,i (2) or i,7,8,i (3) principal rays; without 
filamentous extension to first principal ray of upper lobe. Anal fin with iv,64 (3), iv,7 



Fig. 9 
Hara spinulus, holotype, NRM 52556, 50.3 mm SL; Myanmar: Baw Di Chaung. Scale bar: 25 



A REVIEW OF THE CATFISH GENUS HARA 



497 




498 



H. H. NG & M. KOTTELAT 



Table 9. Morphometric data for Hara spinulus (n=5: 34.7-50.3 mm SL). 





Holotype 


Range 


Mean±SD 


%SL 








Predorsal length 


41.3 


39.4-45.8 


43.6±3.01 


Preanal length 


71.4 


68.6-71.8 


70.4±1.36 


Prepelvic length 


52.7 


52.0-55.3 


53. 2± 1.26 


Prepectoral length 


21.9 


21.4-23.7 


22 .5± 1.07 


Length of dorsal fin base 


21.9 


19.1-22.9 


20.7±1.67 


Dorsal spine length 


18.5 


14.7-19.8 


17.4±2.18 


Anal fin length 


16.9 


15.3-18.3 


16.5+1.26 


Pelvic fin length 


17.5 


15.8-18.1 


16.9±0.96 


Pectoral fin length 


30.6 


29.3-33.3 


31.0+1.47 


Pectoral spine length 


29.2 


27.5-30.3 


29. 0± 1.00 


Caudal fin length 


26.4 


24.5-26.8 


25.9±0.90 


Length of adipose fin base 


13.5 


12.9-16.1 


14.5±1.42 


Dorsal to adipose distance 


8.3 


7.8-11.3 


9.3+1.58 


Post-adipose distance 


14.7 


12.4-17.7 


14.9+1.89 


Caudal peduncle length 


12.9 


11.8-15.0 


13.8±1.40 


Caudal peduncle depth 


8.5 


6.7-8.5 


7.4±0.66 


Body depth at anus 


20.9 


17.2-20.9 


18.5+1.51 


Head length 


28.8 


26.9-28.9 


28.2±0.80 


Head width 


29.0 


25.9-30.0 


28.6+1.69 


Head depth 


23.7 


22.7-24.7 


23.7±0.86 


Length of posterior process on coracoid 


27.6 


24.9-28.0 


26.5+1.31 


%HL 








Snout length 


50.3 


47.4-52.9 


50.4+2.04 


Interorbital distance 


33.6 


33.6-37.3 


35.7+1.71 


Eye diameter 


11.6 


11.2-14.4 


12.4+1.25 


Nasal barbel length 


32.9 


22.1-32.9 


27!±4.14 


Maxillary barbel length 


85.6 


72.5-104.5 


91.3+13.01 


Inner mandibular barbel length 


52.7 


43.1-53.7 


49.0±4.40 


Outer mandibular barbel length 


61.6 


56.9-68.7 


61.3±4.49 



(1) or iv,7,i (1) rays. Pelvic fin with i,5 (4) rays. Pectoral fin with 1,6 (1) or 1,64 (4) rays; 
anterior spine margin with 11-18 small distally directed serrations, posterior margin 
with 7-11 large medially directed serrations. Coracoid with well developed posterior 
processes, extending to two thirds distance between base of posteriormost pectoral-fin 
ray and pelvic-fin origin. 

Colour: Head and body terracotta to light chocolate brown, with a mottling 
caused by numerous smaller, irregular darker brown patches. Belly and ventral sur- 
faces of head light brown. Two thin, irregular light brown bars on body: first located 
between dorsal- and adipose-fin bases, second thicker than first and located on caudal 
peduncle. Dorsal fin dark mottled brown, except for thin hyaline distal margin. Pectoral 
fin dark brown, with thin hyaline distal margin and either a small ovoid hyaline spot or 
a crescentic median hyaline band on middle third of fin. Pelvic and anal fins hyaline, 
with dark brown bases and subdistal stripes. Adipose fin brown as in body, with lighter 
coloured distal margin. Caudal fin hyaline, with irregular W-shaped band subdistally 
and with small dark brown flecks randomly distributed throughout fin. Two small light 
brown patches present at bases of caudal-fin lobes in some individuals. Maxillary and 
mandibular barbels light brown, with dark brown annuii. 



A REVIEW OF THE CATFISH GENUS HARA 499 

Distribution: Hara spinulus is known from the Baw Di Chaung drainage, 
which drains the western face of the Rakhine (Arakan) Yoma, in southwestern 
Myanmar (Fig. 2). 

Etymology: The name comes from the diminutive form of the Latin spina, 
meaning thorn, and refers to the short dorsal spine of this species. A noun in apposition. 

KEY TO THE SPECIES OF HARA 

1 Caudal fin emarginate; vertebrae 27-29 [Brahmaputra and Ganges river 

drainages] H. jerdoni 

Caudal fin deeply forked; vertebrae 30-35 2 

2 Serrations on anterior edge of dorsal spine present [Ganges and 

Brahmaputra river drainages] H. hara 

Serrations on anterior edge of dorsal spine absent 3 

3 Supraoccipital process reaching anterior nuchal plate; posterior process 
on coracoid extending three quarters distance between bases of pectoral 
spine and pelvic fins; total soft pectoral-fin rays modally 5 [Sittang River 

drainage] H. minuscula 

Supraoccipital process not reaching anterior nuchal plate; posterior 
process on coracoid extending two thirds or less distance between bases 

of pectoral spine and pelvic fins; total soft pectoral-fin rays modally 6-7 .... 4 

4 Length of adipose-fin base 10.0-13.7% SL; eye diameter 8.0-9.8% HL 

[Brahmaputra River drainage] H. horai 

Length of adipose-fin base 12.9-17.0% SL; eye diameter 11.2-17.9% HL . . . 5 

5 Length of dorsal spine 14.7-19.8% SL; first principal ray on upper lobe 
of caudal fin not extended into filament [Baw Di Chaung drainage] 

H. spinulus 

Length of dorsal spine 20.2-27.6% SL; first principal ray on upper lobe 

of caudal fin usually extended into filament 6 

6 Posterior process on coracoid reaching to two thirds distance between 
bases of pectoral spine and first pelvic-fin ray, its length 23.6-27.0% SL 
[rivers draining southern extremity of Tenasserim Range] ....//. mesembrina 
Posterior process on coracoid reaching to midway of distance between 
bases of pectoral spine and first pelvic-fin ray, its length 19.9-23.9% SL . . . . 7 

7 Caudal peduncle length 14.9-17.8% SL, depth 6.1-7.2% SL (depth 2.1- 
2.9 times in its length); body depth 13.9-19.2% SL [Ataran, Salween and 

Sittang river drainages] H. filamentosa 

Caudal peduncle length 18.8-21.3% SL, depth 5.0-6.1% SL (depth 3.1- 

3.9 times in its length); body depth 12.5-14.5% SL [Irrawaddy River 
drainage] H. longissima 

DISCUSSION 

Although Thomson & Page (2006) consider Erethistes and Hara to be 
synonyms (the former being the senior name), a phylogenetic analysis of the Sisoridae 
using 190 morphological and 1371 molecular characters carried out as part of HHN's 



500 H. H. NG & M. KOTTELAT 

doctoral dissertation (Ng, 2006) indicates that Hara (including Erethistes) is para- 
phyletic. The analysis of the concatenated dataset indicates that the species currently 
assigned to Hara can be divided into three clades: one consisting of H. filamentosa (the 
type species of Hara)+H. mesembrina, another consisting of H . jerdoni+H . minuscula, 
and another consisting of Erethistes pusillus and all other species of Hara. However, 
the three clades were not recovered in the analysis of the morphological dataset alone 
(most species of Hara were recovered in a polytomy with Erethistoides and 
Ayarnangra) and no morphological synapomorphies could be found to diagnose the 
three clades. The apparent paraphyly of Hara and the conflicting nature of the evidence 
are currently being further investigated and the complete results of the analysis will be 
published elsewhere. The synonymization of Hara with Erethistes by Thomson & Page 
(2006) does not truly reflect the phylogenetic relationships of this group of catfishes. 
Therefore, pending publication of the results of a fine-scale phylogenetic analysis of 
the group, we follow the sole current usage in the literature in recognizing Erethistes 
and Hara as distinct genera. We see no advantage in substituting one non-phylogenetic 
classification scheme with another. 

The species-level taxonomy of Hara was confused, with only one attempt to 
review the genus (Hora, 1950). We consider only three valid species to occur on the 
Indian subcontinent: H. hara (Hamilton, 1822), H. horai Misra, 1976 and H.jerdoni 
Day, 1870. Both H. saharsai Datta Munshi & Srivastava, 1988 and H. serrata 
Vishwanath & Kosygin, 2000 are considered junior synonyms of//, hara because we 
could not find any significant differences in comparing the original descriptions and 
material we identified as H. hara. Hara saharsai (described from the Ganges River 
drainage in northern Bihar, which is within the recorded range for H. hara) is not 
adequately diagnosed from H. hara in the original description. The only difference is 
mentioned in the key to the species (Datta Munshi & Srivastava, 1988: 263), and states 
that H. hara has a pectoral spine shorter than head length, while H. saharsai has a 
pectoral spine longer than head length. Our examination of material indicates that the 
pectoral spine in H. hara is equal to, or very slightly longer than head length and 
because we could not find other characters to distinguish between the two species, we 
consider them to be conspecific. Hara serrata (described from the Jiri River, a tribu- 
tary of the Barak River, itself a tributary of the Brahmaputra River, which is within the 
recorded range for H. hara) is diagnosed as a distinct species based on the serrated 
anterior edge of the dorsal spine, but our examination of H. hara shows that it also has 
the dorsal spine serrated anteriorly. No other characters could be found to distinguish 
H. serrata and H. hara, and we also consider the two conspecific here. 

Hara filamentosa was originally described from the Tenasserim Provinces, 
which was a British administrative unit of colonial Burma (Myanmar) consisting of the 
coastal area east and south of the Irrawaddy Delta [not to be confused with the present 
administrative unit of Tanintharyi (Tenasserim), which constitutes only a fraction of 
the former Tenasserim Provinces] . The Tenasserim Provinces include the present-day 
states of Kayah, Kayin, Mon, Bago (partially) and possibly Tanintharyi in Myanmar 
(Fang & Kottelat, 1999). We have discussed the actual locality of the fishes described 
from "Tenasserim" by Blyth in an earlier work (Ng & Kottelat, 2001), and hypothesize 
that much of this material comes from either the Sittang River drainage or from the 



A REVIEW OF THE CATFISH GENUS HARA 501 

drainages to the south of it. The original description does not state on how many spec- 
imens it is based and Blyth gives only one size (3 inches) which could mean either that 
he examined a single specimen (which would then be the holotype) or several for 
which only the size of the largest is given, a common practice of the time. Edward 
Blyth was the curator of the museum of the Asiatic Society of Bengal, and types of the 
fishes he described were initially deposited at the Society's museum; this material was 
subsequently transferred to the Indian Museum (now the Zoological Survey of India 
(Whitehead & Talwar, 1976). 

Hora (1950) examined six specimens of Hara (ZSI 585) in the ZSI collected by 
Berdmore from "Tenasserim" (Major Hugh Berdmore [May 1812-30 May 1859] 
collected in Tenasserim the type series of several species described by Blyth). As the 
description of H '. filamentosa appeared after Berdmore's death (apparently in U.K.) it 
seems that part (or all) of ZSI 585 could have been used in the original description of 
H. filamentosa. Hora (1950) reidentified five of the six specimens as H . filamentosa 
(which he considered conspecific with H. hara), and the remaining specimen as 
Erethistes pusillus. Other than the size of the specimen reidentified as E.pusillus (39.8 
mm SL, which is smaller than the the size reported by Blyth; possibly the size of his 
largest specimens if he had more than one), no other indication of size was given by 
Hora (1950) for ZSI 585. Hora (1950: 201) commented that "it appears ... that Day's 
[1877: pi. 102 fig. 4] figures of E. conta ... were made from one of these specimens 
[ZSI 585]". Day (p. 453) commented that "Berdmore sent 5 or 6 specimens to the 
Calcutta Museum". This creates some doubts as to whether all of the specimens in ZSI 
585 may indeed be the type(s) of H. filamentosa. 

We note that the specimen out of ZSI 585 that Hora identified as E. pusillus is 
the only specimen of the species ever recorded from Myanmar and this casts doubts as 
to its identity or origin. Erethistes pusillus is otherwise known only from the 
Brahmaputra and Ganges river drainages. Since the drawing of the pectoral spine of 
this specimen in Hora (1950: fig. 1) is unmistakeably that of E.pusillus, this means that 
the material in ZSI 585 almost certainly was obtained from at least two different areas 
(southern Myanmar and somewhere in the Ganges or Brahmaputra river drainage). 
This has implications for the stability of nomenclature (see below). 

The first author visited the ZSI twice, but was neither able to examine ZSI 585 
nor successfully locate the type material of H. filamentosa. This type material is also 
not mentioned in the catalogue of fish types in that collection (Menon & Yazdani, 
1968). The fish collections of the ZSI suffered extensive loss and damage as a result of 
a flood when the collection was temporarily housed in Benares during the Second 
World War (Chopra, 1947; Whitehead & Talwar, 1976), and the type(s) of H. filamen- 
tosa might have been lost at that time or is(are) mislabelled or misplaced. It is not 
known if Hora examined ZSI 585 before or after the war. There is no known other ma- 
terial that could possibly be types of H. filamentosa. Our studies indicate that several 
species of Hara with a filamentous extension of the dorsalmost principal caudal-fin ray 
occur in Myanmar and Thailand; only one is known from southern Myanmar (in the 
former "Tenasserim Provinces") to date, and we identify it as H. filamentosa. However, 
problems remain in objectively defining H. filamentosa with the uncertainties sur- 
rounding the type series, the number of types originally included, the possible type 



502 H. H. NG & M. KOTTELAT 

status and apparent loss of ZSI 585, and the vague type locality. As two species are 
recorded by Hora in the potential type series, the status of the name can only be cleared 
by a lectotype or a neotype designation. 

The designation of a neotype is needed to define the nominal taxon objectively. 
We designate MHNG 2687.038, 43.7 mm SL, from the Ataran drainage in Myanmar, 
as neotype of H. filamentosa, as diagnosed and described above. It agrees with Blyth's 
original description, with Day's drawing of a putative syntype, and it comes from 
within the area that can be considered as the original type locality. 

Kottelat (1983) described Erethistes maesotensis from the Salween River 
drainage in western Thailand. The only difference between Erethistes and Hara is the 
direction of the serrations on the anterior edge of the pectoral spine (divergent in 
Erethistes and antrorse in Hara; Hora, 1950, de Pinna, 1996). The serrations in the pec- 
toral spine of E. maesotensis (Kottelat, 1983: fig. 2b) are not truly divergent, but have 
a few that are retrorse or anteriorly directed, instead of all the serrations being antrorse 
as typically seen in other Hara species. This is a condition commonly seen in juvenile 
Hara, and as such, we attach no significance to its use as a diagnostic character. We 
were unable to find any significant differences between the material of H. filamentosa 
from the Salween River drainage in western Thailand (most collected at or very near 
the type locality of E. maesotensis) with the material we identify as H. filamentosa 
from Myanmar, although it should be noted that most of the Salween material we have 
examined (58 out of 65 specimens) lack the filamentous extension to the dorsalmost 
principal caudal-fin ray. 

Of the other material from Myanmar reported as H. hara by Hora (1950), the 
material from Meetan [=Mitan Chaung] (ZSI Fl 1049/1) is most likely H. filamentosa, 
while that from Indawgyi Lake (ZSI F10878/1) and Mandalay (ZSI 582) is most likely 
H. longissima (the probable identities of these three lots are surmised from the 
collection localities). The specimen reported from Bassein (ZSI 1453) is almost cer- 
tainly H. minuscula, as this specimen was reported as having a proportionately longer 
pectoral spine (a relatively long pectoral spine is a diagnostic character of this species). 

Because H. filamentosa and H. minuscula occur sympatrically in the Sittang 
river drainage, it is necessary to rule out the possibility that the smaller H. minuscula 
are not merely juvenile H. filamentosa. The most striking difference is the extent of the 
supraoccipital process; it reaches the anterior nuchal plate in H. minuscula while it 
does not reach it in H. filamentosa. We note that the contact between the supraoccipital 
process and the anterior nuchal plate is independent of ontogeny, as it remains constant 
for all Hara species (from India, Bangladesh, Thailand and Myanmar) of all size 
ranges we have examined (16-85 mm SL; see individual species account for explicit 
details of size ranges of each species). 

The number of vertebrae also distinguishes the two species (Table 10). While 
the number of abdominal vertebrae does not really distinguish Hara species, H. mi- 
nuscula differs from H. filamentosa by the number of caudal vertebrae (14-16, vs. 17- 
19) and the total number of vertebrae (30-32, vs. 31-34). Hara minuscula is distin- 
guished from all other Indochinese Hara species in having most specimens (29 out of 
36) with more abdominal vertebrae than caudal or equal number of both; caudal 
vertebrae outnumber abdominal vertebrae in only 7 specimens out of 36 examined 
(Table 10). In H. filamentosa there are 2 to 5 more caudal than abdominal vertebrae. 



A REVIEW OF THE CATFISH GENUS H ARA 



503 



^•'tON'iOWOj. 
rM <N ^- 1 _i ,_ i _ iffi^f 






i ' ' (N 

HNfsH 

, rt M H 



LT, 

en 

^ on Ti ln^ 

£ O O ,_ m © 

cm ^. -h 



(T) — i 



fi s 



r^, 



m 



"* r-î O 



co ^_, 



ÏÏ fi "t - 1 



i_ r- ^t cN -h " 



CN £i CN 



^ r-~ ^ 



r^oo^ 



^J Os ^ CN oo Tt y 



2, Tf 00 00 's!- ' <— I 



a s 



| . | £ s s» 

~ S °5 " ^ S 

J I S * § e | 5, 



504 H. H. NG & M. KOTTELAT 

Hara minuscula possesses fewer soft pectoral fin rays (modally 5 vs. 6-7) than 
most congeners. This reduction also occurs only in H. jerdoni, another diminutive 
Hara species, and we treat this reduction as a pedomorphic character associated with 
miniaturization (see discussion in Weitzman & Vari, 1988). 

The original description of H. horai Misra (1976: 245, pi. 9 fig. 1) does not 
include explicit information on type material and Tilak & Talwar (1976: 246) 
designated a neotype. This neotype designation is not valid because a neotype can only 
be designated if all primary types (holotype, lectotype, neotype or syntypes) are lost 
(ICZN art. 75.3). Tilak & Talwar assumed that there was a holotype and that it is lost. 
This is erroneous because Misra did not state that he had a single specimen. All 
specimens on which Misra's H. horai is based are syntypes. These syntypes include all 
specimens examined by Misra [if any] and the specimens of//, hara sensu Hora (1950: 
200) explicitly listed by Misra as belonging to the present species, that is those 
collected by Shaw & Shebbeare in Terai and Duars; Hora (1950: 201) listed only ZSI 
Fl 1390/1 [2]; so there are at least 2 syntypes and one of them is still extant (the one 
Tilak & Talwar designated as neotype). Therefore, the Code does not permit the 
designation of a neotype. This surviving syntype is designated here as lectotype of 
H. horai. 

The length of the posterior process on coracoid is a useful diagnostic character 
for Hara species. Its use in diagnosing members of the Erethistidae is reported here for 
the first time, although it has previously been shown to be useful in diagnosing species 
of the Aspredinidae, another siluriform group with a prominent posterior process on the 
coracoid (Mees, 1989). 

ACKNOWLEDGEMENTS 

We thank James Maclaine (BMNH), David Catania (CAS), Sonia Fisch-Muller, 
Claude Weber (MHNG), Sven Kullander (NRM), Douglas Nelson (UMMZ), Kelvin 
Lim (ZRC) and A. K. Karmakar (ZSI) for permission to examine material under their 
care. Thanks are also due to Katsuma Kubota for assistance in the field and gift of 
material, Kamphol Udomritthiruj and U Tin Win for providing material for study, and 
an anonymous reviewer for useful comments. Funding from the Horace H. Rackham 
School of Graduate Studies, University of Michigan to HHN is gratefully 
acknowledged. 

REFERENCES 

Blyth, E. 1860. Report on some fishes received chiefly from the Sitang River and its tributary 
streams, Tenasserim Provinces. Journal of the Asiatic Society of Bengal 29: 138-174. 

Chopra, B. 1947. Zoological Survey of India, 1942-1945. Records of the Indian Museum 44: 
347-355. 

Datta Munshi, J. S. & Srivastava, M. R 1988. Natural history of fishes and systematics of 
freshwater fishes of India. Narendra Publishing House, Delhi, 403 pp. 

Day, F. 1870. Notes on the genus Hara. Journal of the Asiatic Society of Bengal 39: 37-40. 

Day, F. 1875-78. The fishes of India; being a natural history of the fishes known to inhabit the 
seas and fresh waters of India, Burma, and Ceylon. Quaritsch, London, (1875): 1-168, 
pis 1-40; (1876): 169-368, pis 41-78; (1877): 369-552, pis 79-138; (1878): i-xx, 
553-778, pis 139-195. 



A REVIEW OF THE CATFISH GENUS HARA 505 



de Pinna, M. C. C. 1996. A phylogenetic analysis of the Asian catfish families Sisoridae, 

Akysidae, and Amblycipitidae, with a hypothesis on the relationships of the neotropical 

Aspredinidae (Teleostei, Ostariophysi). Fieldiana. Zoology (New Series) 84: 1-83. 
Fang, F. & Kottelat, M. 1999. Danio species from northern Laos, with descriptions of three 

new species (Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters 10: 

281-295. 
Hora, S. L. 1950. Siluroid fishes of India, Burma and Ceylon. XIII. Fishes of the genera 

Erethistes Müller and Troschel, Hara Blyth and of two new allied genera. Records of the 

Indian Museum Al: 183-202. 
Husain, A. & Tilak, R. 1978. On the redescription and distribution of Hara jerdoni Day 

(Sisoridae: Siluriformes). Journal of the Inland Fisheries Society of India 9: 92-91. 
Kottelat, M. 1983. A new species of Erethistes Müller & Troschel from Thailand and Burma 

(Osteichthyes: Siluriformes: Sisoridae). Hydrobiologia 107: 71-74. 
Kottelat, M. 1989. Zoogeography of fishes from Indochinese inland waters with an annotated 

check-list. Bulletin Zoologisch Museum Universiteit van Amsterdam 12: 1-55. 
M'Clelland, J. 1844. Description of a collection of fishes made at Chusan andNingpo in China, 

by Dr. G. R. Playfair, Surgeon of the Phlegethon, War Steamer, during the late Military 

operations in that country. Calcutta Journal of Natural History, Miscellaneous Arts and 

Sciences in India A: 390-413, pis 21-25. 
Mees, G. F. 1989. Notes on the genus Dysichthys, subfamily Bunocephalidae, family 

Aspredinidae (Pisces, Nematognathi). Proceedings of the Koninklijke Nederlandse 

Akademie van Wetenschappen (Series C. Biological and Medical Sciences) 92: 189-250. 
Menon, A. G. K. & Yazdani, G. M. 1968. Catalogue of type-specimens in the Zoological Survey 

of India, Part 2. Fishes. Records of the Zoological Survey of India 61: 91-190. 
Misra, K. S. 1976. The fauna of India and adjacent countries. Pisces. Vol. III. Teleostomi: 

Sypriniformes: Siluri. Zoological Survey of India, Calcutta, 367 pp. 
Ng, H. H. 2006. A Phylogenetic Analysis of the Asian Catfish Family Sisoridae (Teleostei: 

Siluriformes), and the Evolution of Epidermal Characters in the Group. PhD dissertation, 

University of Michigan, Ann Arbor, 448 pp. 
Ng, H. H. & Kottelat, M. 1998. Hyalobagrus, a new genus of miniature bagrid catfish from 

Southeast Asia (Teleostei: Siluriformes). Ichthyological Exploration of Freshwaters 9: 

335-346. 
Ng, H. H. & Kottelat, M. 2001. A review of the genus Batasio (Teleostei: Bagridae) in 

Indochina, with the description of B. tigrinus n. sp. from Thailand. Revue suisse de 

Zoologie 108:495-511. 
Roberts, T. R. 1994. Systematic revision of Asian bagrid catfishes of the genus Mystus sensu 

stricto, with a new species from Thailand and Cambodia. Ichthyological Exploration of 

Freshwaters 5: 241-256. 
Thomson, A. W. & Page, L. M. 2006. Genera of the Asian Catfish Families Sisoridae and 

Erethistidae(Teleostei: Siluriformes). Zootaxa 1345: 1-96. 
Tilak, R. & Talwar, P. K. 1976. A taxonomic reassessment of Hara horai Misra (Pisces: 

Sisoridae) with a designation of neotype. Newsletter of the Zoological Survey of India 2: 

245-247. 
Ukkatawewat, S. & Vidthayanon, C. 1998. [Fishes of the Salween River basin, Thailand]. 

Aquatic Natural Resources Museum, Technical Paper 2: 1-89 [Thai, English abstract]. 
Vinciguerra, D. 1890. Viaggio di Leonardo Fea in Birmania e regioni vicine XXIV. Pesci. 

Annali del Museo Civico di Storia Naturale di Genova (Serie 2a) 9: 129-362. 
Vishwanath, W. & Kosygin, L. 2000. On a new species of the genus Hara Blyth from Manipur, 

India. Indian Journal of Fisheries Al: 143-147. 
Weitzman, S. H. & Vari, R. P. 1988. Miniaturization in South American freshwater fishes; an 

overview and discussion. Proceedings of the Biological Society of Washington 101: 

444-465. 
Whitehead, P. J. P. & Talwar, P. K. 1976. Francis Day (1829-1889) and his collection of Indian 

fishes. Bulletin of the British Museum (Natural History) Historical Series 5: 1-189. 



Revue suisse de Zoologie 114 (3): 507-511; septembre 2007 



Deroplatys indica, nouvelle espèce de l'Inde 
(Dictyoptera, Mantodea) 

Roger ROY 

Muséum national d'Histoire naturelle, Entomologie, 45 rue Buffon, F - 75005 Paris. 

Email: entomol@mnhn.fr 



Deroplatys indica, new species from India (Dictyoptera, Mantodea). - A 

new species of Deroplatys is described after a single male from India, 
preserved in the Muséum d'Histoire naturelle de la ville de Genève. 

Keywords: Dictyoptera - Mantodea - Mantidae - Deroplatyinae - 
Deroplatys - région orientale. 

INTRODUCTION 

Le genre Deroplatys Westwood, 1839, comprend dans l'état actuel des connais- 
sances (Anisyutkin, 1998; Ehrmann, 2002) onze espèces répandues en Asie du Sud-Est 
depuis le Myanmar jusqu'à la Nouvelle-Guinée, de taille moyenne à grande, de colo- 
ration générale brune, bien ailées dans les deux sexes, qui se caractérisent surtout par 
des expansions foliacées latérales sur le pronotum, de forme différente suivant les 
espèces et les sexes dans chacune. La tête est sans grandes particularités avec les yeux 
latéralement arrondis, les pattes antérieures sont de proportions ordinaires pour des 
Mantes, avec un nombre d'épines moyen, tandis que les pattes médianes et 
postérieures ont les fémurs avec un lobe préapical plus ou moins développé, ainsi 
qu'une épine géniculaire. 

Le spécimen étudié ici, qui fait partie de la collection du Muséum d'Histoire 
naturelle de la ville de Genève (MHNG), avait déjà été repéré par 1 'eminent spécialiste 
des Mantes Max Beier lors de son passage au Muséum en octobre 1973, et il s'était 
contenté de lui ajouter une étiquette imprimée à son nom avec la mention manuscrite 
«Deroplatys â spec. vie. Saravaka Westw.». En effet son pronotum ressemble surtout 
à celui du type femelle de D. sarawaca Westwood, 1889, espèce décrite de Borneo, 
jamais citée d'autres régions et dont les mâles n'ont pas encore été signalés à ma 
connaissance. Mais les similitudes s'arrêtent là, le nouveau spécimen différant de 
sarawaca et aussi de toutes les autres espèces connues de Deroplatys par l'absence de 
lobes préapicaux aux fémurs médians et postérieurs, et par sa localisation en Inde, ce 
qui accroît notablement vers l'ouest l'aire de répartition du genre. De plus, la forme de 
son pronotum apparaît inhabituelle pour un mâle. Cependant, toutes ses autres particu- 
larités étant conformes à la diagnose de Deroplatys, l'espèce qu'il représente doit bien 
se situer dans ce genre. 



Manuscrit accepté le 02.03.2007 



508 



R. ROY 



DESCRIPTION 

Deroplatys indica n. sp. Figs 1-9 

Holotype S: India (Kerala), Cardamon Hills, Periyar, env. de l'hôtel Aranga, 
959 m, 4.II.1972, leg. R. Mussard, C. Besuchet & I. Lobi, genitalia R. Roy n°4043, 
collection MHNG, seul spécimen connu. 

Longueur du corps 60 mm, coloration générale brun sombre, organes du vol 
dépassant largement l'extrémité de l'abdomen (Fig. 1). 




Fig. 1 
Deroplatys indica n. sp., â holotype, x 1,5. 



Tête (Fig. 2) longue d'environ 5 mm et large de 6,5 mm, avec les yeux régu- 
lièrement arrondis très saillants et les ocelles bien développés. Vertex à peine bombé 
entre ses sillons latéraux. Antennes longues d'environ 30 mm avec les deux premiers 
articles brun clair, les suivants plus fins, progressivement plus sombres, courtement 



DEROPLATYS INDICA, NOUVELLE ESPECE DE L INDE 



509 




Figs 2-9 
Deroplatys indica n. sp., 6 holotype. (2) tête. (3) pronotum. (4) région apicale du fémur médian 
gauche et base du tibia en vue dorsale. (5) région apicale du fémur postérieur gauche en vue 
dorsale. (6) plaque suranale avec la base des cerques. (7) région apicale d'un cerque. (8) partie 
postérieure de la plaque sous-génitale avec les styles. (9) genitalia en vue ventrale. 



510 R.ROY 

ciliés. Ecusson frontal un peu plus large que haut, à bord supérieur arrondi, finement 
rebordé, de teinte relativement claire. Clypéus, labre et pièces buccales y compris les 
palpes brun sombre. 

Pronotum (Fig. 3) long de 21 mm dont 15,5 mm pour la métazone et large de 
14 mm, avec les bords foliacés formant deux élargissements successifs, le second plus 
grand et à bords moins denticulés. Prosternum assombri au niveau de l'insertion des 
hanches, sa métazone relativement claire avec une série d'une dizaine de doubles 
taches sombres, les dernières étant plus allongées. 

Pattes antérieures conformées comme celles des autres espèces de Deroplatys. 
Hanches longues de 12,5 mm à face dorsale brune avec deux grandes taches beiges et 
à face ventrale d'un brun plus clair parsemée de petits granules beiges, les lobes 
apicaux étant brun-noir et se recouvrant partiellement; leur bord externe avec 6 ou 7 
grandes épines aplaties à base brun sombre entre lesquelles des plus petites concolores. 
Fémurs longs de 15 mm, brun assez sombre avec des marbrures du côté dorsal, brun 
clair du côté ventral avec des parties brun sombre dont une grande tache au niveau des 
huit premières épines internes; 4 épines discoïdales très sombres, 4 épines externes, les 
deux premières étant plus sombres, et 14-15 épines internes, les quatre premières et 
ensuite seulement les plus grandes brun-noir. Tibias longs de 8 mm avec l'épine 
terminale, brun marbré, armés de 9-10 épines externes et 15 épines internes, les 
dernières plus sombres. Premier article des tarses long d'environ 4,5 mm, les suivants 
courts (les deux derniers ont été perdus). 

Pattes médianes et postérieures grêles, d'un brun un peu marbré, sans lobes 
préapicaux aux fémurs (Figs 4 et 5) dont la région terminale est seulement arquée et un 
peu élargie, et se termine par des prolongements dorsal et ventral relativement longs, 
en pointe mousse pour les médians, longs de 13 mm, en arrondi plus large pour les 
postérieurs, longs de 15 mm; l'épine géniculaire est présente, mais très courte. Les 
tibias, longs de 12 mm pour les médians, de 16 mm pour les postérieurs, sont très 
minces. Le premier article des tarses est environ aussi long que les suivants réunis. 

Elytres longs de 47 mm, brun sombre dans l'aire costale, large de 3,5 mm, et à 
l'avant de l'aire discoidale, d'un brun plus clair ensuite avec toutefois l'apex plus 
foncé; stigma beige clair, très apparent. Ailes longues de 43 mm, brun clair puis sombre 
dans l'aire costale et l'avant de l'aire discoidale, laquelle présente une éclaircie 
préapicale, brun sombre avec les nervures transverses soulignées de clair dans l'aire 
anale. 

Abdomen d'un brun sombre presque uniforme, tergites avec de petits lobes 
latéraux. Plaque suranale (Fig. 6) brune, transverse avec le bord postérieur arrondi à 
peine échancré; cerques grêles, apparemment de 17 articles, le premier brun sombre, 
les suivants clairs, d'abord courts, puis plus longs, le dernier environ deux fois et demie 
plus long que large (Fig. 7). Plaque sous génitale très grande, un peu plus longue que 
large, d'un brun un peu marbré, portant des petits styles plus clairs (Fig. 8). 

Genitalia (Fig. 9) avec l'hypophallus terminé par un prolongement double, en 
pointe et en lobe arrondi, beaucoup plus grand que chez D. desiccata Westwood, 
espèce type du genre; épiphallus droit sans particularités marquantes; titillateur plutôt 
court, renflé dans sa partie moyenne; phallus et pseudophallus très allongés, ce dernier 
en pointe courbe. 



DEROPLATYS INDICA, NOUVELLE ESPECE DE L'iNDE 5 1 1 



DISCUSSION 



Cette nouvelle espèce est bien caractéristique par la forme de son pronotum et 
par l'absence de lobes préapicaux aux fémurs médians et postérieurs, lobes présents 
chez les autres espèces, petits chez les mâles, beaucoup plus grands chez les femelles; 
on ne peut donc la confondre avec aucune autre actuellement connue. Cette absence ne 
me semble pas suffisante à elle seule pour exclure l'espèce du genre Deroplatys, tous 
les autres caractères étant conformes à la diagnose du genre (Ehrmann, 2002). 
D'ailleurs il n'est pas impossible que la femelle inconnue soit pourvue de tels lobes 
plus ou moins réduits. 

REMERCIEMENTS 

Je remercie vivement les Dr Bernd Hauser et Peter Schwendinger, du MHNG, 
qui ont attiré mon attention sur ce remarquable spécimen et m'ont permis de le décrire. 

REFERENCES 

Anisyutkin, L.N. 1998. Notes on the genus Deroplatys Westwood, with description of a new 
species from Vietnam (Mantodea: Mantidae: Deroplatyinae). Zoosystema rossica 7: 
95-99, 21 fig. 

Ehrmann, R. 2002. Mantodea, Gottesanbeterinnen der Welt. Münster, Natur und Tier - Verlag 
GmbH. 519 pp., 56 fig., 32 pi. couleurs. 



Revue suisse de Zoologie 1 14 (3): 513-519; septembre 2007 



Further additions to the scorpion fauna of the Guayana region 
of South America 1 

Wilson R. LOURENÇO 

Département de Systématique et Evolution, USM 0602, Section Arthropodes 
(Arachnologie), Muséum national d'Histoire naturelle, CP 053, 61 rue Buffon 
75005 Paris, France. E-mail: arachne@mnhn.fr 



Further additions to the scorpion fauna of the Guayana region of South 
America. - This paper presents the results of a study of a new collection of 
South American scorpions now deposited in the Geneva Museum. The 
collection comprises two families, four genera and nine species. Among 
them is one new species, Broteochactas danielleae sp. n. (Chactidae), 
described here from Guyana and Brazil. 

Keywords: Scorpiones - Broteochactas - new species - Guyana. 

INTRODUCTION 

Contributions to our knowledge of the scorpion fauna of the Neotropical region 
and to material deposited in the Natural History Museum of Geneva have been 
presented in previous studies (Lourenço, 1997, 2002a). As already mentioned in 
numerous publications, South America is probably one of the regions of the world best 
studied for its fauna of scorpions. Studies began in the first half of the 19 th century and 
were for the first time compiled in a monograph by Mello-Leitäo (1945). Since then 
other contributions have been published, notably by Maury (1979), Gonzâlez-Sponga 
(1996), Lourenço, (2002b,c) and Ojanguren Affilastro (2005). On account of the 
diversity and richness of the scorpion fauna of South America, the discovery and 
description of new species and even of new genera is by no means unusual (Lourenço, 
2003, 2006; Lourenço et al, 2004). 

The present contribution, once again, is the result of a study of a more or less 
heterogeneous collection of scorpions now deposited in the Natural History Museum 
of Geneva. In addition to the description of a new species in the genus Broteochactas 
Pocock, a list of all the species studied here is given. This list is presented in an 
alphabetical order of families, genera and species. Species from the following countries 
have been included: Brazil, Guyana, Suriname and Venezuela. Most of the material 
examined was collected in the Guayana region, a floristic lowland province that has 
been delineated botanically (see Mori, 1991). 



Study supported by the "Département municipal des affaires culturelles de la Ville de Genève' 
Manuscript accepted 25.05.2007 



514 w - R- LOURENÇO 



METHODS 



Illustrations and measurements were made using a Wild M5 stereo-microscope 
with a drawing tube and an ocular micrometer. Measurements follow those of Stahnke 
(1970) and are given in mm. Trichobothrial notations are those developed by Vachon 
(1974) and the morphological terminology mostly follows that of Vachon (1952) and 
Hjelle (1990). 

TAXONOMY 

Family Buthidae C.L. Koch, 1837 

Genus Tityus C.L. Koch, 1836 
Subgenus Atreus Gervais, 1843 

Tityus cambridgei Pocock, 1 897 

Material: Suriname, between Moengo and Albina, 5-15/III/1987 (collected by local 
village people and received by J.-C. Lacroix), 2 males, 2 females. 

Distribution: Eastern Amazonia and Guayana, Brazil, French Guiana and 
Suriname. 

Tityus discrepans (Karsch, 1879) 

Material: Venezuela, Caracas IV/2005 (A. Borges leg.), 2 males, 2 females. 
Distribution: Trinidad, Venezuela. 

Tityus falconensis Gonzâlez-Sponga, 1974 

Material: Venezuela, Estado Falcon, 11/2005 (E. Ythier leg.), 1 female. 
Distribution: Venezuela. 

Tityus magnimanus Pocock, 1 897 

Material: Venezuela, Estado Mènda, X/1977 (P. Augusto leg.), 1 female. 
Distribution: Venezuela. 



Family Chactidae Pocock, 1893 
Genus Broteochactas Pocock, 1893 

Broteochactas danielleae sp. n. Figs 1-12 

Material: Guyana (on the border with Brazil), between Lethem and Conceiçâo do Mau, 
6/IV/1987 (collected by local people and received by J.-C. Lacroix): female holotype. Brazil, 
State of Para, on the border with Guyana, X/1965 (F. Castro leg.); female paratype. Type material 
deposited in the Natural History Museum of Geneva (MHNG). 

Etymology: The species name is a patronym in honor of Dr Danielle Decrouez, 
director of the Natural History Museum of Geneva. 

Diagnosis: Small scorpions, 23 to 24 mm in total length. Coloration reddish 
yellow to reddish brown. Body and appendages weakly granulated or smooth, with 
minute punctation. Pectines with 7-8 teeth in females. Trichobothrial pattern of type C 
neobothriotaxic 'majorante'. 



FURTHER ADDITIONS TO THE SCORPION FAUNA OF THE GUAYANA 



515 



1 & 






2 










M# 
















JH 


■Vy 



Figs 1-2 
Broteochactas danielleae sp. n. Female holotype, dorsal and ventral aspects. 



Broteochactas danielleae sp. n. can be included in the 'Broteochactas' species 
group (Lourenço, 2002b). The new species can be distinguished from others in the 
genus Broteochactas and in particular from Broteochactas santanai Gonzalez-Sponga, 
1978 which occurs in the nearby region of Estado Bolivar in Venezuela, and from 
Broteochactas delicatus (Karsch, 1879) which is distributed throughout the Guayana 
region, by the following features: (i) carapace, pedipalps and metasomal tegument 
exclusively punctuated (ii) overall size smaller (iii) larger number of pectinial teeth. 
Moreover, the new species is found in a savannah-like open vegetation habitat, 
the 'Amazon terra firme savannahs' (Murça Pires & Prance, 1985), whereas the other 
two species are found in tropical forest. 

Description: (based on female holotype and paratype). Coloration. Basically 
reddish yellow to reddish brown (Figs 1-2). Prosoma: carapace reddish brown. Tergites 
reddish brown, slightly paler than carapace, with two longitudinal yellowish strips. 
Metasomal segments reddish yellow, with darker zones over carinae; vesicle yellowish. 
Chelicerae reddish yellow, with diffuse variegated blackish spots; fingers uniformly 
deep reddish yellow. Pedipalps reddish; femur darker than patella and chela. Legs red- 
dish yellow to yellow. Venter and sternites yellowish; pectines and genital operculum 
paler than sternites. 

Morphology: Carapace lustrous and acarinate, with dense minute punctation; 
furrows shallow. Sternum pentagonal, wider than long. Tergites acarinate, with only 
minute granulations, almost smooth and shiny. Pectinal teeth count 7-7 (7-8), fulcra 
absent (Fig 2). Sternites smooth and shiny, VII acarinate. Metasomal segments IV and 
V longer than wide; metasomal tegument almost lustrous with only a few small granu- 
lations and a few punctuations; segment V with small spinoid granulations ventrally 



516 



W. R. LOURENÇO 




8 



11 



Figs 3-11 
Broteochactas danielleae sp. n. Female holotype. (3) Chelicera, dorsal aspect. (4) Disposition of 
granules on the dentate margins of the pedipalpal chela movable finger. (5) Metasomal segment 
V and telson, lateral aspect. (6-11) Trichobothrial pattern. Chela, dorso-external and ventral 
aspects (6-7); femur, dorsal aspect (8); patella, dorsal, external and ventral aspects (9-11). 



FURTHER ADDITIONS TO THE SCORPION FAUNA OF THE GUAYANA 



517 




Fig. 12 
Map of the Guianas region, comprising the Guayana floristic province. Are indicated the loca- 
lities of Broteochactas danielleae sp. n., in Guyana and Brazil (black circle with white star), and 
the new localities in Suriname for Tityus cambridgei, Broteochactas delicatus, Brotheas 
gervaisii, Brotheas granulatus and Hadrurochactas schaumii (black circle). 



(Fig 5). Carinae on segments I-V moderately developed; ventral carina absent from 
segments I and II, weakly marked on segments III-IV. Pedipalps: femur with dorsal 
internal, dorsal external and ventral internal carinae moderately marked; ventral 
external carina vestigial; dorsal and ventral aspects with minute granulations; internal 
aspect weakly granular. Patella with minute granulations; dorsal internal, ventral 
internal, ventral external and external carinae weak; other carinae vestigial. Chela with 
minute granulations; ventral and dorsal median carina moderately to strongly 
developed; other carinae weakly marked; internal aspect with a few moderate granules. 
Dentate margins on movable and fixed fingers with 6 rows of granules (Fig 4). 
Chelicerae with a dentition typical of the family Chactidae (Vachon, 1963), and with 
dense setation ventrally and internally (Fig 3). Trichobothriotaxy of type C (Figs 6-11); 
neobothriotaxic 'majorante' (Vachon, 1974). 

Morphometric values of the female holotype: Total length excluding the vesicle, 
24.1. Carapace: length, 4.2; anterior width, 2.8; posterior width, 4.7. Metasomal 
segments. I: length, 1.4; width, 2.5; II: length, 1.7; width, 2.1; III: length, 1.7; width, 



518 W. R. LOURENÇO 

1.9; IV: length, 2.1; width, 1.8; V: length, 3.8; width, 1.7; depth, 1.5. Vesicle: width, 
1.5; depth, 1.1. Pedipalp: femur length, 2.9, width, 1.3; patella length, 3.5, width, 1.5; 
chela length, 6.8, width, 2.5, depth, 3.0; movable finger length, 3.8. 

Broteochactas delicatus (Karsch, 1 879) 

Material: Suriname, between Moengo and Albina, 5-15/III/1987 (collected by local 
village people and received by J.-C. Lacroix), 9 males, 3 females. 

Distribution: Brazil, French Guiana, Guiana, Suriname, Venezuela (?). 

Genus Brotheas C.L. Koch, 1837 

Brotheas gervaisii Pocock, 1 893 

Material: Suriname, between Moengo and Albina, 5-15/III/1987 (collected by local 
village people and received by J.-C. Lacroix), 2 males. 

Distribution: Brazil, French Guiana, Suriname. 

Brotheas granulatus Simon, 1877 

Material: Suriname, between Moengo and Albina, 5-15/111/1987 (collected by local 
village people and received by J.-C. Lacroix), 1 male, 3 females. 

Distribution: Brazil, French Guiana, Suriname. 

Genre Hadrurochactas Pocock, 1 893 

Hadrurochactas schaumii (Karsch, 1880) 

Material: Suriname, north of Benzdorp, 18/25/111/ 1987 (collected by local village 
people and received by J.-C. Lacroix), 2 males, 2 females. 

Distribution: Brazil, French Guiana, Guiana, Suriname, Venezuela. 

ACKNOWLEDGEMENTS 

I am very grateful to Dr Danielle Decrouez, director of the Natural History 
Museum of Geneva, and to the 'Département municipal des affaires culturelles de la 
Ville de Genève' for supporting this study the results of which are presented here. I 
thank Prof. John L. Cloudsley-Thompson (London) and Dr Peter Schwendinger 
(MHNG), for reviewing the manuscript. Claude Ratton (MHNG) produced the photo- 
graphs and Florence Marteau (MHNG) arranged them into a plate. 

REFERENCES 

Gonzàles-Sponga, M. A. 1996. Gufa para identificar escorpiones de Venezuela. Cuadernos 

Lagoven, Caracas, 204 pp. 
Hjelle, J. T. 1990. Anatomy and morphology (pp. 9-63). In: Polis, G. A. (ed.). The Biology of 

Scorpions. Stanford University- Press, Stanford, 587 pp. 
Lourenço, W. R. 1997. Additions à la faune de scorpions néotropicaux (Arachnida). Revue 

suisse de Zoologie 104 (3): 587-604. 
Lourenço, W. R. 2002a. Nouvelles additions à la faune de scorpions néotropicaux (Arachnida). 

Revue suisse de Zoologie 109 (1): 127-141. 
Lourenço, W. R. 2002b. Scorpions of Brazil. Les Editions de L'IF, Paris, 320 pp. 



FURTHER ADDITIONS TO THE SCORPION FAUNA OF THE GUAYANA 519 

Lourenço, W. R. 2002c. Scorpions (pp. 399-438), In: J. Adis (ed.) Amazonian Arachnida and 
Myriapoda. Pensoft Publishers, Series Faunistica N° 24, Sofia-Moscow, 590 pp. 

Lourenço, W. R. 2003. Humicolous buthoid scorpions: a new genus and species from French 
Guiana. Comptes rendus Biologies 326: 1149-1 155. 

Lourenço, W. R. 2006. Nouvelle proposition de découpage sous-générique du genre Tityus 
C. L. Koch, 1836 (Scorpiones, Buthidae). Boletin de la Sociedad Entomològica Arago- 
nesa 39: 55-67. 

Lourenço, W. R. Cerqueira Baptista, R. L. & Ponce de Leao Giupponi, A. 2004. Troglobitic 
scorpions: a new genus and species from Brazil. Comptes rendus Biologies 327: 
1151-1156. 

Maury, E. A. 1979. Apuntes para une zoogeografia de la escorpiofauna argentina. Acta 
Zoologica Lilloana 35: 703-719. 

Mello Leitäo, C. 1945. Escorpiòes Sul Americanos. Arquivos do Museu Nacional, Rio de 
Janeiro 40: 1-468. 

Mori, S. A. 1991. The Guayana lowland floristic Province. Comptes rendus des séances de la 

Société de Biogéographie 67 (2): 67-75. 
Murça Pires, J. & Prance, G. T. 1985. The vegetation types of the Brazilian Amazon (pp. 109- 

145). In: Prance, G. T. & Lovejoy, T. E. (eds). Amazonia. Pergamon Press, Oxford, New 

York, 442 pp. 

Ojanguren Affilastro, A. A. 2005. Estudio monografico de los escorpiones de la Repûblica 

Argentina. Revista Ibèrica de Aracnologfa 11: 75-241. 
Stahnke, H. L. 1970. Scorpion nomenclature and mensuration. Entomological News 81: 

297-316. 
Vachon, M. 1952. Etudes sur les Scorpions. Institut Pasteur d'Algérie, Alger, 482 pp. 
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, 2e sér., 35 

(2): 161-166. 
Vachon, M. 1974. Etude des caractères utilisés pour classer les familles et les genres de 

Scorpions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles trichobothriaux 

et types de trichobothriotaxie chez les Scorpions. Bulletin du Muséum national 

d'Histoire naturelle, Paris 3e sér., n° 140, Zool. 104: 857-958. 



Revue suisse de Zoologie 1 14 (3): 521-572; septembre 2007 



Records of Pauropoda (Pauropodidae; Brachypauropodidae; 
Eurypauropodidae) from Singapore, Indonesia and Malaysia with 
the description of 18 new species 
(Pauropoda and Symphyla of the Geneva Museum XV) 

Ulf SCHELLER 

Häggeboholm, Häggesled, S-53194 Järpas, Sweden. 

Records of Pauropoda (Pauropodidae; Brachypauropodidae; Eury- 
pauropodidae) from Singapore, Indonesia and Malaysia with the 
description of 18 new species (Pauropoda and Symphyla of the Geneva 
Museum XV). - A collection of 104 Pauropoda (Myriapoda) from South- 
east Asia was studied. Twenty-seven species were identified, 1 8 of them are 
new to science and are described here. These are: Allopauropus serapiensis 
sp. n., A. cibodasensis sp. n., A.javanus sp. n., A. bidentulus sp. n., A. arcua- 
tus sp. n., A. acuticaudis sp. n., A. prolongatus sp. n., A. divaricatus sp. n., 
A. bakoensis sp .n., A. crassus sp. n., A. curtus sp. n., A. quadrispinus sp. n., 
A. trapezoides sp. n., Scleropauropus singapuranus sp. n., Samarangopus 
trilix sp. n., S. tuberosus sp. n., S. interstinguus sp. n., S. sarawakensis sp. 
n. Most species in this collection have not been found elsewhere, indicating 
a high degree of endemism. The wide range element is poor in species. 

Keywords: Myriapoda - taxonomy - soil fauna - Southeast Asia - bio- 
geography. 

INTRODUCTION 

In November and December 1987 an entomological expedition from the Natural 
History Museum, Geneva, visited Singapore, Indonesia (Java, Bali) and Malaysia 
(Sarawak). The members, Drs Bernd Hauser, Charles Lienhard and Pierre Strinati, col- 
lected 104 specimens of Pauropoda from litter and subsoil, mainly by means of auto- 
matical extraction and to a less degree manually. Twenty-seven species are present be- 
longing to six genera in three families: Pauropodidae {Allopauropus, Rabaudauropus 
and Scleropauropus), Brachypauropodidae (Brachypauropoides) and 
Eurypauropodidae (Samarangopus and Sphaeropauropus). This material made it 
possible to report Pauropoda for the first time from Singapore, Bali and Sarawak. Two 
species are now known from Singapore {Scleropauropus singapuranus sp. n. and 
Samarangopus tuberosus sp. n.), one species from Bali {Allopauropus quadrispinus sp. 
n.) and 18 species from Sarawak, 13 of them new to science (Allopauropus serapiensis 
sp. n., A. bidentulus sp. n., A. arcuatus sp. n., A. acuticaudis sp. n., A. prolongatus sp. 
n., A. divaricatus sp. n., A. bakoensis sp. n., A. curtus sp. n., A. crassatus sp. n., A. 
trapezoides sp. n., Samarangopus trilix sp. n., S. interstinguus sp. n., S. sarawakensis 
sp. n.) and five species previously known from other countries (Allopauropus man- 



Manuscript accepted 24.05.2007 



522 U. SCHELLER 

jakotompensis Remy, A. proximus Remy, Rabaudauropus dispar Scheller, 
Brachypauropoides penanorum Scheller, Samarangopus longipenes Scheller). Six 
additional species can be listed for the Javanese fauna {Allopauropus mortensenii 
(Hansen), A. pulcher (Remy), A. pumilio Remy, A. javanus sp. n., A. cibodasensis 
sp. n. and Sphaewpauropus arcuatus Scheller). 

MATERIAL AND METHODS 

Abbreviations: ad. ..., subad. ... and juv. ... = an adult, a subadult or a juvenile 
specimen with the number of pairs of legs indicated. Descriptive terms are listed in 
Scheller, 1988. 

Measurements: body length of holotype and range of body lengths of adult 
paratypes are given in mm in brackets. Other indications of absolute lengths are given 
in urn. When relative lengths are used the reference value is stated in the text. 

Collecting has been made either by hand or by soil samples, the latter extracted 
in Berlese funnels at places indicated. 

The material, preserved in alcohol, is deposited in the pauropod collection of the 
Department of Arthropods and Entomology I, Natural History Museum of Geneva, 
Switzerland. 

In the section Systematics the collectors are given by surnames only. 

PREVIOUS KNOWLEDGE 

The Pauropoda fauna of the south-eastern Oriental region is poorly known by a 
few records only: Hansen (1902), a small collection of nine species from the island of 
Koh Chang in the Gulf of Siam; Silvestri (1930), two species from Java; Remy (1933), 
one species from central Vietnam; Attems (1938) and Scheller (2004), one and eight 
species respectively from Vietnam; Remy (1957d), six species from the Philippines, the 
Palau Islands and Guam; Scheller et al. (1994) and Scheller (2001), six and 14 species 
respectively from Sabah on northern Borneo and Scheller (1995), 11 species from 
north-western Thailand. 

The picture of a poorly investigated Pauropoda fauna in this part of the world is 
strengthened by the almost complete lack of knowledge about the northern part of the 
Australian region. Only three publications deal with that region: Remy (1957b) and 
Scheller (1993), one and 22 species, respectively, from New Caledonia, and Scheller 
(1996) one species from Papua New Guinea. 

With few exceptions the collections studied by the above authors are small, but 
taken together they indicate a most diversified fauna. 

SYSTEMATICS 

Order Tetramerocerata 

Pauropodidae 

Pauropodinae 

Genus Allopauropus Silvestri, 1902 

Subgenus Decapauropus Remy, 1957 (Remy. 1957a). 



RECORDS OF PAUROPODA 523 



1 . Alllopauropus (D.) proximus Remy 

Allopauropus proximus Remy, 1948a: 572-573, fig. 4. 

Material examined: Malaysia, Sarawak, Serian District, Penrissen Road, 12 mis from 
Kuching, "Semongok Wildlife Rehabilitation Centre, Nursery Centre of the Forest Department", 
soil sample from between buttresses of large trees (extraction at Kuching, Sarawak), alt. 50 m, 4 
ad. 9(9), 8.XII.1987 (loc. Sar-87/60, leg. Hauser); Bako National Park, Jalan Lintang, soil sam- 
ple (extraction in Geneva) from between buttresses of Austrobuxus nitidus Miq. [= Longetia 
malayana (Benth.) P. & H.] (Euphorbiaceae), alt. 30 m, 2 ad. 9( 9 ), 2 juv. 6, 2 juv. 5, 1 1. XII. 1987 
(loc. Sar-87/76, leg. Hauser). - Alltogether 10 specimens. 

General distribution: This species is widely and discontinuously distributed 
in the tropics and subtropics of the Americas, Africa and southern Asia. It is here 
reported for the first time from Sarawak. 

2. Allopauropus (D.) mortensenii (Hansen) 

Pauropus mortensenii Hansen, 1902: 382-385, pi. 5, figs la-c. 

Material examined: Indonesia, Java, Bogor, Botanical Garden, soil sample from 
between buttresses of large trees near "Guest House" (extraction at Bogor), alt. =250 m, 2 ad. 
9(9), 24.XI.1987 (loc. Sar-87/8, leg. Hauser). - Altogether 2 specimens. 

General distribution: This species is here reported for the first time from 
Java. It was described from the island of Koh Chang in the Gulf of Thailand (Hansen, 
1902) and has later been reported also from Egypt (Remy, 1950), Réunion (s. n. 
Allopauropus cf. mortensenii, Remy, 1956e), Mauritius (s. n. Allopauropus cf. 
mortensenii, Remy, 1959), Sri Lanka (Remy, 1962; Scheller, 1970). There is also a 
doubtful record from Australia. 

3. Allopauropus (D.) manjakotompensis Remy & Bello 

Allopauropus manjakotompensis Remy & Bello, 1960: 86-88, fig. 8. 

Material examined: Malaysia, Sarawak, Bako National Park, Jalan Tg. Sapi, soil sam- 
ple from between buttresses of large trees (extraction in Geneva), alt. 10 m, 1 juv. 6, 2 juv. 5, 
11.XTI.1987 (loc. Sar-87/80, leg. Hauser). - Altogether 3 specimens. 

General distribution: This species is previously known from two localities on 
Madagascar only (Remy & Bello, 1960). 

4. Allopauropus (D.) pulcher Remy 

Allopauropus pulcher Remy, 1956c: 445-446, figs 1-3. 

Material examined: Indonesia, Java, Cibodas, Botanical Garden, alt. 1300 m, 1 ad. 
9(9), 25.XI.1987 (loc. Sar-87/18, leg. Hauser). 

General distribution: This species is here reported for the first time from 
Java. It was previously known only from the type locality in South Africa and from a 
few places in the USA: Texas (Remy, 1956d) and Great Smoky Mountains National 
Park (Scheller & Bernard, 2005). 

5. Allopauropus (D.) pumilio Remy 

Allopauropus pumilio Remy, 1956e: 148-149, fig. 7. 

Material examined: Indonesia, Java, Bogor, Botanical Garden, soil sample from 
between buttresses of large trees near the two lakes (extraction at Bogor), alt. 260 m, 2 ad. 9( 9 ), 
24.XI.1987 (loc. Sar-87/11, leg. Hauser). - Altogether 2 specimens. 



524 U. SCHELLER 

General distribution: The species is here reported for the first time from Java. 
It was known earlier from the type locality on La Réunion only. 

6. Allopauropus (D.) serapiensis sp. n. Figs 1-11 

Type material: Holotype: ad. 9(9), Malaysia, Sarawak, Gunung Serapi at road 
Kuching-Matang, soil sample in forest at the road to the television station (extraction at Kuching, 
Sarawak), alt. 670 m, 9.XII.1987 (loc. Sar-87/64, leg. Hauser). Paratype: same data as for holo- 
type, 1 ad. 9( 9 ). 

Other material: Malaysia, Sarawak, Bako National Park, Jalan Lintang, soil sample 
from between buttresses of Austrobuxus nitidus Miq. [= Longetia malayana (Benth.) P. & H.] 
(Euphorbiaceae) (extraction in Geneva), alt. 30 m, 4 ad. 9(2o\2$), 1 subad. 8(sex?), 2 juv. 6, 
2 juv. 5, 11. XII. 1987 (loc. Sar-87/76, leg. Hauser). - Altogether 11 specimens. 

Etymology: A latinized adjective of the name Serapi. 

Diagnosis: Several pygidial characters (shape of anal plate, setae of tergum and 
sternum, shape of posterior margin of tergum) indicate that A. serapiensis sp. n. is close 
to A. (D.) absimilis Serieller from southern Vietnam (Serieller, 2004). Good distinctive 
characters are the shape of the antennal globulus (proportionately long, with ovoid 
capsule in A. serapiensis sp. n.; short, with almost spherical capsule in A. absimilis), 
the proportions of the tergal antennal branch (5.3 times as long as its greatest diameter 
and 1.8 times as long as sternal branch in A. serapiensis sp. n.; 2.2-2.6 and 0.9-1.1 
respectively in A. absimilis) and the shape of the tarsus of the last pair of legs (2.7-3.2 
times as long as its greatest diameter in A. serapiensis sp. n., 4.5 in A. absimilis). 

Description: Length = 0.41 (-0.43) mm. 

Head (Fig. 1): Most tergal setae lost, those present short, very thin, tapering, 
pointed, striate. Relative lengths of setae (holotype only), 1 st row: flj = 10, a 2 = ?» 2 nd 
row: «!= 13, a 2 and a 3 = ?; 3 rd row: a x = 10, a 2 = ?; 4 th row: a x - a 2 - 10, a 3 = ?, a 4 
= 12; lateral group: l x = ?, l 2 = 8, l 3 = 15. Ratio a l /a l - a l in 1 st and 2 nd row 0.6, 3 rd 
row 0.8, 4 th row 0.7. Temporal organs large, at least twice longer than their shortest 
interdistance; small pistil in posterior part. Head cuticle glabrous. 

Antennae (Fig. 2): Segment 4 with setae p, p\ p" and r; /?'" not ascertained. 
Setae cylindrical, densely striate. Relative lengths of setae: p - 100, p' = (57-)60, p" 
= (28-)30(-32), r = 60(-76). Tergal seta p about as long as tergal branch t. The latter 
somewhat fusiform, (2.2-)2.6 times as long as its greatest diameter and about as long 
as sternal branch s, this (1.5-)1.7 times as long as its greatest diameter and with its 
anterodistal corner distinctly truncate. Seta q cylindrical, blunt, densely striate, almost 
as long as (-1.3 times as long as) length of s. Relative lengths of flagella (basal 
segments included) and basal segments: F t = 100, bs^ = (4-)5; F 2 = (24-)33(-37), bs 2 
= 3(-4); F 3 = 70(-78), bs 3 = 5(-6). F 1 4.9(-5.8) times as long as t, F 2 and F 3 (1.6-)1.8(- 
2.0) and (3.5-)4.5 times as long as 5, respectively. Distal calyces with small caps and 
distal part of flagella axes fusiformly widened. Globulus g 1.6(-1.8) times as long as 
wide and its width (0.9-)l.l of greatest diameter of t; =8 bracts present. Antennae 
glabrous. 

Trunk: Setae of collum segment (Fig. 3) simple, somewhat davate, blunt, 
annulate. Sublateral seta (1.6-)1.9 times as long as submedian seta; sternite process 
narrow anteriorly and without apical incision; appendages barrel-shaped, caps with 
distinctly bent surface. Process and appendages glabrous. 



RECORDS OF PAUROPODA 



525 




,a 


20 Mm 

j 


,b 


« 



Figs 1-11 
Allopauropus (D.) serapiensis sp. n., 1-5, 7-11, holotype, ad. 9(9); 6, paratype, ad. 9(a). (1) 
head, median and right part, tergal view; (2) left antenna, sternal view; (3) collum segment, 
median and left part, sternal view; (4) tergite VI, posterior part; (5) T^; (6) genital papillae, 
anterior view; (7) seta on coxa of leg 9; (8) seta on trochanter of leg 9; (9) tarsus of leg 9; (10) 
tarsus of leg 8; (1 1) pygidium, tergal view. Scale a: Figs 1, 5; b: Figs 3-4, 6-10; c: Figs 2, 11. 

Setae on tergites as submedian setae on head; 4+4 setae on tergite I, 6+6 on 
II-IV, 6+? on V and 4+2 on VI. Posterior setae on tergite VI (Fig. 4) 0.2(-0.3) of inter- 
distance and about 0.5 of length of pygidial setae a v 



526 ü. SCHELLER 

Bothriotricha (Fig. 5): Relative lengths: T { = 100, T 2 = ?(125), T 3 = (101-)114, 
T 4 = (108-)130, T 5 = (146-)182). All with straight and very thin axis. T 3 thickest and 
with apical ovoid swelling, 0.1 of length of bothriotrix. Pubescence very short except 
below end-swelling of T 3 , the latter with very short, simple hairs. 

Genital papillae (Fig. 6, paratype): 2.1 times as long as their greatest diameter, 
proximal half subcylindrical, distal half conical, with outer side strongly curved, 
glabrous; distal seta short, very thin, 0.4 of length of papilla. 

Legs: Setae on coxa (Fig. 7) and trochanter (Fig. 8) of leg 9 short, furcate, 
branches thin, cylindrical, striate, secondary branch of seta on trochanter very thin. 
Corresponding setae on more anterior legs (including setae on coxa of leg 2 in male), 
simple, cylindrical, without rudiments of secondary branch. Tarsus of leg 9 (Fig. 9) 
tapering, (2.7-)3.1(-3.2) times as long as its greatest diameter. Setae striate-annulate, 
proximal seta short, thin, tapering, curved, V4 of length of tarsus and very little longer 
than distal seta; the latter cylindrical, striate, blunt, almost as long as proximal seta. 
Cuticle of tarsus glabrous. Distal setae on tarsi of legs 1-8 furcate (Fig. 10), branches 
cylindrical, striate, blunt. 

Pygidium (Fig. 11): Tergum. Posterior margin between st and above anal plate 
with a low triangular bulge protruding from a trapezoid base. Relative lengths of setae: 
flj = 10, a 2 = (7-)8, a 3 = (12-)13(-14), st = 7(-8). Setae a^, a 2 , and a 3 thin, cylindrical, 
somewhat curved inwards and converging, only a 3 with faint pubescence distally; st 
tapering, pointed, somewhat curved inwards and with a knee in proximal third. 
Distance a 1 - a x 1.3(-1.5) times as long as a^, distance a x - a 2 as long as distance a x - 
ûj and somewhat shorter than distance a 2 - ay, distance st - st 2.2(-2.6) times as long 
as st and 1.2(-1.3) times as long as distance öj - a x . Tergum glabrous. 

Sternum. Posterior margin between b\ straight. Relative lengths of setae 
(flj=10): by = 40(-46), b 2 - 10(-11). Setae tapering, striate, b 2 diverging, somewhat 
curved inwards, by 1.4(-1.5) times as long as interdistance; b 2 0.6 of distance b x - b 2 . 

Anal plate large, subquadrate, somewhat longer than broad, with convex lateral 
margins and broadly V-shaped posteromedian indentation. Each posterolateral corner 
with two appendages: one directed posteriorly, as long as plate, tapering, the other very 
short and protruding from sternal side. Long appendages somewhat curved inward, 
short ones straight, diverging. Plate and sternum glabrous. 

7. Allopauropus (D.) cibodasensis sp. n. Figs 12-19 

Type material: Holotype: ad. 9( 9 ), Indonesia, Java, Cibodas, Botanical Garden, alt. 
«1300 m, 25.XI.1987 (loc. Sar-87/18, leg. Hauser). 

Etymology: A latinized adjective that refers to the name Cibodas. 

Diagnosis: A. (D.) cibodasensis sp. n. seems to be a very close relative of 
A. (D.) bedosae Serieller from north-western Thailand (Serieller, 1995). They can be 
distinguished by the shape of the posterior part of the pygidial sternum (margin straight 
in A. cibodasensis sp. n., with broad indentation in A. bedosae) and by the shape of the 
anal plate (plate short with long appendages in A. cibodasensis sp. n., plate longer with 
short appendages, especially the sternal ones, in A. bedosae). 

Description: Length = 0.51 mm. 



RECORDS OF PAUROPODA 



527 




Figs 12-19 
Allopauropus (D.) cibodasensis sp. n., holotype, ad. 9(9). (12) head, median and right part, 
tergal view; (13) right antenna, sternal view; (14) collum segment, median and left part, sternal 
view; (15) tergite VI, posterior part; (16) T 3 ; (17) seta on trochanter of leg 9; (18) tarsus of leg 
9; (19) pygidium, tergal view. Scale a: Fig. 16; b: Figs 14, 17-18; c: Figs 12, 13, 15, 19. 



Head (Fig. 12): Some tergal setae lost, those present of medium length, some- 
what davate, with distinct endsegment, striate. a 3 in 2 nd row tapering, pointed. 
Relative lengths of setae: 1 st row: a x = a 2 = 10; 2 nd row: a x = 12, a 2 = 10, a 3 = 15; 3 rd 
row: a x and a 2 = ?; 4 th row: a Y = \3, a 2 = ?, a 3 = 25, a 4 = ?; lateral group: / } = 27, l 2 
= 20, l 3 = 22. Ratio a x /a x - a x in 1 st row 0.6, in 2 nd row 0.4 and 4 th row 0.8. Temporal 
organs 0.8 of shortest interdistance. Head cuticle glabrous. 



528 U. SCHELLER 

Antennae (Fig. 13): Segment 4 with setae p,p\ p " and r, p '" not ascertained. 
Setae cylindrical, annulate. Relative lengths of setae: p = 10Q,p'=10,p"=42,r = 53. 
Tergal seta/? 1.2 times as long as tergal branch t. The latter somewhat fusiform, widest 
near the middle, 2.4 times as long as its greatest diameter and as long as sternal branch 
5, this 1.7 times as long as its greatest diameter and with its anterodistal corner dis- 
tinctly truncate. Seta q cylindrical, blunt, striate, 0.9 of length of s. Relative lengths of 
flagella (basal segments included) and basal segments: Fj = 100, bs x = 6; F 2 = 45, bs 2 
= 3; F 3 = 81, bs^, = 7. F ] 5 times longer than t, F 2 and F 3 2.2 and 4.1 times as long as 
s, respectively. Distal calyces small, distal part of flagella axes fusiformly widened. 
Globulus g 1 . 1 times as long as wide and its width 0.9 of greatest diameter of t ; 9 bracts 
present. Antennae glabrous. 

Trunk: Setae of collum segment (Fig. 14) simple, somewhat davate, blunt, 
annulate. Sublateral seta 3.4 times as long as submedian seta; sternite process small, 
narrow anteriorly and without apical incision; appendages directed inwards, small, 
indistinct. Process and appendages glabrous. 

Setae on anterior tergites cylindrical, striate, more posteriorly growing glabrous, 
pointed; 4+4 setae on tergite I, 6+6 on II- V, and 4+2 on VI. Posterior setae on tergite 
VI (Fig. 15) 0.4 of interdistance and 0.9 of length of pygidial setae flj. 

Bothriotricha: Relative lengths: T x = 100, T 2 broken, T 3 = 95, T 4 = 113, T 5 = 
181. All with straight, thin axis, T 3 (Fig. 16) thickest, with apical swelling, this almost 
0.2 of length of bothriotrix and broadest near base. Pubescence short, longest below 
endswelling of r 3 , the latter with very short erect hairs. 

Legs: Setae on coxa of leg 9 not studied, seta on trochanter of that leg (Fig. 17) 
furcate, branches subsimilar, cylindrical, striate, blunt. Corresponding setae on more 
anterior legs simple, striate, blunt. Tarsus of leg 9 (Fig. 18) tapering, 2.7 times as long 
as its greatest diameter. Setae subequal in length, 0.3 of length of tarsus. Proximal seta 
tapering, pointed, with short pubescence distally; distal seta cylindrical, blunt, striate. 
Cuticle of tarsus glabrous. 

Pygidium (Fig. 19): Tergum. Posterior margin evenly rounded but with small 
median triangular lobe between aj and st. Relative lengths of setae: a x = 10, a 2 = 9, a 3 
= 24, st = 14. Setae a^ and a 2 straight, somewhat diverging, glabrous, the former cylin- 
drical, the latter tapering, a 3 curved inward, diverging, tapering, st straight, somewhat 
davate and converging, glabrous. Distance a.\ - a^ 1.2 times as long as a ls distance a^ 
- a 2 3.2 times as long as distance a 2 - a 3 ; distance st - st 1.2 times as long as st and 
1.4 times as long as distance a^ - a-y. Tergum glabrous. 

Sternum. Posterior margin between b\ straight. Relative lengths of setae (a^ = 
10): b { = 33, b 2 = 13. Setae b x cylindrical, faintly striate, b 2 somewhat diverging and 
curved inward, tapering, pointed. b x as long as interdistance; b 2 as long as distance b\ 
-b 2 . 

Anal plate subquadrate, with concave lateral margins; distal part with 4 
posteriorly directed cylindrical blunt appendages, tergal ones thickest, somewhat 
curved inward, those protruding from sternal side shorter and thinner, somewhat 
converging; tergal and sternal appendages 1.5 and 1.3 times as long as plate, respec- 
tively. Plate and sternum glabrous. 



RECORDS OF PAUROPODA 529 

8. Allopauropus (D.) javanus sp. n. Figs 20-28 

Type material: Holotype: ad. 9( 9 ), Indonesia: Java, Cibodas, Lithocarpus-Castanopsis 
forest along gorge above the Botanical Garden, under rotten wood, alt. =1400 m, 25. XI. 1987 
(loc. Sar-87/17, leg. Lienhard). 

Etymology: A latinized adjective of the name Java. 

Diagnosis: Among several similar species in the subgenus the new species can 
be identified by the following combination of characters: flattened base of capsule in 
antennal globulus, setae of collum segment somewhat claviform and with distinct 
endsegment, T 3 with distal endswelling with very short pubescence, anal plate with U- 
shaped posterior incision. 

A. javanus sp. n. might be most closely related to two species known from Sri 
Lanka, A. (D.) excavatus Serieller and A. (D.) baculatus Scheller (Scheller, 1970), and 
to the Madagascan A. (D.) delphini Remy and A. (D.) barroisi Remy (Remy, 1956a). 
Good distinctive characters are: in relation to A. excavatus the proportions of the tergal 
antennal branch (as long as sternal branch in A. javanus sp. n., distinctly longer in A. 
excavatus) and the number of appendages of the anal plate (only two sternal ones in A. 
javanus, two sternal and two tergal ones in A. excavatus); in relation to A. baculatus, 
the shape of the pubescence of the bothriotricha (very short in A. javanus sp. n., long 
and with partly branched hairs in A. baculatus) and the shape of the T 3 (with distal 
swelling in A. javanus sp. n., no swelling in A. baculatus); in relation to A. delphini, 
the shape of the pubescence on the endswelling of the 7 3 (endswelling almost glabrous 
in A. javanus sp. n., with long ramose hairs in A. delphini) and the shape of the 
posterior margin of the pygidial tergum (with triangular bulge in A. javanus sp. n., no 
bulge in A. delphini); in relation to A. barroisi: the shape of the st (long, cylindrical in 
A. javanus sp. n., short, davate in A. barroisi). Among these four species the new 
species may be most close to A. barroisi because both have very similar anal plates and 
the submedian setae a x in the 4 th row of the tergal setae on the head are lacking in both. 

Description. Length = 0.46 mm. 

Head (Fig. 20): Many tergal setae lost, those present short, somewhat davate, 
striate. fl 3 in 2 nd row cylindrical. Relative lengths of setae (for holotype only), 1 st row: 
a l =a 2 = 10; 2 nd row: a { = 11, a 2 = 10, a 3 = 13; 3 rd row: a x = ?, a 2 = 14; 4 th row: a x 
not identified, a 2 = 10, a 3 and a 4 = ?; lateral group, not identified. Ratio a x /a x - a x not 
studied. Temporal organs about as long as shortest interdistance. Head cuticle glabrous. 

Antennae (Fig. 21 ): Segment 4 with setae p, p', p" and r; p "' not ascertained. 
Setae/? and p' somewhat davate, annulate, p" cylindrical, striate. Relative lengths of 
setae: p = 100, p '■ = 53, p" = 32, r = ?. Tergal seta p 1 .5 times as long as tergal branch 
t. The latter somewhat fusiform, widest near the middle, 1.6 times as long as its greatest 
diameter and 1.1 times as long as sternal branch s, this 1.7 times as long as its greatest 
diameter and with its anterodistal corner moderately truncate. Seta q cylindrical, blunt, 
densely striate, 1.3 times as long as length of s. Relative lengths of flagella (basal 
segments included) and basal segments: F x - 100, bs x = 6; F 2 = 45, bs 2 = 5; F 3 = 90, 
bs 3 = 4. F i 4.5 times as long as t, F 2 and F 3 2.2 and 4.3 times as long as s, respectively. 
Distal calyces small, distal part of flagella axes fusiformly widened. Globulus g 1.2 
times as long as wide and its width 0.6 of greatest diameter of t; =9 bracts present. 
Antennae glabrous. 



530 



U. SCHELLER 




Figs 20-28 
Allopauropus (D.) javanus sp. n., holotype, ad. 9(9). (20) head, median and right part, tergal 
view; (21) right antenna, tergal view; (22) collum segment, median and left part, sternal view; 
(23) tergite VI, posteromedian part, tergal view; (24) T 3 ; (25) seta on coxa of leg 9; (26) seta on 
trochanter of leg 9; (27) tarsus of leg 9; (28) pygidium, posteromedian and left part, sternal view. 
Scale a: Fig. 24; b: Figs 20, 22, 25-27; c: Figs 21, 23, 28. 



Trunk: Setae of collum segment (Fig. 22) simple, somewhat davate, blunt, 
annulate. Sublateral seta 2.4 times as long as submedian seta; sternite process small, 
narrow anteriorly and without apical incision; appendages indistinct, caps thin. Process 
and appendages glabrous. 

Setae on tergites cylindrical, blunt, striate; 4+4 setae on tergite I and 4+2 on VI. 
Posterior setae on tergite VI (Fig. 23) 0.6 of interdistance and 0.8 of length of pygidial 
setae a^. 



RECORDS OF PAUROPODA 531 

Bothriotricha: Relative lengths: 7\ = 100, T 2 = 107, T 3 - 101, T 4 - 123, T 5 = 
179. All with straight and very thin axis. r 3 (Fig. 24) thickest, with apical ovoid 
swelling, this well 0.1 of length of bothriotrix. Pubescence very short except below 
endswelling of r 3 , the latter almost glabrous. 

Legs: Setae on coxa (Fig. 25) and trochanter (Fig. 26) of leg 9 furcate, branches 
cylindrical, striate, blunt; branches subequal in length on coxal seta, one of the 
branches distinctly longer than the other on trochanter. Corresponding setae on more 
anterior legs with somewhat davate main branch and rudimentary secondary branch. 
Tarsus of leg 9 (Fig. 27) tapering, 2.8 times as long as its greatest diameter. Setae equal 
in length, 0.3 of length of tarsus. Proximal seta tapering, pointed, with short pubescent 
distally; distal seta cylindrical, blunt, very densely striate. Cuticle of tarsus glabrous. 

Pygidium (Fig. 28): Tergum. Posterior margin between st and above anal plate 
with a triangular lobe. Relative lengths of setae: a x = 10, a 2 = st = 1 1, a 3 = 19. Setae 
ai straight, directed posteriorly, glabrous; a 2 , and a 3 curved inward, converging, the 
former glabrous, the latter with very faint distal pubescence; st somewhat S-shaped, 
converging, tapering, glabrous. Distance a^- a^ 1.2 times as long as a 1s distance a j - 
a 2 0.6 of distance ci\ - a x and 1.8 times as long as distance a 2 - fl 3 ; distance st - st 1.2 
times as long as st and 1.1 times as long as distance a± -ay Tergum glabrous. 

Sternum. Posterior margin between b\ with broad, shallow, bow-shaped inden- 
tation. Relative lengths of setae (öj = 10): b x = 41, b 2 = 12. Setae tapering, striate, b 2 
somewhat diverging and curved inwards. b\ 1.9 times as long as interdistance; b 2 0.7 
of distance b\ - b 2 . 

Anal plate 2.3 times as long as greatest breadth, lateral margins concave, distal 
part protruding into two rounded lobes separated by a U-shaped indentation, 0.3 of 
length of plate. Two davate striate appendages originating from sternal sides of these 
lobes and projecting backwards-downwards; these appendages 0.4 of length of plate. 
Plate and sternum glabrous. 

9. Allopauropus (D.) bidentulus sp. n. Figs 29-37 

Type material: Holotype: ad. 9(a), Malaysia: Sarawak, Gunung Serapi at road 
Kuching-Matang, soil sample (extraction at Kuching) from forest at the road to the television sta- 
tion, alt. 670 m, 9.XII.1987 (loc. Sar-87/64, leg. Hauser). Paratypes: same data as for the holo- 
type, 4 ad. 9(2 o\ 29). -Altogether 5 specimens. 

Etymology: From the Latin bi = two and dens = tooth (referring to the distal 
part of the anal plate). 

Diagnosis: A. (D.) bidentulus sp. n is a close relative of A. (D.)facetus Remy 
from Nosy Be, Madagascar (Remy, 1956a). The two species are very alike regarding 
the general chaetotaxy of the pygidium and the shape of the anal plate. They are well 
distinguished by the shape of the following characters: antennal globulus g (1.5 times 
as long as its greatest diameter and its length 0.6-0.7 of length of s in A. bidentulus 
sp. n., longish, 2.2 times as long as greatest diameter and its length as long as s in 
A. facetus); base segments of the antennal flagella (about twice longer than wide in 
A. bidentulus sp. n., 5-6 times longer than wide in A. facetus Remy); pygidial setae öj 
(cylindrical and converging in A. bidentulus sp. n., somewhat lanceolate and diverging 
in A. facetus). 



532 



U. SCHELLER 




Figs 29-37 
AUopaiiropus (D.) bidentulus sp. n.. holotype, ad. 9{6). (29) head and anterior part of tergite I, 
median and right part, tergal view; (30) left antenna, sternal view; (31) collum segment, median 
and left part, sternal view; (32) tergite VI, posterior part; (33) 7 3 ; (34) genital papillae, anterior 
view; (35) seta on coxa of leg 9; (36) tarsus of leg 9; (37) pygidium, median and left part, tergal 
view. Scale a: Fig. 33; b: Figs 29, 31-32, 34-36; c: Figs 30, 37. 



RECORDS OF PAUROPODA 533 

Description: Length = (0.28-)0.35 mm. 

Head (Fig. 29): Setae (holotype only) on anterior part of tergal side short, much 
longer on posterior part. Setae ajand a 2 of 4 th row not verified. Setae thin, striate. 
Relative lengths of setae: 1 st row: a x = a 2 = 10; 2 nd row: a x = a 2 = a 3 = 14; 3 rd row: a x 
= 26, a 2 = 36; 4 th row: a 3 = 28, a 4 = ?; lateral group setae not studied. Ratio a l /a i -a ] 
in 1 st row 1.0, in 2 nd row 0.7, in 3 rd row 1.6. Temporal organs proportionally large, 3.1 
times as long as their shortest interdistance. Small posterior aperture close to posterior 
margin of temporal organ. Head cuticle glabrous. 

Antennae (Fig. 30): Segment 4 with setae p,p\ p " and r; p '" not ascertained. 
Setae cylindrical, annulate. Relative lengths of setae (holotype only): p = 100, /?'=?, 
p" = 27, r = 46. Tergal seta/? 1.2 times as long as tergal branch t. The latter somewhat 
fusiform, widest near the middle, 1.5 times as long as its greatest diameter and 0.8 of 
length of sternal branch s, this (1.4-)1.6 times as long as its greatest diameter and with 
its anterodistal corner distinctly truncate. Seta q thin, cylindrical, blunt, faintly striate, 
about as long as s. Relative lengths of flagella (basal segments included) and basal 
segments: F 1 = 100, bs x = 5(-6); F 2 = 33(-38), bs 2 = 3; F 3 = 68(-76), bs 3 = (6-)7. F x 
(5.6-)6.7 times as long as t, F 2 and F 3 1.8(-1.9) and (3.5-)3.6 times as long as s, 
respectively. Distal calyces small, distal part of flagella axes somewhat widened in F 1 
and F 3 , not in F 2 . Globulus g 1.4 times as long as wide and its width 0.8 of greatest 
diameter of t; 9 bracts present. Antennae glabrous. 

Trunk: Setae of collum segment (Fig. 31) simple, cylindrical, blunt, striate. 
Sublateral seta 2.1 (-2.2) times as long as submedian seta; sternite process small, 
narrow anteriorly and without apical incision; appendages small, indistinct. Both 
process and appendages glabrous. 

Setae on anterior tergites cylindrical, striate, more posteriorly growing glabrous. 
4+4 setae on tergite I, 4+2 on VI. Posterior setae on tergite VI (Fig. 32) 0.4 of inter- 
distance and 0.8 of length of pygidial setae a x . 

Bothriotricha: Relative lengths (holotype only): T x = 100, T 2 = 88, T 3 = 80, T 4 = 
68, T 5 = 121. All but r 3 with straight, very thin axis, T 3 (Fig. 33), with apical swelling, 
this 0.2 of length of bothriotrix. Pubescence short, on 7\, T 2 , T4 and T 5 extremely short, 
longest pubescence below endswelling of T 3 , the latter with short almost erect hairs. 

Genital papillae (Fig. 34): Short, conical with rounded outer sides, 1.8 times as 
long as their greatest diameter, glabrous; distal seta very thin, 0.5 of length of papilla. 

Legs: Setae on coxa of leg 9 (Fig. 35) short, furcate, striate, branches subsimilar 
in length, seta on trochanter seems to be simple. Corresponding setae on more anterior 
legs simple, striate. Tarsus of leg 9 (Fig. 36) tapering, (3.1-)3.2 times as long as its 
greatest diameter. Proximal seta tapering, pointed, distal one cylindrical, blunt, striate, 
both 0.2 of length of tarsus. Cuticle of tarsus glabrous. 

Pygidium (Fig. 37): Tergum. Posterior margin evenly rounded. Relative lengths 
of setae: a x = 10, a 2 = 10(-11), a 3 = (15-) 16, st = 8. Setae cylindrical, somewhat curved 
inward, blunt, glabrous, converging. Distance a\ - öj 1.5(-1.6) times as long as a l5 
distance aj - a 2 about 3 times longer than distance a 2 - a 3 ; distance st - st (2.3-)2.6 
times as long as st and (1.1-)1.3 times as long as distance a x - a x . Tergum glabrous. 

Sternum: Posterior margin between b\ with deep, broadly V-shaped indentation. 
Relative lengths of setae (a l = 10): b x = 56(-58), b 2 = 14(-15). Seta b x cylindrical, 



534 U. SCHELLER 

somewhat tapering, faintly striate distally, b 2 subcylindrical, tapering distally, 
glabrous, somewhat diverging. b\ 1.4 times as long as interdistance, b 2 0.6 of distance 
b x - b 2 . 

Anal plate broadest at base, longish, triangular, with straight sides and two ap- 
pendages most distally; appendages very short, blunt, close to each other. Plate and 
sternum glabrous. 

10. Allopauropus (D.) arcuatus sp. n. Figs 38-45 

Type material: Holotype: ad. 9(2), Malaysia, Sarawak, Gunung Serapi at road 
Kuching-Matang, soil sample (extraction at Kuching) from forest at the road to the television sta- 
tion, alt. 670 m, 9.XH1987 (loc. Sar-87/64, leg. Hauser). 

Etymology: From the Latin arcus = bow (referring to pygidial setae st). 

Diagnosis: Many species in the subgenus have similar anal plates but A. (D.) 
arcuatus sp. n. is distinguished by having also davate bothriotricha F 3 and cylindrical 
st evenly curved inward like an arc of a circle. The setae a x of the 4 th row of setae on 
the tergal side of the head have not been verified. 

Description: Length = 0.32 mm. 

Head (Fig. 38): Tergal setae short, somewhat davate, annulate, with distinct 
endsegment. Relative lengths of setae: 1 st row: only one pair of setae verified 1 = 10; 
2 nd row: a x = 9, a 2 = 1 1, tf 3 = 15; 3 rd row: a x = 9, a 2 = ?; 4 th row: a x not verified, a 2 = 
?, a 3 = 9, a 4 = ?; lateral group of setae not studied. Ratio a^/a^ - flj in 2 nd row 0.3, in 
3 rd row 0.8. Temporal organs proportionally small, as long as shortest interdistance. 
Head cuticle glabrous. 

Antennae (Fig. 39): Segment 4 with at least setae p, p', and p", r and p'" not 
ascertained. Setae thin, cylindrical, indistinctly striate. Relative lengths of setae: p = 
100, p' - 62, p" - 55. Tergal seta p as long as tergal branch t. The latter somewhat 
fusiform, widest outside the middle, 1.7 times as long as its greatest diameter and 0.9 
of length of sternal branch s, this 1 .7 times as long as its greatest diameter and with its 
anterodistal corner distinctly truncate. Seta q thin, cylindrical, blunt, densely striate, 
1.5 times as long as s. Relative lengths of flagella (basal segments included) and basal 
segments: Fj = 100, bs\ = 5; F 2 = 32, bs 2 = 4; F 3 = 84, bs^ = 5. Fj 6.3 times as long 
as t, F 2 and F 3 1.9 and 4.8 times as long as s, respectively. Distal calyces hemi- 
spherical, distal part of flagella axes not widened. Globulus g 1.3 times as long as wide 
and its width 0.8 of greatest diameter of t; ~1 bracts present. Antennae glabrous. 

Trunk: Setae of collum segment (Fig. 40) simple, cylindrical, tapering, striate. 
Sublateral seta 3 times longer than submedian seta; sternite process small, narrow, with 
small incision anteriorly; appendages small. Process and appendages glabrous. 

Setae on anterior tergites cylindrical, striate, more posteriorly growing glabrous. 
4+4 setae on tergite I, 4+2 on VI. Posterior setae on tergite VI (Fig. 41) 0.5 of inter- 
distance and 0.8 of length of pygidial setae a x . 

Bothriotricha: 7\, T-, and F 4 lost or broken. Lengths: F 3 = 49, T 5 = 50 urn. All 
but F 3 seems to be very thin, F 3 (Fig. 42) with proximal 1/3 of axis cylindrical, middle 
1/3 forming a club-shaped swelling, distal 1/3 very thin; proximal part glabrous, 
middle part maybe with very short pubescence, distal part with long branched hairs 
arranged in whirls. 



RECORDS OF PAUROPODA 



535 



Ì •? f 




Figs 38-45 
Allopauropus (D.) arcuatus sp. n., holotype, ad. 9(9). (38) head, median and right part, tergal 
view; (39) left antenna, sternal view; (40) collum segment, median and left part, sternal view; 
(41) tergite VI, posterior part; (42) T 3 ; (43) seta on coxa of leg 9; (44) tarsus of leg 9; (45) 
pygidium, median and left part, sternal view. Scale a: Figs 38, 40; b: Figs 41-43; c: Figs 39, 
44, 45. 



536 U. SCHELLER 

Legs: Setae on coxa (Fig. 43) and trochanter of leg 9 simple, cylindrical, blunt, 
striate. Corresponding setae on more anterior legs simple, striate, coxal seta somewhat 
thicker than seta on trochanter. Tarsus of leg 9 (Fig. 44) tapering, 2.4 times as long as 
its greatest diameter. Proximal seta tapering, pointed, distal one cylindrical, blunt, 
striate, the former 0.4 and the latter 0.7 of length of tarsus. Cuticle of tarsus glabrous. 

Pygidium (Fig. 45): Tergum. Posterior margin evenly rounded. Relative lengths 
of setae: a^ = 10, a 2 = 7 ', a 3 = st = 13. Setae cylindrical, blunt, almost glabrous; a± and 
a 2 straight, a 3 and st curved inward, st curved like an arch, a 2 an d «3 diverging, st 
converging. Distance a x - a x 1.3 times as long as a 1? distance a^- a 2 \2> times as long 
as distance a 2 ~ a ^ distance st - st 1.6 times as long as st and distance a^ - a^. Tergum 
glabrous. 

Sternum. Posterior margin between b x with low median bulge. Relative lengths 
of setae (a-^ = 10): b\ = 34, b 2 = 8. Setae striate, by subcylindrical, b 2 tapering, some- 
what diverging. b\ 1.2 times as long as interdistance, b 2 0.8 of distance b^ - b 2 . 

Anal plate 1.4 times as long as broad, broadest in distal third, with concave 
lateral margins; distal part protruding into two rounded somewhat diverging lobes; two 
cylindrical appendages originating from sternal sides of these lobes and projecting 
backward; these appendages 0.3 of length of plate, cylindrical, blunt, striate, somewhat 
diverging. Plate and sternum glabrous. 

1 1. Allopauropus (D.) acuticaudis sp. n. Figs 46-52 

Type material: Holotype: subad. 8( 9 ), Malaysia, Sarawak, Bau, in the surroundings of 
the cave "Gua Puang" near Kampong Pelaman Sekiang, on steep slope of Gunung Jambusan, un- 
der bark, alt. 10-30 m, 4.XII.1987 (loc. Sar-87/52, leg. Lienhard). 

Etymology: From the Latin acutus = pointed and cauda = tail. 

Diagnosis: The specimen is not in the best condition (many setae lost) but it is 
well defined by the peculiar shape of the anal plate in combination with the 
endswelling of the T 3 and the unusual shape of the proximal seta of the tarsus of the 
last pair of legs. A. acuticaudis sp. n. appears to be most closely related to two 
Madagascan species, A. baudoti Remy and A. lienharti Remy (Remy, 1956a). It can be 
distinguished from A. lienharti by the shape of the r 3 (thick, simple, with distal 
swelling in A. acuticaudis sp. n.; thin, branched in A. lienharti) and the shape of the 
distal seta on the tarsi of the last pair of legs (simple in A. acuticaudis sp. n.; furcate in 
A. lienharti). Good distinctive characters in relation to A. baudoti are the size ratio q/s 
(0.6 in A acuticaudis sp. n.; 1.7 in A. baudoti), the shape of the r 3 (thick, simple, with 
distal swelling in A. acuticaudis sp. n.; thin, tapering in A. baudoti) and the shape of 
the proximal seta on the tarsi of the last pair of legs (very short, blunt, less than 0.1 of 
length of tarsus and about 0.3 of length of distal seta in A. acuticaudis sp. n.; 1/4 of 
length of tarsus and longer than distal seta in A. baudoti). The new species is also re- 
lated to A. (D.) digitiger Remy (Remy, 1956a), but to a less degree. 

Description: Length = 0.42 mm. 

Head: Not available for study. 

Antennae (Fig. 46): Segment 4 with at least setae p, p ', and p ", r and p '" not 
ascertained. Setae thin, cylindrical, indistinctly striate. Relative lengths of setae: p = 
10, p ' = 6, p " - 4. Tergal seta p 0.9 of length of tergal branch t. The latter cylindrical, 



RECORDS OF PAUROPODA 



537 




Figs 46-52 
Allopauropus (D.) acuticaudis sp. n., holotype, subad. 8(9). (46) left antenna, sternal view; (47) 
7/3; (48) seta on coxa of leg 8; (49) seta on trochanter of leg 8; (50) tarsus of leg 8; (51) pygi- 
dium, median and left part, sternal view; (52) anal plate, lateral view. Scale a: Figs 48-49; b: Figs 
47, 50-52; c: Fig. 46. 



1.5 times as long as its greatest diameter and 0.9 of length of sternal branch s, this 1.3 
times as long as its greatest diameter and with anterodistal corner distinctly truncate. 
Seta q proportionately short, thin, cylindrical, blunt, densely striate, 0.6 of length of s. 
Relative lengths of flagella (basal segments included) and basal segments: Fj = 100, 
bsi = 9; F 2 = 36, bs 2 = 5; F 3 = 83, bs^ = 13. Fj 5.3 times as long as t, F 2 and F 3 1.7 
and 3.9 times as long as s, respectively. Distal calyces small, hemispherical, distal part 
of flagella axes not widened. Globulus g 1 .4 times as long as wide and its width 0.9 of 
greatest diameter of t; «9 bracts present. Antennae glabrous. 

Trunk: Collum segment and setae on tergites not available for study. 

Bothriotricha: Only r 3 (Fig. 47) studied, the others broken. Length of r 3 = 
35 urn, axis thick, with distal swelling, pubescence very short. 

Legs: Setae on coxa of leg 8 (Fig. 48) short, furcate, striate, main branch some- 
what davate, striate, secondary branch much thinner and 0.3 of length of main branch; 
seta on trochanter (Fig. 49) simple, cylindrical, blunt, striate. All corresponding setae 
on more anterior legs simple, striate. Tarsus of leg 8 (Fig. 50) tapering, 3.3 times as 
long as its greatest diameter. Proximal seta very short, cylindrical, blunt, glabrous, 



538 u - SCHELLER 

distal one cylindrical, blunt, striate; proximal seta less than 0.1 of length of tarsus and 
0.3 of length of distal seta. Cuticle of tarsus glabrous. 

Pygidium (Fig. 51): Tergum. Posterior margin with broad and shallow indenta- 
tion between st. Lengths of setae: öj = ?, a 2 = 7, «3 = ?, st = 8 urn. a 2 and st subcylin- 
drical, blunt, glabrous, somewhat curved inward, converging. Distance st - st 2.1 times 
as long as st. Tergum glabrous. 

Sternum. Posterior margin between b\ almost straight, with one small bulge 
only at each side of anal plate. Lengths of setae: b\ = 23, b 2 = 8 urn. Seta b\ cylindrical, 
striate, b 2 tapering, faintly striate distally, somewhat curved inward. b x as long as 
interdistance, b 2 0.8 of distance b-y - b 2 . 

Anal plate (Figs 51-52) broadest in proximal half, subcircular, with triangular 
posterior prolongation; this with one marginal row of hairs on each side and with two 
short appendages protruding backward from sternal side; appendages subparalleli, 
cylindrical, blunt, glabrous. Plate with faint pubescence, sternum glabrous. 

12. Allopauropus (D.) prolongatus sp. n. Figs 53-57 

Type material: Holotype: ad. 9( 9 ), Malaysia, Sarawak, Serian District, Penrissen Road, 
12 mis from Kuching, "Semongok Wildlife Rehabilitation Centre, Nursery Centre of the Forest 
Department", soil sample (extraction at Kuching) from between buttresses of large trees, alt. 50 
m, 8.XII.1987 (loc. Sar-87/60, leg. Hauser). Paratype: 1 ad. 9(9), same data as holotype. - 
Altogether 2 specimens. 

Etymology: From the Latin pro- = in relation to, and longus = long (referring 
to the lengthened posterior part of pygidial tergum). 

Diagnosis: The type specimens are not in the best condition (many setae lost) 
but they are well defined by the shape of the anal plate (with balloon-like submedian 
appendages), the posterior part of the pygidial tergum (with a large posterior lobe) and 
the st (balloon-like). The general morphology of the anal plate is alike that in A. (D.) 
ludovicae Remy from Corsica (Remy, 1945) but otherwise they are not close to each 
other. At present the relationships of this species can not be established. 

Description: Length = 0.75 mm. 

Head: Head not available for study. 

Antennae (Fig. 53): All setae lost. Tergal antennal branch elongate, somewhat 
fusiform, distal end obliquely truncate, 3.0 times longer than wide and as long as 
sternal branch s, this 2.2 times as long as its greatest diameter and with its anterodistal 
corner roundly truncate. Relative lengths of flagella (basal segments included) and 
basal segments (for holotype only): Fj = 100, bs j = 8; F 2 = 53, bs 2 = 6; F 3 = ?, bs^ = 
8. F { 3.1 times as long as t, F 2 1.7 times as long as s. Distal calyces of F l conical, those 
of F 2 small, hemispherical; distal part of flagella axes fusiformly widened. Globulus g 
1.2 times as long as wide and its width 0.9 of greatest diameter of t; 13 bracts present. 
Antennae glabrous. 

Trunk: Setae of collum segment (Fig. 54) furcate; main branch subcylindrical, 
blunt, striate, secondary branch rudimentary, pointed. Setae long, sublateral one 1.3 
times as long as submedian seta; sternite process small, narrow anteriorly and without 
apical incision; appendages subglobular, caps very small. Process and appendages 
glabrous. Setae on tergites I- VI not studied. 



RECORDS OF PAUROPODA 



539 




Figs 53-57 
Allopauropus (D.) prolongatus sp. n., holotype, ad. 9(2). (53) left antenna, sternal view; 
(54) collum segment, median and left part, sternal view; (55) seta on trochanter of leg 9; (56) 
tarsus of leg 9; (57) pygidium, median and left part, tergal view. Scale a: Figs 55-56; b: Fig. 54; 
c: Figs 53, 57. 

Bothriotricha: Not available for study. 

Legs: Setae on coxa and trochanter (Fig. 55) of leg 9 furcate, branches subequal 
in length, cylindrical, somewhat widened distally, striate, blunt. Corresponding setae 
on more anterior legs furcate, with rudimentary secondary branch. Tarsus of leg 9 (Fig. 
56) tapering, 4.1 times as long as its greatest diameter. Proximal seta thin, tapering, 
pointed, with very short pubescent most distally, seta 0.4 of length of tarsus and 2.1 
times as long as distal seta; the latter cylindrical, blunt, striate. Cuticle of tarsus 
glabrous. 

Pygidium (Fig. 57): Tergum. Posterior margin between widely separated st with 
large broadly linguiform lobe above anal plate; between lobe and st a distinct but 
shallow indentation. Relative lengths of setae: ay-a^= 10, a 2 = ?, st «7. Setae a l and 



540 U. SCHELLER 

a 3 tapering, pointed, curved inward, glabrous, the former diverging, the latter 
converging, st balloon-shaped, converging, glabrous, at most twice longer than wide. 
Distance a x - a x 0.9 of length of a ls distance a x - a 2 0.8 of distance a x - a x and 0.9 of 
distance a 2 -a^\ distance st - .st 2.8 times as long as st and 2. 1 times as long as distance 
a i -ay. Tergum glabrous. 

Sternum. Posterior margin with shallow indentation between by. Relative 
lengths of setae (öj = 10): by = 35, b 2 = 12. Setae tapering, striate distally, b 2 pointed, 
somewhat curved inward, by 1.5 times as long as interdistance; b 2 ì.ì times as long as 
distance by - b 2 . Tergum glabrous. 

Anal plate somewhat longer than broad, broadest at base, lateral margins 
concave, distal margin with shallow median indentation and 4 large appendages: two 
posterolateral, 0.8 of length of plate, somewhat diverging, pointed, outer margin almost 
straight, inner margin convex; and two submedian appendages, 0.7 of length of plate, 
protruding backward from posterior margin of sternal side of plate, these appendages 
very alike st but with very short pubescence. Plate and sternum glabrous. 

13. Allopauropus (D.) divaricatus sp. n. Figs 58-66 

Type material: Holotype: ad. 9(9), Malaysia, Sarawak, Bako National Park, Jalan 
Lintang, soil sample (extraction in Geneva) from between buttresses of Austrobuxus nitidus Miq. 
[=Longetia malayana (Benth.) P. & H.] (Euphorbiaceae), alt. 30 m, 1 1. XII. 1987 (loc. Sar-87/76, 
leg. Hauser). 

Etymology: From the Latin divarico = stretch out (referring to the appendages 
of the anal plate). 

Diagnosis: A. (D.) divaricatus sp. n. is well delimited by several characters of 
the pygidium (see below: setae by and st, anal plate), and also by the shape of the 
genital papillae and of the tarsus of the last pair of legs. These partly rare or unique 
characters, make it impossible to trace the relationships of this species at present. 

Description: Length = 0.66 mm. 

Head (Fig. 58): Setae on tergal side thin striate, of medium length, shortest on 
posteromedian part. Relative lengths of setae: 1 st row: a y = 10, a 2 = 8; 2 nd row: ay = 7, 
a 2 = 13, a 3 = 11; 3 rd row: a x - 6, a 2 = 8; 4 th row: a x - 6, a 2 = 13, a 3 = 17, a 4 = 9; lat- 
eral group setae: Zj = 17, l 2 = 13, / 3 = 1 1. Ratio a x /a x - a x in 1 st row 1.8, in 2 nd row 
0.4, in 3 rd row 0.8 and in 4 th row 1.4. Temporal organs 1.8 times as long as their 
shortest interdistance. Small posterior aperture close to posterior margin of temporal 
organ. Head cuticle glabrous. 

Antennae (Fig. 59): Segment 4 with setae p, p\ p " and r; p '-" not ascertained. 
Setae subcylindrical, blunt, striate. Relative lengths of setae: p = 100, p ' - 46, p " = 43, 
r = 49. Tergal seta/? 1.1 times as long as tergal branch t. The latter slender, cylindrical, 
4.7 times as long as its greatest diameter and 1 .5 times as long as sternal branch s, this 
2.2 times as long as greatest diameter and with anterodistal corner distinctly truncate. 
Seta q as seta/?, 1.2 times as long as s. Lengths of flagella (basal segments included) 
and basal segments: F x and bs x lost; F 2 = 52, bs 2 = 6; F 3 = 120, bs^ = 8 urn. F 2 and F 3 
2.4 and 5.6 times as long as s, respectively. Distal calyces small, distal part of flagella 
axes fusiformly widened on F 2 , cylindrically widened on F 3 . Globulus g 1.4 times as 
long as wide, its width 1 . 1 times as wide as greatest diameter of t; «9 bracts present. 
Antennae glabrous. 



RECORDS OF PAUROPODA 



541 



i & ? èmmM é& 




Figs 58-66 
Allopauropus (D.) divaricatus sp. n., holotype, ad. 9(6). (58) head, median and right part, tergal 
view; (59) right antenna, sternal view; (60) collum segment, median and left part, sternal view; 
(61) r 3 ; (62) genital papillae and seta on coxa of leg 2, anterior view; (63) seta on coxa of leg 9; 
(64) seta on trochanter of leg 9; (65) tarsus of leg 9; (66) pygidium, median and left part, sternal 
view. Scale a: Figs 58, 61, 65; b: Figs 59-60, 62-64, 66. 



Trunk: Setae of collum segment (Fig. 60) long, simple, blunt, striate. Sublateral 
seta 1.4 times as long as submedian seta; sternite process with anterior incision; 
appendages subglobular, caps very thin. Process and posterior halves of appendages 
with short pubescence. 



542 U. SCHELLER 

Setae on anterior tergites similar to submedian setae on tergal side of head. 4+4 
setae on tergite I, 6+6 on II- V, VI not available for study. 

Bothriotricha: Relative lengths: T x = 100, T 2 = 120, T 3 = 115, T 4 = 123, T 5 = 
170. All with straight simple axes, all but r 3 very thin with, extremely short pubescence 
most distally. r 3 (Fig. 61) with axis thickest in proximal 2/3, then growing very thin, 
pubescence strong, hairs simple, oblique. 

Genital papillae (Fig. 62): 2.7 times as long as greatest diameter, conical, base 
obliquely trucate, distal part thin, strongly tapering, glabrous, distal seta propor- 
tionately long, 0.8 of length of papilla. 

Legs: Setae on coxa (Fig. 63) and trochanter (Fig. 64) of leg 9 long, furcate, 
striate, main branch somewhat davate, secondary branch cylindrical, branches sub- 
similar in length. Corresponding setae on more anterior legs simple, striate. Tarsus of 
leg 9 (Fig. 65) strongly tapering, 3.5 times as long as its greatest diameter. Proximal se- 
ta lost, distal one cylindrical, blunt, striate, 0.3 of length of tarsus. Cuticle of tarsus 
glabrous. 

Pygidium (Fig. 66): Tergum. Posterior margin rounded, with broad rounded 
bulge between st. Relative lengths of setae: a\ - 100, a 2 = 68, a 3 = 1 18, st » 50. These 
setae tapering, a^, a 2 and « 3 curved inward, a x and a 2 glabrous, a 3 with very short 
pubescence distally, st curved downward, striate. Distance a x - a x 1.4 times as long as 
flj, distance a j - a 2 3 times longer than distance a 2 - a 3 ; distance st - st 1.2 times as 
long as st and 1.2 times as long as distance a x - a x . Tergum glabrous. 

Sternum. Posterior margin between b\ with broad median indentation. Relative 
lengths of setae (flj = 100): £>j = 240, b 2 = 98. Seta by curved downward-forward, 
somewhat tapering, distal half very densely striate, b 2 subcylindrical, tapering distally, 
striate, somewhat diverging, b^ 1-1 times as long as interdistance, b 2 as long as distance 

h \ - b 2- 

Anal plate large, 1.1 times as broad as long, narrowest anteriorly, with large 

lateral corners and rounded posterior margin; two widely separated, subcylindrical 

appendages protruding from posterosternal side of lateral corners, appendages strongly 

diverging, curved inward, appendages and posterior part of plate distinctly pubescent, 

lateral parts with shorter hairs. Posteromedian part of sternum pubescent. 

14. Allopauropus (D.) bakoensis sp. n. Figs 67-75 

Type material: Holotype: ad. 9($), Malaysia, Sarawak, Bako National Park, Jalan 
Lintang, soil sample (extraction in Geneva) from between buttresses of Austrobuxus nitidus Miq. 
[= Longetia malayana (Benth.) P. & H.] (Euphorbiaceae), alt. 30 m, 1 1 .XII. 1987 (loc. Sar-87/76, 
leg. Hauser). Paratypes: Malaysia: Sarawak: Serian District, Penrissen Road, 12 mis from 
Kuching, "Semongok Wildlife Rehabilitation Centre, Nursery Centre of the Forest Department", 
soil sample (extraction at Kuching) from between buttresses of large trees, alt. 50 m, 2 ad. 
9((?,9), 8.XII.1987 (loc. Sar-87/60, leg. Hauser). - Altogether 3 specimens. 

Etymology: A latinized adjective of the name Bako (referring Bako National 
Park). 

Diagnosis: An anal plate similar to that in A. (D.) bakoensis sp. n. has not been 
found in other species of the genus and is, together with the unusually long st and the 
T^ with distal swelling, delimiting this species. Its relationships cannot be traced at 
present. 



RECORDS OF PAUROPODA 



543 




Figs 67-75 
Allopauropus (D.) bakoensis sp. n., holotype, ad. 9(9). (67) head and anterior part of tergite I, 
right part, tergal view; (68) right antenna, sternal view; (69) collum segment, median and left 
part, sternal view; (70) tergite VI, posteromedian part and right posterior corner; (71) T 3 ; (72) 
genital papillae and seta on coxa of leg 2, anterior view; (73) seta on coxa of leg 9; (74) tarsus 
of leg 9; (75) pygidium, posteromedian and left part, sternal view. Scale a: Fig. 67; b: Figs 69, 
71-74; c: Figs 70, 75; d: Fig. 68. 



544 U. SCHELLER 

Description: Length = 0.51 (-0.53) mm. 

Head (Fig. 67): Tergal setae thin, striate. Relative lengths of setae (holotype on- 
ly), 1 st row: a x = a 2 = 10; 2 nd row: a l = a 3 = 15, a 2 = 19; 3 rd row: a x = 10, a 2 = 8; 4 th 
row: a x = 9, a 2 and a 3 = ?, a 4 = 10; lateral group: /j = 15, l 2 = 19, / 3 = ?. Ratio a/u^ - 
flj in 1 st row 1.1, 2 nd row 0.9, 3 rd row 0.7, 4 th row 0.8. Temporal organs twice longer 
than shortest interdistance; small pistil present in posterior part. Head cuticle glabrous. 

Antennae (Fig. 68): Segment 4 with setae p,p\ p" and r\p'" not ascertained. 
Setae striate, p,p\ p " cylindrical, blunt, r tapering. Relative lengths of setae: p = 100, 
p'= (35-)37, p" =■ (30-)32(-33), r = (51-)53. Tergal seta/? 1.9 times as long as tergal 
branch t. The latter fusiform, (1.8-)1.9 times as long as greatest diameter and 0.9 of 
length of sternal branch s, this twice longer than greatest diameter and with anterodis- 
tal corner distinctly truncate. Seta q long, cylindrical, striate, 1.8(-1.9) times as long as 
length of s. Relative lengths of flagella (basal segments included) and basal segments: 
F 1 = 100, bs i = (5-)6; F 2 = 39(-44), bs 2 = 2; F 3 = (82-)85, bs 3 = 5. F 1 (6.0-)6.1 times 
as long as t, F 2 and F 3 2.2(-2.4) and 4.7(-4.9) times as long as s, respectively. Distal 
calyces hemispherical and distal part of flagella axes fusiformly widened, strongest in 
F 2 . Globulus g (1.4-) 1.5 times as long as wide and its width (0.9-) as long as greatest 
diameter of t; «8 bracts present. Antennae glabrous. 

Trunk: Setae of collum segment (Fig. 69) simple, cylindrical, blunt, annulate. 
Sublateral seta (2.7-)2.8 times as long as submedian seta; sternite process small, 
narrow anteriorly and without apical incision; appendages short, hemispherical, with 
small caps. Both process and appendages glabrous. 

Setae on tergites short, thin, striate; 4+4 setae on tergite I, 6+6 on II- V, and 4+2 
on VI. Posterior setae on tergite VI (Fig. 70) 0.4(-0.5) of interdistance and about 
0.6(-0.7) of length of pygidial setae a\. 

Bothriotricha: Relative lengths (holotype only): T x = 100, T 2 = 97, T 3 = 83, T 4 
= 93, T 5 = 140. All with straight, simple axes, all but T 3 very thin. Axis of T 3 (Fig. 71) 
thickest in the middle, subdistally thin and with distal swelling, this twice longer than 
wide and almost 0.2 of length of bothriotrix. Pubescence short, thin, almost vertically 
erect except on T 3 ; the latter with coarse hairs, oblique on axis, erect on endswelling. 

Genital papillae (Fig. 72, of paratype): Proximal half subcylindrical, with 
concave outer side distally, 2.2 times as long as greatest diameter, glabrous; distal seta 
short, thin, 0.3 of length of papilla. Seta on coxa of leg 2 furcate, main branch cylin- 
drical blunt striate, secondary branch rudimentary, pointed, glabrous. 

Legs: Setae on coxa (Fig. 73) and trochanter of leg 9 furcate, branches thin, 
cylindrical, striate, secondary branch somewhat thinner than primary branch. 
Corresponding setae on more anterior legs with rudimentary secondary branch. Tarsus 
of leg 9 (Fig. 74) tapering, 3.3(-3.6) times as long as greatest diameter. Proximal seta 
somewhat curved, tapering, pointed, with distinct pubescence; distal seta almost 
straight, cylindrical, blunt, striate. Proximal seta (0.2-)0.3 of length of tarsus and as 
long as distal seta. Cuticle of tarsus glabrous. 

Pygidium (Fig. 75): Tergum. Posterior margin between st straight. Relative 
lengths of setae: a x = 10, a 2 = (7-)8, a 3 = 13(-14), st = 17(-19). Setae a h a 2 and a 3 , 
thin, cylindrical, tapering, directed posteriorly, a x also straight, with short pubescence, 
a 2 somewhat curved inward, a 3 somewhat curved inward and with faint pubescence 



RECORDS OF PAUROPODA 545 

distally; st tapering, somewhat curved inward, converging, glabrous. Distance a x - a x 
2.0(-2.1) times as long as a x , distance a x -a 2 0.3 of distance a x -a x and 3 times longer 
than distance a 2 - «3; distance st - st 1.4(-1.5) times as long as st and 1.2(-1.3) times 
as long as distance a x - a x . Tergum glabrous. 

Sternum. Setae b x on small but distinct posterior lobes, margin between lobes 
straight. Relative lengths of setae {a x = 10): b x = (39-)44, b 2 = 10(-1 1). Setae striate, 
b x tapering, b 2 cylindrical, somewhat curved inwards. b\ 1.4(-1.5) times as long as 
interdistance; b 2 0.8 of distance b x - b 2 . 

Anal plate narrowest anteriorly, anterior part of lateral margins somewhat 
concave, distal part convex, forming lateral rounded lobes at broadest part; posterior 
margin with shallow median indentation; plate 1.8 times as broad as long. Two 
appendages, protruding posteriorly from hindmost part of sternal side; appendages 0.5 
of the length of plate, thick, flattened distally, somewhat diverging. Plate with 
appendages distinctly pubescent, hairs longest on distal part of appendages; sternum 
glabrous. 

15. Allopauropus (D.) crassatus sp. n. Figs 76-81 

Type material: Holotype: ad. 9( 2 ), Malaysia, Sarawak, Bau, near "Fairy Cave I", soil 
sample (extraction in Geneva) from between buttresses of large trees, alt. 20 m, 13.XII.1987 (loc. 
Sar-87/82, leg. Hauser). Paratypes: Malaysia, Sarawak, Serian District, Penrissen Road, 12 mis 
from Kuching, "Semongok Wildlife Rehabilitation Centre, Nursery Centre of the Forest 
Department", soil sample (extraction in Kuching) from between buttresses of large trees, alt. 50 
m, 1 ad. 9( 2 ), 8.XII.1987 (loc. Sar-87/60, leg. Hauser). Gunung Serapi at road Kuching-Matang, 
soil sample (extraction in Kuching) in forest at the road to the television station, alt. 670 m, 
2 subad. 8(9), 9.XII.1987 (loc. Sar-87/64, leg. Hauser). 

Other material: Same data as for holotype, 1 juv. stad.?. - Altogether 5 specimens. 

Etymology: From the Latin crassus = thick; here meaning thickened (referring 
to 7/3, st and anal plate). 

Diagnosis: A. (D.) crassatus sp. n. is a close relative of A. (D.) bakoensis sp. n. 
described above. Distinctive characters are: the shape of the bothriotricha T 3 (with 
longish ends welling, almost half of length of bothriotrix in A. crassatus sp. n., about 
0.2 ofthat length in A. bakoensis sp. n.), the st (directed posteriorly and sparsely striate 
in A. crassatus sp. n., converging and glabrous in A. bakoensis sp. n.) and the shape of 
the anal plate (lateral margins convex and posterior appendages sickle-shaped in A. 
crassatus sp. n., lateral margins concave and posterior appendages with flattened distal 
surface in A. bakoensis sp. n.). Due to similarities in the antennae, the bothriotricha T 3 , 
the legs and the general shape of the anal plate A. crassatus sp. n. may also be related, 
but to a less degree, to A. (D.) thailandensis Serieller from the Chiang Mai Province in 
northern Thailand (Serieller, 1995). 

Description: Length = (0.32-)0.35(-0.39) mm. 

Head: Not available for study. 

Antennae (Fig. 76): Segment 4 with setae p, p\ p " and r; p '" not ascertained. 
Setae cylindrical, striate. Relative lengths of setae: p = 100, p ' = 63(-78), p "■=■ 68(-73), 
r = (54)58. Tergal seta p 0.9 of the length of tergal branch t. The latter fusiform, 
1.7 times as long as its greatest diameter and 1.1 times as long as sternal branch s, this 
1.7(-1.8) times as long as its greatest diameter and with its anterodistal corner distinctly 



546 



U. SCHELLER 




Figs 76-81 
Allopauropus (D.) crassatus sp. n., holotype ad. 9(9). (76) right antenna, sternal view; (77) 
collum segment, median and left part, sternal view; (78) r 3 ; (79) seta on trochanter of leg 9; (80) 
tarsus of leg 9; (81) pygidium, sternal view. Scale a: Figs 79, 80; b: Fig. 77; c: Figs 78, 81; d: 
Fig. 76. 



RECORDS OF PAUROPODA 547 

truncate. Seta q, cylindrical, striate, (1.1-)1.2 times as long as length of s. Relative 
lengths of flagella (basal segments included) and basal segments: F y = (100), bsy = 
(3-5); F 2 = (35-41), bs 2 = (3-5); F 3 = (82-91), bs 3 = (5-6). F 1 (4.8-5.4) times as long 
as t, F 2 and F 3 (1.9-2.1) and (4.2-4.6) times as long as s, respectively. Distal calyces 
hemispherical, distal part of flagella axes fusiformly widened. Globulus g 1.3(-1.4) 
times as long as wide and its width 0.6(-0.7) of greatest diameter off; »9 bracts present. 
Antennae glabrous. 

Trunk: Setae of collum segment (Fig. 77) thin, simple, cylindrical, blunt, striate. 
Sublateral seta (2.8-)2.9 times as long as submedian seta; sternite process narrow, with 
extremely small anterior incision; appendages somewhat barrel-shaped, caps low. 
Process and appendages glabrous. 

Setae on tergites short, thin, striate; 4+4 setae on tergite I, 6+6 on II- V, and 4+2 
on VI. 

Bothriotricha: Relative lengths (for holotype only): Ty = 100, T 2 = ?, T 3 = 78, 
T 4 = 98, T 5 = 170. All with straight, simple axes, all but T 3 very thin. Axis of r 3 
widening outward into a longish distal swelling, thickest in the middle, 0.4 of length of 
bothriotrix. Pubescence very short, thin, almost erect except on r 3 ; the latter (Fig. 78) 
with coarse oblique hairs. 

Legs: Setae on coxa and trochanter (Fig. 79) of leg 9, furcate, branches thin, 
cylindrical, striate, secondary branch at least on trochanter shorter than main branch. 
Corresponding setae on more anterior legs simple. Tarsus of leg 9 (Fig. 80) somewhat 
tapering, 3 times longer than greatest diameter. Setae cylindrical, striate, proximal seta 
somewhat curved and a little thinner than distal seta, the latter straight, blunt. Proximal 
seta almost 0.2 of length of tarsus and 0.8(-0.9) of length of distal seta. Cuticle of tarsus 
glabrous. 

Pygidium (Fig. 81): Tergum. Posterior margin rounded and with broad lobe 
between st, lobe about 4 times broader than long and not protruding. Relative lengths 
of setae: a x = 10, a 2 = (12-)13, a 3 = (18-)20(-21), st = (25-)27(-29). Setae tapering, a h 
a 2 and a 3 with short pubescence, ay also straight, directed posteriorly, a 2 also curved 
inward and directed posteriorly, a 3 also curved inward and diverging, st almost straight, 
directed posteriorly, sparsely striate. Distance ay - ay 3.3(-3.4) times as long as ay, dis- 
tance öj - a 2 0.6 of distance ay - ay and 3 times longer than distance a 2 - a 3 ; distance 
st - st (1.4-)1.5 times as long as st and 1.2 times as long as distance ay - ay. Tergum 
glabrous. 

Sternum. Posterior margin between by straight. Relative lengths of setae (ay = 
10): by = (60-)64, b 2 = 20(-22). Setae very little tapering, striate, b 2 somewhat curved 
inward, converging, by 1.2(-1.3) times as long as interdistance; b 2 0.7 of distance 
by - b 2 . 

Anal plate narrowest anteriorly, anterior part of lateral margins straight, 
posterior part convex; broadest part at middle, 1.4 times as broad as long; posterior 
margin with two shallow median indentations inside posterolateral corners. Two 
appendages protruding posteriorly from hindmost part of sternal side inside the inden- 
tations, appendages 0.5 of length of plate, thick, sickle-shaped, diverging. Plate with 
appendages distinctly pubescent, hairs longest at lateral margins of plate and on 
appendages. Sternum glabrous. 



548 U. SCHELLER 

Stage subad. 8. Tergite V with 6+4 setae. Setae in posterior row short, lanceo- 
late, with oblique pubescence. 

16. Allopauropus (D.) curtus sp. n. Figs 82-89 

Type material: Holotype: subad. 8(9), Malaysia, Sarawak, Serian District, Penrissen 
Road, 12 mis from Kuching, "Semongok Wildlife Rehabilitation Centre, Nursery Centre of the 
Forest Department", soil sample (extraction at Kuching) from between buttresses of large trees, 
alt. 50 m, 1 ad. subad. 8( 9 ), 8.XII.1987 (loc. Sar-87/60, leg. Hauser). 

Diagnosis: A. (D.) curtus sp. n. seems to be most closely related to A. (D.) 
adjacens Remy from the Cape Province, South Africa, (Remy, 1956b) and A. (D.) 
hortulanus Remy from Pondichéry (Remy, 1961). They have distinct similarities in the 
general shape of the bothriotricha r 3 and the anal plate. The following characters are 
useful for distinguishing these species: the shape of the axis of bothriotricha T$ (with 
short ovoid swelling in the middle in A. curtus sp. n., swelling very longish in A. 
adjacens, large davate swelling in A. hortulanus), and the shape of the pygidial setae 
st (long, distally both widened and cut obliquely in A. curtus sp. n., subcylindrical and 
blunt in A. adjacens and A. hortulanus). 

Etymology: From the Latin curtus = truncated (referring to the distal part of 
pygidial setae st). 

Description: Length = 0.45 mm. 

Head (Fig. 82): Posteromedian part of tergal side not available for study. Setae 
thin, striate. Relative lengths of setae, 1 st row: öj = 10, a 2 = 13; 2 nd row: a^ = 13, a 2 
=20, a 3 = ?, 3 rd row and first 3 setae in 4 th row not available for study, a 4 in 4 th row = 
15; lateral group: /j = 20, l 2 and / 3 = ?. Ratio a x /a x - a x in 1 st row 0.9 and in 2 nd row 
0.5. Temporal organs 1.1 times as long as their shortest interdistance. Head cuticle 
glabrous. 

Antennae (Fig. 83): Segment 4 with setae p, p', p " and r; p '" not ascertained. 
Setae thin, tapering, p, p\ p " striate, r glabrous. Relative lengths of setae: p - 100, p ' 
= 92, p" = 29, r = 42. Tergal seta p 1.2 times as long as tergal branch t. The latter 
fusiform, 1.7 times as long as greatest diameter and 0.9 of length of sternal branch s, 
this 1.4 times as long as greatest diameter and with anterodistal corner distinctly trun- 
cate. Seta q thin, cylindrical, striate, 1.2 times as long as length of s. Lengths of F\ = 
44, bs^ 5.5 urn (F 2 and F 3 lost). F 1 4.4 times as long as t. Distal calyx of Fj very small, 
distal part of flagella axis distinctly widened. Globulus g 1.4 times as long as wide and 
its width as long as greatest diameter of /; »9 bracts present. Antennae glabrous. 

Trunk: Setae of collum segment (Fig. 84) simple, cylindrical, blunt, annulate. 
Sublateral seta 1.7 times as long as submedian seta; sternite process small, pointed an- 
teriorly; appendages small, with low caps. Process and appendages glabrous. 

Setae on tergites thin, striate; 4+4 setae on tergite I, 6+6 on LT-V, and 4+2 on VI. 
Posterior setae on tergite VI not available for study. 

Bothriotricha (Figs 85-86): Relative lengths: T x = 100, T 2 = ?, T 3 = 94, T 4 = ?, 
T 5 = 115. All with straight simple axes, all thin except F 3 . The latter with davate 
proximal half, distal half very thin. Pubescence short oblique on davate part of r 3 , 
longer, erect, arranged in whirls on distal part of other bothriotricha studied. 



RECORDS OF PAUROPODA 



549 




20|jm 



86 




Figs 82-89 
Allopauropus (D.) curtus sp. n., holotype, subad. 8(9). (82) head, anterior and right side, tergal 
view; (83) right antenna, tergal view; (84) collum segment, median and left part, sternal view; 
(85) T 3 ; (86) T 5 ; (87) seta on coxa of leg 8; (88) tarsus of leg 8; (89) pygidium, posteromedian 
and right part, tergal view. Scale a: Figs 85-88; b: Figs 82, 84; c: Fig. 89; d: Fig. 83. 



Legs: Setae on coxa (Fig. 87) and trochanter of leg 8 furcate, branches similar 
to each other, thin, cylindrical, striate. Corresponding setae on more anterior legs with 
rudimentary secondary branch, coxal setae somewhat davate. Tarsus of leg 8 (Fig. 88) 
tapering, 3.4 times as long as greatest diameter. Distal seta straight, cylindrical, blunt, 
striate; proximal seta curved, tapering, pointed, glabrous. Proximal seta 0.3 of length 
of tarsus and twice longer than distal seta. Cuticle of tarsus glabrous. 



550 U. SCHELLER 

Pygidium (Fig. 89): Tergum. Posterior margin with shallow indentation between 
st. Relative lengths of setae: a x = a^ = 10, a 2 = 8, st = 12. Setae a l5 a 2 and a 3 , cylin- 
drical, glabrous, a l5 also straight, somewhat diverging, a 2 and a 3 somewhat curved 
inward; st straight, converging, somewhat widening outward, distal end obliquely 
truncate, flattened, proximal half glabrous, distal half with short pubescence. Distance 
a 1 - a y 1.1 times as long as a l5 distance a ^ - a 2 4 times longer than distance a 2 - a 3 ; 
distance st -st 1.8 times as long as st and 1.9 times as long as distance a± - ay Tergum 
glabrous. 

Sternum. Posterior margin with shallow indentation between setae b x . Relative 
lengths of setae (flj = 10): by = 32, b 2 = 10. Setae thin, somewhat tapering, b^ densely 
striate distally, b 2 glabrous. b x 1.2 times as long as interdistance; b 2 0.8 of distance 
b x -b 2 . 

Anal plate 1.8 times as long as broad, lateral margins concave, posterior part 
bilobate, lobes rounded, diverging. Four appendages protruding backwards from hind- 
most part of lobes: two in tergal position, short, cylindrical, blunt, striate, and two 
protruding from sternal side, thin, tapering, pointed, glabrous, somewhat converging. 
Tergal appendages 0.4 and sternal appendages 0.5 of length of plate. Plate and sternum 
glabrous. 

17. Allopauropus (D.) quadrispinus sp. n. Figs 90-96 

Type material: Holotype: ad. 9(9), Indonesia, Bali, Ubud, "Monkey Forest", at the 
curve in the road through the forest, under stones on slope, alt. 200 m, 30.XI.1987 (loc. Sar- 
87/37, leg. Hauser). 

Etymology: From the Latin quadric-, from quattuor = four and spina = spine, 
jag (referring to the anal plate). 

Diagnosis: A. (D.) quadrispinus sp. n. may be most closely related to A. (D.) 
vinsoni Remy from Mauritius (Remy, 1959). The anal plates have the same groundplan 
and the setae of the pygidial tergum are similar. Both species can be distinguished by 
the shape of the posterior margin of the tergum (with distinct posteromedian lobe in A. 
quadrispinus sp. n., no lobe in A. vinsoni) and by the shape of the anal plate (sub- 
median appendages 3.5 times as long as plate and no sternal appendages in A. 
quadrispinus sp. n., 2.3 times as long as plate and with two short sternal appendages in 
A. vinsoni). 

Description: Length = 0.82 mm. 

Head: Not available for detailed study, anterior setae davate, posterior ones 
cylindrical. 

Antennae (Fig. 90): Segment 4 with at least 3 setae, p,p',p", r and p'" not as- 
certained. Setae cylindrical, annulate-striate. Relative lengths of setae: p = 100, p' = 
71, p " - 54. Tergal seta p 1.4 times as long as tergal branch t. The latter fusiform, 2.5 
times as long as greatest diameter and as long as sternal branch s, this 1.8 times as long 
as greatest diameter and with its anterodistal corner distinctly truncate. Seta q thin, 
striate, at least 0.5 of length of s. Flagella Fj and F 3 broken, only base segments 
remaining; length of base segments of Fj = 5.5 urn, those of F 3 = 5 urn, length of 
flagella F 2 = 56 (im and its base segment = 3 urn. F 2 2.8 times as long as s. Distal 



RECORDS OF PAUROPODA 



551 




Figs 90-96 
Allopauropus (D.) quadrispinus sp. n., holotype, ad. 9(9). (90) left antenna, tergal view; (91) 
tergite VI, posterior part; (92) T 3 ; (93) seta on coxa of leg 9; (94) seta on trochanter of leg 9; (95) 
tarsus of leg 9; (96) pygidium, sternal view. Scale a: Figs 91-95; b: Figs 90, 96. 



calyces F 2 hemispherical, distal part of flagella axes fusiformly widened. Globulus g 
1.8 times as long as wide and its width 0.8 of greatest diameter of t; «9 bracts present. 
Antennae glabrous. 

Trunk: Collum segment not available for study. Setae on anterior tergites short, 
cylindrical, striate; on tergite VI 4+2 all short, lanceolate, pubescent; length of 
posterior setae on tergite VI (Fig. 91) 0.1 of interdistance. 

Bothriotricha: Most bothriotricha broken off but seems to be very thin. Lengths: 
r 3 = 98, T 5 = 130 urn. Axes straight, simple, pubescence short, on 7 3 (Fig. 92) longer, 
with oblique, simple hairs. 

Legs: Setae on coxa (Fig. 93) and trochanter (Fig. 94) of leg 9, large, furcate, 
their branches subequal, subcylindrical, striate, blunt. Corresponding setae on leg 8 
similar, on legs 1-7 main branches somewhat davate, secondary branches rudimentary. 
Tarsus of leg 9 (Fig. 95) tapering, 4.3 times as long as its greatest diameter. Setae 



552 U. SCHELLER 

striate, proximal seta tapering, pointed, distal seta cylindrical, blunt; the latter 0.3 of 
length of tarsus and 0.5 of length of distal seta. Cuticle of tarsus glabrous. 

Pygidium (Fig. 96): Tergum. Posterior margin with broad and low rounded lobe 
posterior of a 2 , trus l Qrj e with an extra median semicircular lobe protruding from 
between st. Relative lengths of setae: ay = 10, a 2 = 14, a 3 = 22, st = 5. Setae tapering, 
a i straight, lanceolate, diverging, striate; a 2 , a 3 and st subcylindrical, a 2 also thin, 
curved inward, converging, striate distally, a 3 also somewhat S-shaped, weakly 
pubescent distally, st somewhat S-shaped, thin, converging. Distance ay -ay 1.3 times 
as long as ay, distance a^ - a 2 2.2 times as long as distance a 2 - a 3 ; distance st - st 3 
times longer than st and 1.6 times as long as distance ay - a x . Tergum glabrous. 

Sternum. Posterior margin between by with a low median bulge below anal plate 
and shallow indentations between bulge and insertion areas of by. Relative lengths of 
setae (ay = 10): by = 34, b 2 = 15. Setae tapering, striate distally, b 2 somewhat curved 
inward, diverging, by 1.1 times as long as interdistance; b 2 0.7 of distance b x - b 2 . 

Anal plate broadest at base, lateral margins concave, converging, posterior part 
with 4 long, thin, subparalleli appendages protruding backward; submedian 
appendages longest, 3.5 times as long as plate, converging distally and with short 
pubescence there; lateral appendages 1.5 times as long as plate, glabrous, somewhat 
diverging distally. Plate with appendages and sternum glabrous. 

18. Allopauropus (D.) trapezoides sp. n. Figs 97-103 

Type material: Holotype: ad. 9(a), Malaysia, Sarawak, Bako National Park, Jalan 
Lintang, soil sample (extraction in Geneva) from between buttresses of Au strobuxus nitidus Miq. 
[= Longetia malayana (Benth.) P. & H.] (Euphorbiaceae), alt. 30 m, 1 1. XII. 1987 (loc. Sar-87/76, 
leg. Hauser). Paratype: same data as for holotype, 1 ad. 9( 9 ). - Altogether 2 specimens. 

Etymology: From the Latin trapezium = trapezium (referring to the shape of 
the anal plate). 

Diagnosis: A. (D.) trapezoides sp. n. is a very small species characterised by the 
following combination of characters not found in other species of the subgenus 
Decapauropus: antennal globulus proportionally large and with many bracts, setae of 
collum segment simple, annulate, with hemispherical endsegment, the r 3 with distinct 
pubescence of simple oblique-erect hairs, main branch of seta on coxa of leg 9 annu- 
late with hemispherical endsegment, tarsi short, anal plate linguiform, with two short 
submedian appendages on distal part of sternal side. 

Description: Length = 0.25(-0.30) mm. 

Head: Not available for study. 

Antennae (Fig. 97): Segment 4 with setae p, p\ p " and r, p '" not ascertained. 
Setae cylindrical, all but r striate, r very thin, glabrous. Relative lengths of setae (holo- 
type only):p= 100,/?'= 92,/?"= 67, r = ll. Tergal seta p 0.7 of length of tergal branch 
t. The latter somewhat fusiform, twice longer than greatest diameter and 0.9 of length 
of sternal branch s, this 1 .4 times as long as greatest diameter and with anterodistal 
corner truncate. Seta q thin, cylindrical, blunt, densely striate, 0.9 of length of s. 
Relative lengths of flagella (basal segments included) and basal segments: F y = 100, 
bs x = 7; F 2 = 47(-49), bs 2 = 3(-4); F 3 = (80-)84, bs 3 = 7. F x 5.0(-5.1) times as long as 



RECORDS OF PAUROPODA 



553 



==~****?3&3#^ 




Figs 97-103 
Allopauropus (D.) trapezoides sp. n., holotype, ad. 9(9). (97) right antenna, sternal view; (98) 
Collum segment, median and left part, sternal view; (99) T 3 ; (100) genital papillae and seta on 
coxa of leg 2, anterior view; (101) seta on coxa of leg 9; (102) tarsus of leg 9; (103) pygidium, 
posterior and left part, sternal view. Scale a: Fig. 99; b: Figs 98, 100-102; c: Fig. 103; d: Fig. 97. 



t, F 2 and F 3 2.2 and 4.0(-4.1) times as long as s, respectively. Distal part of F 2 with 
very small calyces, distal part of flagella axes cylindrically widened in F ls indistinctly 
widened in F 2 . Globulus g proportionally large, 1.5 times as long as wide, 0.7 of length 
of s and its width somewhat wider than greatest diameter of t; at least 10 bracts present. 
Antennae glabrous. 

Trunk: Setae of collum segment (Fig. 98) simple, somewhat davate, annulate, 
with hemispherical endsegment. Sublateral seta 3 times longer than submedian seta; 
sternite process triangular, without anterior incision; appendages barrel-shaped, low, 
flat. Process and appendages glabrous. Setae on tergites not available for study. 



554 U. SCHELLER 

Bothrìotricha: Most bothriotricha broken off. Lengths: T 2 = 60, T 3 = 60(62), 
r 5 = 67(75) |nm. Axes straight, thin. 7 3 (Fig. 99) thickest, with distinct pubescence of 
simple, oblique - erect hairs. 

Genital papillae (Fig. 100): Genital papillae conical, 1.8 times as long as 
greatest diameter, glabrous; distal seta thin, 0.8 of length of papilla. 

Legs: Setae on coxa (Fig. 101) of leg 9 short, furcate, main branch davate, 
annulate, apical segment hemispherical, secondary branch thin, somewhat shorter than 
main branch. Seta on trochanter and corresponding setae on more anterior legs not 
available for study, except on coxa of leg 2, seta there short, furcate, with branches 
subequal in length, main branch, cylindrical, striate, blunt, secondary branch thin, 
pointed, striate. Tarsus of leg 9 (Fig. 102) short tapering, 2.4(2.6) times as long as its 
greatest diameter. Proximal seta thin, somewhat curved, tapering distally, almost 
glabrous; distal seta somewhat davate, striate. Proximal seta 0.3 of length of tarsus and 
1.2 times as long as distal seta. Cuticle of tarsus glabrous. 

Pygidium (Fig. 103): Tergum. Posterior margin rounded, two small posterior 
submedian bulges between st. Relative lengths of setae: a^-a 2 = st =10, a 2 = (8)10. 
a 3 = (11)12. Setae a^,a 2 and a 3 thin, cylindrical, glabrous, a x also straight, posteriorly 
directed, a 2 and a 3 somewhat curved inward, a-, converging, a 3 directed backward; st 
straight, somewhat davate, striate, directed posteriorly. Distance a x - a^ (1.4)1.5 times 
as long as ûj, distance a^ - a 2 2.3 times as long as distance a 2 - a 3 ; distance st - st 1.5 
times as long as st and as long as distance a^ —&\- Tergum glabrous. 

Sternum. Posterior margin between b\ with broad shallow indentation. Relative 
lengths of setae {a x = 10): b x - 25(27), b 2 = 10(11). b\ cylindrical, blunt, striate, b 2 
tapering, somewhat curved inward, b^ 1.1 times as long as interdistance; b 2 0.7 of 
distance by - b 2 . 

Anal plate narrowest at base, trapezoid, lateral margins somewhat convex, 
posterior margin rounded with small median indentation; two short, subglobular, sub- 
median, diverging appendages protruding downward from posterior part of sternal side. 
Plate and sternum glabrous. 

Genus Rabaudauropus Remy, 1953 

19. Rabaudauropus dispar Serieller 

Rabaudauropus dispar Serieller, 1994 (in Scheller et al, 1994): 5-7, figs 20-30. 

Material examined: Malaysia, Sarawak, Road Kuching-Serian, 18 km from Kuching, 
near Kampong Kuap, secondary forest, soil sample (extraction in Geneva) from between 
buttresses of a large tree, alt. 30 m, 1 juv., 13.XII.1987 (loc. Sar-87/86, leg. Hauser). 

General distribution: The species was previously known from two sites in 
Sabah only (Scheller et al, 1994). 

Scleropauropodinae 

Genus Scleropauropus Silvestri, 1902 
Subgenus Scleropauropoides Remy, 1957c 

20. Scleropauropus (S.) singapuranus sp. n. Figs 104-109 

Type material: Holotype: ad. 9(9), Singapore, Botanical Garden, in the section 
"Jungle", under bark, alt. 25 m, 16.XII.1987 (loc. Sar-87/89, leg. Hauser). 



RECORDS OF PAUROPODA 



555 




Figs 104-109 
Scleropauropus (S.) singapuranus sp. n. holotype, ad. 9(2). (104) head, median and right part, 
tergal view; (105) right antenna, sternal view; (106) tergite VI, posteromedian part; (107) T 3 ; 
(108) tarsus of leg 9; (109) pygidium, posteromedian and left part, sternal view. Scale a: Figs 
104, 106-108; b: Figs 105, 109. 



Etymology: A latinized adjective of the name Singapura (Singapore). 

Diagnosis: Only four species have been described in this subgenus: 
S. squameus Remy from the Ivory Coast (Remy, 1948b), S. mameti Remy from 
Mauritius (Remy, 1959) and Sri Lanka (Scheller, 1970), S. prunilis Scheller from Sri 
Lanka (Scheller, 1970) and S. quadriramosus Scheller from Thailand (Scheller, 1995). 
S. singapuranus sp. n. is closest to S. mameti but can be distinguished from that species 
by the shape of the setae of the 2 nd row of the tergal side of the head (cylindrical and 



556 U. SCHELLER 

striate in S. singapuranus sp. n., lanceolate, with pubescence in S. mameti) and by the 
shape of the temporal organs of the head (in tergal view of the same breadth in anterior 
and posterior parts in S. singapuranus sp. n., much broader in anterior part than 
posteriorly in S. mameti). 

Description: Length = 0.76 mm. 

Head (Fig. 104): Setae in the two anterior rows cylindrical, blunt, striate, in the 
two posterior rows lanceolate, glabrous or with indistinct pubescence. Relative lengths 
of setae, 1 st row: ay = 10, a 2 = 16; 2 nd row: ay = a 2 = 14, a 3 = 20, 3 rd row: ay = a 2 = 
10, 4 th row: a x - a 2 - a 3 - 15, ö 4 = 20; setae of lateral group tapering, pointed, with 
oblique pubescence: l\ = l 2 = 34, / 3 = 36. Ratio ay/ay - ay in 1 st and 3 rd rows 0.8, in 
2 nd and 4 th rows 0.6. Temporal organs mainly in lateral position, length 0.9 of shortest 
interdistance. Head cuticle distinctly granular anteriorly and inside temporal organs, 
coarsely granular in posteromedian and posterolateral part. 

Antennae (Fig. 105): Segment 4 with setae p, p',p" and r. Setae p, p',p" cylin- 
drical, blunt, striate, r very thin, tapering, glabrous. Relative lengths of setae: p = 100, 
p' ' = 83,/?"and r- 28. Tergal seta/? 0.8 of length of tergal branch t. The latter fusiform, 
3.1 times as long as its greatest diameter and 1.2 times as long as sternal branch s, this 
1.8 times as long as greatest diameter and with its anterodistal corner distinctly trun- 
cate. Seta q thin, tapering, striate, 0.6 of length of s. Relative lengths of flagella (basal 
segments included) and basal segments: Fy = 100, bsy = 5; F 2 = 57, bs 2 = 3; F 3 = 52, 
bs 3 = 5. Fy 4.5 times as long as t, F 2 and F 3 3.1 and =2.8 times as long as s, respec- 
tively. Distal calyces small, hemispherical, distal part of flagella axis only slightly 
widened. Globulus g 1.3 times as long as wide and its width 1.1 times as long as 
greatest diameter of t; =9 bracts present. Small posterior pistil in temporal organs. 
Antennae glabrous. 

Trunk: Collum segment not available for study. Setae on tergites lanceolate, on 
anterior tergites glabrous, on posterior tergites with sparse oblique pubescence; 4+4 
setae on tergite I, 6+6 on II- V, and 4+2 on VI. Posterior setae on tergite VI (Fig. 106) 
0.2 of interdistance. All tergites with coarsely granular cuticle. 

Bothriotricha: Relative lengths: T x = 100, T 2 = 99, T 3 = 62, T 4 = 57, T 5 = 135. 
All with straight simple axes, all thin except T 3 . The latter (Fig. 107) with thick axis 
and distal endswelling, 3.3 times as long as wide, 0.2 of length of bothriotrix. 
Pubescence on Tj, T 7 , T 4 and T 5 short, erect distally, F 3 glabrous in proximal half, 
endswelling with distinct pubescence of short, simple, oblique hairs. 

Legs: Setae on coxa and trochanter not available for study. Tarsus of leg 9 (Fig. 
108) slender, tapering, 3.8 times as long as its greatest diameter. Setae curved, proximal 
seta tapering, pointed, glabrous, distal seta similar but not pointed. Proximal seta 0.3 
of length of tarsus and 2.4 times as long as distal seta. Cuticle of tarsus glabrous. 

Pygidium (Fig. 109): Tergimi. Posterior margin evenly rounded. Relative lengths 
of setae: ay — a 2 - <3 3 = 10, st = 7. Setae short, lanceolate, glabrous, ay, a 2 and a 3 also 
somewhat diverging, st directed posteriorly. Distance ay - ay 1.4 times as long as ay, 
distance ay - a 2 0.5 of distance a 2 - a 3 ; distance st - st 3.6 times as long as st and 1.8 
times as long as distance a y —cl\. Tergum glabrous. 

Sternum. Posterior margin between setae by with two rounded bulges and a dis- 
tinct median indentation. Relative lengths of setae (ay - 10): by = 43, b 2 = 14. by sub- 



RECORDS OF PAUROPODA 557 

cylindrical, blunt, very densely striate most distally, directed downward, b 2 lanceolate; 
b i 0.9 of interdistance; b 2 0.4 of distance b x -b 2 . 

Anal plate narrowest at base, with one pair of lateral lobes and one pair of sub- 
median posterior lobes; lobes separated by U-shaped indentations; each lobe with a 
posteriorly directed, thin, tapering appendage. Appendages protruding from lateral 
lobes short, 0.3 of length of plate, appendages of posterior lobes 1.4 times as long as 
plate. Plate with appendages and sternum glabrous. 

Brachypauropodidae 

Genus Brachypauropoides Remy, 1952 

21. Brachypauropoides penanorum Serieller 

Brachypauropoides penanorum Serieller, 1994 (in: Scheller et al, 1994): 8-11, figs 31-42. 

Material examined: Malaysia, Sarawak, Serian District, Penrissen Road, 12 mis from 
Kuching, "Semongok Wildlife Rehabilitation Centre, Nursery Centre of the Forest Department", 
soil sample (extraction at Kuching) from between buttresses of large trees, alt. 50 m, 1 juv. 3, 
8.XIL1987 (loc. Sar-87/60, leg. Hauser). 

General distribution: Previously known from a single locality in Sabah only 
(Scheller et al, 1994). 

EURYPAUROPODIDAE 

Eurypauropodinae 

Genus Samarangopus Verhoeff, 1934 

22. Samarangopus longipenes Serieller 

Samarangopus longipenes Scheller, 2001: 965-969, figs 39-53. 

Material examined: Malaysia, Sarawak, Serian District, Penrissen Road, 12 mis from 
Kuching, "Semongok Wildlife Rehabilitation Centre, Nursery Centre of the Forest Department", 
soil sample (extraction at Kuching) from between buttresses of large trees, alt. 50 m, 6 ad. 9( 9 ), 
8.XII.1987 (loc. Sar-87/60, leg. Hauser). - Altogether 6 specimens. 

General distribution: The species was previously known from Sabah only 
(Scheller, 2001). 

23. Samarangopus trilix sp. n. Figs 110-124 

Type material: Holotype: ad 9( 6 ), Malaysia, Sarawak, Serian District, Penrissen Road, 
12 mis from Kuching, "Semongok Wildlife Rehabilitation Centre, Nursery Centre of the Forest 
Department", soil sample (extraction at Kuching) from between buttresses of large trees, alt. 
50 m, 8.XE.1987 (loc. Sar-87/60, leg. Hauser). 

Etymology: From the Latin trilix = woven with three threads (referring to the 
cuticle of the tergites). 

Diagnosis: S. trilix sp. n. closely resembles S. browni Remy from New 
Caledonia (Remy, 1957b). Distinctive character are e.g., the shape of the stalk of the 
antennal globulus g (long and subcylindrical in S. trilix sp. n., shorter and conical in 
S. browni), the shape of the sternal antennal branch (3.2 times as long as greatest 
diameter in S. trilix sp. n., about 2.3 in S. browni) and the shape of setae Z? 3 of the 



558 



U. SCHELLER 




Figs 110-124 
Samarangopus trilix sp. n., holotype, ad. 9(9). (110) left antenna, sternal view; (111) collum 
segment, median and left part, sternal view; (112) body, tergal view, tergites I- VI showing areas 
with concentration of funnel-shaped organs; (113) tergite I, anterior margin, sternal view 
showing marginal protuberances, internal latice and funnel-shaped organs; (114) tergite IV, 
lateral margin anterior of r 3 , tergal view; (115) tergite I, central part, cuticular pattern with 
internal latice and funnel shaped organs; (116) tergite II, right posterolateral part; (117) tergite 
III, central part, cuticular structures; (118) wart-like structure with funnel-shaped organ; (119) 
r 3 ; (120) genital papillae and seta on coxa of left leg 2; (121) seta on coxa of leg 9; (122) tarsus 
of leg 9; (123) femur of leg 1 with appendage; (124) pygidium, median and left part, sternal view. 
Scale a: Figs 121-122; b: Fig. 120;c: Figs 110-111, 113-117, 119, 123-124; d: Fig. 118. 



RECORDS OF PAUROPODA 559 

pygidial sternum (glabrous and pointed, length 0.2 of interdistance in S. trilix sp. n., 
cylindrical, blunt, striate, 0.5 of interdistance in S. browni). 

Description: Length. 1.56 mm. 

Antennae (Fig. 110): Antennae glabrous; segment 4 with 3 tapering striate se- 
tae, their lengths 15 urn. Sternal branch s with shallow anterior indentation, anterior 
margin = 23 urn, posterior margin = 38 um, of base = 7 um, maximum 0=12 urn, 
q thin, cylindrical, annulate- striate, 1 = 17 (im. Posterior margin/length of g 2.1, 
posterior margin/maximum = 3.0, maximum 0/0 of base = 1.7. Tergal branch sub- 
cylindrical, 1 = 43 urn, of base = 6 urn, maximum 0=11 um; pore not ascertained; 
length of f/maximum = 3.9. Globulus g, 1 = 17 urn, maximum = 8 urn; length/- 
maximum = 2.1; number of bracts 9, their length = 7 urn, capsule subspherical, = 
4 urn. Relative lengths of flagella (base segments included) and base segments: Fj = 
100, bsi = 22, F 2 = 87, bs 2 = 8, F 3 = ?, bs^ = 22. F 1 3 times longer than tergal branch 
t, F 2 3 times longer than sternal branch s. Calyces of F^ largest, subhemispherical, 
those of F 2 and F 3 , hemispherical. 

Trunk: Submedian setae of collum segment (Fig. Ill) short, furcate, branches 
subcylindrical, with short pubescence, 1 = 6 urn. Sternite process broad, rounded 
anteriorly, no anterior incision. Appendages large but short, cylindrical, caps flat, some- 
what larger than appendage itself. Process and appendages glabrous. 

Tergites (Fig. 112): Tergite I with a single marginal row of short, somewhat 
davate, glabrous protuberances (Fig. 113); lateral margins of other tergites (Fig. 114) 
with short spatulate protuberances. Cuticle thick, inner part of all tergites with cylin- 
drical canals or supports around cavities in a complicated three-dimensional network 
(Figs 113, 115). Inner parts of tergites with surface structures of peculiar shape and 
unknown function (Figs 116-117). Cuticular surface near margins and especially on 
posterior parts of tergites with many wart-like structures with a small transparent 
funnel on top and a dense collar of small davate transparent appendages surrounding 
its base (Fig. 118). 

Length/width ratio of tergites: I = 0.6, II and IV = 0.4, III, V and VI = 0.5. 

Bothriotricha: All bothriotricha but F 3 with very thin axes; these glabrous 
except for a minute pubescence on their distal third. F 3 (Fig. 1 19) with thicker axes and 
distal endswelling with weak pubescence. Relative lengths of bothriotricha: Fj = 100, 
T 2 = 106, F 3 = 50, F 4 = 94, T 5 = 82. 

Genital papillae (Fig. 120): Base segments cylindrical. Length of papillae = 
56 urn, greatest = 35 urn, length of seta = 20 urn. Proximal part of genital papillae 
subcylindrical, distal part conical, seta 0.4 of length of papilla, this 1.6 times as long as 
greatest diameter. Cuticle glabrous. Coxal seta of leg 2 as on leg 1, length = 20 urn. 

Legs: All legs 5-segmented. Seta on coxa (Fig. 121) and trochanter of leg 9 
similar to each other, thin, furcate, striate, with glabrous base; length of secondary 
branch 0.7 of primary one. More anteriorly these setae with rudimentary pointed 
glabrous secondary branches. Tarsi short, tapering, those of leg 9 (Fig. 122) 2.3 times 
as long as greatest diameter, with two tergal setae, both pointed and glabrous. Proximal 
seta 15 urn, distal one 10 urn. Proximal setae 0.3 of length of tarsus and 1.5 times as 
long as distal seta. Cuticle of tarsus glabrous. No proximal seta on tarsus of leg 1. All 
legs with large main claw and small setose anterior secondary claw; in leg 9 the former 



560 U. SCHELLER 

reaching 0.5 of length of tarsus. A blunt glabrous appendage, length = 7 urn, on 
anterior side of femur of leg 1 (Fig. 123). 

Pygidium (Fig. 124): Tergum. Posterior margin evenly rounded. Setae glabrous, 
a x very short, davate, curved inward, diverging; a 2 and a 3 straight, cylindrical, 
posteriorly directed, somewhat diverging; st straight, broadly lanceolate, diverging. 
Lengths of setae: a\ = st = 6, a 2 = 12, a 3 = 20 urn. Distance a x - a x = 6, a 2 - a 2 = 34, 
a 3 - a 3 = 54, flj - a 2 = 9, a 2 ~ fl 3 = H> ^ ~ 5? = 19 urn. Distance a^ - ay as long as flj, 
distance a^ - a 2 0.8 of distance a 2 - a 3 ; distance st - st 3.2 times as long as st and 0.8 
of distance a x - a x . Cuticle glabrous. 

Sternum. Posterior margin between b\ almost straight. Setae by lost, b 2 tapering, 
pointed, distal 2/3 with short pubescence, /> 3 lanceolate, glabrous. Lengths of setae: b 2 
= 25, b 3 = 7 urn. Distance b 1 -b l = 48, b 2 - b 2 = 85, b 3 - b 3 = 31, b { - b 2 = 68, /3 2 - ^3 
= 55 urn. & 2 0.7 distance b^ - b 2 , £> 3 0.2 of interdistance. 

Anal plate 1.2 times as long as wide, widest in anterior third, broadest part 
forming indistinct lateral corners, from there two short, thin, cylindrical, blunt, 
glabrous appendages protruding obliquely backward; appendages 0.4 of length of 
plate; posterior 2/3 of plate divided into two tapering branches by a narrow V-shaped 
incision, each branch with two appendages: a submedian, very short, tapering, glabrous 
point, and outside it a stalked bladder, in sternal view similar to a knife-blade. The 
latter 0.6 of length of plate. Plate and sternum glabrous. A shield-shaped plate with 
digitiform posterior appendage protrudingbackward from between st. 

24. Samarangopus tuberosus sp. n. Figs 125-137 

Type material: Holotype: ad 9( 9 ), Singapore, Labrador Hill, dry forest, soil sample 
(extraction at Bogor, Java) from under trees with small buttresses, alt. 40 m, 21. XI. 1987 (loc. 
Sar-87/1, leg. Hauser). 

Etymology: From the Latin tuberosus = full of protuberances, nodose 
(referring to the tergites). 

Diagnosis: S. tuberosus sp. n. is close to S. segniter Scheller from Sabah 
(Scheuerer ai, 1994). Both have similar antennae, the same type of cuticular structures 
on the tergites and the pygidial setae b 2 are large and lanceolate. Distinguishing char- 
acters are: the shape of the distal part of the lateral protuberances of tergites I-V (cut 
almost squarely or rounded in S. tuberosus sp. n., triangular in S. segniter), the surface 
of r 3 (glabrous in S. tuberosus sp. n., with distinct pubescence in S. segniter), the 
length of the branches of the seta on the trochanter of leg 9 (primary branch twice 
longer than secondary branch in S. tuberosus sp. n., equal in length in S. segniter), the 
shape of the pygidial setae a^ and st (flj straight and st large and lanceolate in S. tu- 
berosus sp. n., a i curved inward and st short, cylindrical, blunt in S. segniter). 

Other allied species are S. umbraculus Scheller and S. spathaceus Scheller from 
New Caledonia (Scheller, 1993), S. poculifer Scheller from Thailand (Scheller, 1995) 
and, S. jacobsoni (Silvestri) from Java (Silvestri, 1930). 

Description: Length = 0.80 mm. 

Antennae (Fig. 125): Glabrous; segment 4 with 3 striate setae: p and p' cylin- 
drical blunt, p " tapering pointed, their lengths: p and p " - 10 um, p ' - 8 urn. Sternal 
branch s: anterior margin = 1 1 urn. posterior margin = 14 urn, of base = 7 urn, 



RECORDS OF PAUROPODA 



561 




Figs 125-137 
Samarangopus tuberosus sp. n., holotype, ad. 9(9). (125) left antenna, sternal view; (126) 
collum segment, median and left part, sternal view; (127) body, tergal view, tergites I-VI showing 
areas with elevated cuticle; (128) tergite I, anterior marginal protuberances, sternal view; (129) 
tergile I, right posterolateral corner, sternal view; (130) tergite II, anterolateral comer at insertion 
area of 7^, marginal protuberances and cylindrical organs with umbrella-shaped top, tergal view; 
(131) tergite II, posterolateral corner with marginal protuberances and cylindrical organs with 
umbrella- shaped top, tergal view; (132) tergite VI, posterior part, tergal view; (133) T 3 ; (134) 
seta on coxa of leg 9; (135) tarsus of leg 9; (136) femur of leg 1 with appendage; (137) pygi- 
dium, median and left part, sternal view. Scale a: Fig. 132; b: Figs 128-131, 134-136; c: Figs 
125-126, 133, 137; d: Fig. 134. 



562 U. SCHELLER 

maximum = 9 um, q tapering, striate, 1 = 17 urn. Posterior margin/length of g 1.6, 
posterior margin/maximum = 1.6, maximum 0/0 of base = 1.3. Tergal branch 
fusiform, 1 = 29 urn, of base = 4 urn, maximum = 6 urn; pore not ascertained; 
length of //maximum = 3.3. Globulus g, 1 = 9 urn, maximum = 5 urn; length/- 
maximum = 1.8; number of bracts 7, their length = 4 urn, capsule subspherical, 
bottom somewhat flattened, = 3 urn. Relative lengths of flagella (base segments 
included) and base segments: Fj = 100, bs^ = 13, F 2 = 73, bs 2 = 7, F 3 = 58, bs^ = 13. 
F-[ 3.6 times as long as tergal branch t, F 2 1.9 and F 3 3.4 times as long as sternal branch 
s, respectively. Calyces of F x largest, conical, those of F 2 and F 3 hemispherical. 

Trunk: Submedian setae of collum segment (Fig. 126), furcate, main branch 
cylindrical and striate, secondary branch rudimentary, pointed, glabrous; submedian 
seta 1 = 10 urn, lateral seta 1 = 9 urn. Sternite process broad, pointed anteriorly. 
Appendages barrel-shaped, directed posteriorly, caps somewhat larger than distal part 
of appendage. Process glabrous, appendages almost so. 

Tergites (Fig. 127): With 4 types of protuberances: 1. campanulate on anterior 
and lateral margins of tergite I (Figs 128-129); 2. wedge- or leaf-shaped on lateral 
margins of tergites II- VI (Fig. 130), these small or rudimentary at insertion cavities of 
bothriotricha; 3. smaller, cylindrical, cut squarely distally and with a small umbrella- 
like structure protruding from an apical cavity (Figs 130-132), these partly concen- 
trated in groups on rounded elevations of the cuticle or in dense rows near margins of 
tergites; 4. Small, flat, blunt teeth in groups of 3-5 at posterior margins of tergites I-V 
(Fig. 131). 

Number of marginal protuberances: I, 23; II, 1 small -7^-9 (one smaller); III, 
1 small - 3 - T 2 - 9; IV, 1 small - 4 - T 3 - 4; V, 1 small - 5 - T 4 - 1 small - 3; VI, 6 
-1*5-1. Length/width ratio of tergites: I = 0.7, II-IV = 0.4, V = 0.5 and VI = 0.6. 

Bothriotricha: All bothriotricha but F 3 with very thin axes; these glabrous, 
curled distally. F 3 (Fig. 133) with glabrous distal swelling, this 0.3 of length of 
bothriotrix. Relative lengths of bothriotricha: T } = 100, T 7 = 96, T 3 = 54, T 4 = ?, 

r 5 = 78. 

Legs: All legs 5-segmented. Seta on coxa (Fig. 134) and trochanter of leg 9 
similar to each other, thin, furcate, striate, with glabrous base; length of secondary 
branch 0.5 of primary one. More anteriorly these setae thinner and with rudimentary 
pointed glabrous secondary branches. Tarsi short, tapering, those of leg 9 (Fig. 135) 
twice longer than greatest diameter, with two tergal setae, both pointed and glabrous. 
Proximal seta 10, distal one 15 urn. Proximal setae 0.3 of length of tarsus and twice 
longer than distal seta. Cuticle of tarsus glabrous. No proximal seta on tarsus of leg 1. 
All legs with large main claw and small setose anterior secondary claw; in leg 9 the for- 
mer reaching 0.5 of length of tarsus. A pointed glabrous shortly pubescent appendage 
on anterior side of femur of leg 1 (Fig. 136), its length = 4 urn. 

Pygidium (Fig. 137): Tergum. Posterior margin rounded but with a pentagonal 
plate above anal plate protruding backward from between st, this plate narrowest 
anteriorly and with obtuse posterolateral corners. Two small posteriorly directed jags 
protrude backward at the level of setae a->. Setae glabrous, a ls a 2 and a 3 straight, cylin- 
drical, glabrous, öj somewhat diverging, a 2 converging, a 3 straight, cylindrical, 
posteriorly directed; st lanceolate, curved inward, directed posteriorly. Lengths of 



RECORDS OF PAUROPODA 563 

setae: a x -l pm, a 2 = 5 pm, a 3 = 1 1 pm, st = 13 pm. Distance a x - a x - 10 pm, a 2 - 
At = 30 pm, a 3 - a 3 = 35 pm, a x - a 2 = 12 pm, a 2 - a 3 = 2 urn, st-st= 13 pm. Distance 
flj - aj 1.4 times as long as a^, distance ay - a 2 6 times longer than a 2 - a 3 ; distance 
.st - 5/ as long as st and 1.3 times as long as distance a x - atj. Cuticle glabrous. 

Sternum. Posterior margin between £>j almost straight. Setae &j thin, tapering, 
striate most distally, b 2 large, lanceolate, with transparent wings, glabrous, b 3 tapering, 
pointed, striate most distally. Lengths of setae: b x = 30 urn, b 2 = 20 urn, Z? 3 = 10 urn. 
Distance by - b\ = 39 urn, b 2 - b 2 = 130 um, & 3 - b 3 = 22 pm, b { - b 2 = 21 pm, b 2 - 
b 3 = 22 urn. b 2 as long as distance b\ - b 2 , & 3 0.4 of interdistance. 

Anal plate 1.2 times as long as broad, broadest behind middle, lateral margins 
convex anteriorly, concave posteriorly; distal part cleft by a deep V-shaped incision in- 
to two tube-like branches; each branch with a short, submedian, straight, thorn-like, 
glabrous appendage, and a stalked leaf-shaped, knife-like posteriorly directed bladder, 
with granular surface, length of the latter appendage 0.7 of length of plate. Plate and 
sternum glabrous. 

25. Samarangopus interstinctus sp. n. Figs 138-150 

Type material: Holotype: ad 9( 3 ), Malaysia, Sarawak, Kuching-Matang road, Gunung 
Serapi, soil sample (extraction at Kuching) from forest along road to the TV-station, alt. 320 m, 
9.XII.1987 (loc. Sar-87/66, leg. Hauser). Paratypes: 3 ad. 9(1 o\ 2 2), 1 subad. 8(9), same data 
as for holotype; 1 ad (ct), 1 subad. 8( 9 ), Kuching-Matang road, Gunung Serapi mountain, soil 
sample (extraction at Kuching) from forest along road to the TV-station, alt. 670 m, 9. XII. 1987 
(loc. Sar-87/64, leg. Hauser). 

Other material: Malaysia, Sarawak, Kuching-Serian road, near Kampong Kuap 
(18 km from Kuching), secondary forest, soil sample (extraction in Geneva) from between but- 
tresses of a large tree, alt. 30 m, 6 ad. 9 (lo\59), 2 subad. 8(2), 13.XII.1987 (loc. Sar-87/86, 
leg. Hauser). -Altogether 15 specimens. 

Etymology: From the Latin interstinguo, interstinctus = here and there 
bestrewed with (small fungiform organs on the tergites). 

Diagnosis: The species described here is close to S. poculifer Serieller from 
Thailand (Serieller, 1995). These two species are generally alike in their antennae, their 
cuticular appendages of the tergites and in the shape of the anal plate. They can be dis- 
tinguished by the shape of the process of the collum segment (rounded anteriorly in S. 
interstingctus sp. n.; cleft anteriorly in S. poculifer), the shape of the protuberances of 
the anterior margin of tergite I (large and campanulate in S. interstinctus sp. n.; small 
and wedge-shaped in S. poculifer), by the shape of the specialised protuberances of the 
tergites (davate central column with umbrella-like transparent covering disc in S. 
interstinctus sp. n.; funnel-like, consisting of a central column surrounded by a mem- 
brane in the shape of an upside down umbrella or a wide funnel in S. poculifer). Good 
distinctive characters are also found in the shape of the appendages of the postero- 
lateral corners of the pygidial tergum and its setae a 3 . There are also similarities, but to 
a less degree, with S. papuensis Serieller from Papua New Guinea (Serieller, 1996) and 
5. segniter Serieller from Sabah (Scheller et al, 1994) as to the antennae, the cuticle of 
the tergites and some pygidial characters. 

Description: Length = (0.41-)0.42(-0.48) mm. 

Antennae (Fig. 138): Antennae glabrous; segment 4 with 3 cylindrical, blunt, 
sparsely and weakly striate setae, their lengths: p = p" ' = 7(-8) pm, p' = 6(-7) pm. 



564 



U. SCHELLER 




Figs 138-150 
Samarangopus interstinctus sp. n., holotype, ad. 9(S). (138) right antenna, sternal view; (139) 
collum segment, median and left part, sternal view; (140) tergite I, anterior margin with campa- 
nulate protuberances and fungiform organs, tergal view; (141) tergite I, posterolateral corner 
with marginal protuberances, sternal view; (142) tergite I, central part with elevated cuticle and 
fungiform organs; (143) tergite I, posterior margin; (144) tergite VI, posterior margin between 
insertion cavities of T 5 tergal view; (145) fungiform organ; (146) T 3 ; (147) genital papillae and 
seta on coxa of leg 2, anterior view; (148) seta on coxa of leg 9; (149) tarsus of leg 9; (150) 
pygidium, median and left part, sternal view. Scale a: Figs 140-143, 145, 147; b: Fig. 139; c: Figs 
138, 144, 146, 148-150. 



RECORDS OF PAUROPODA 565 

Sternal branch s, with shallow anterior indentation, anterior margin = 7 urn, posterior 
margin = 11(-12), of base = 5(-6), maximum = 7(-8) um, q cylindrical, blunt, 
striate, 1 = 12(-14) urn. Posterior margin/length of g 1.7(-2.9), posterior margin/- 
maximum = 1.5(-1.6), maximum 0/0 of base = (1.3-)1.5. Tergal branch t somewhat 
fusiform, 1 = (12-) 13 urn, of base = 5, maximum = 6 urn; pore not ascertained; 
length off/maximum = (2.0-)2.2 . Globulus g, 1 = 6(-7) (im, maximum = (4-)5 urn; 
length/maximum = 1.2(-1.4); number of bracts 8(-9), their length = 5 um, capsule 
hemispherical with flattened bottom, = 3 (-4) urn. Relative lengths of flagella (base 
segments included) and base segments: F x = 100, bs^ = 16(-17), F 2 = 42(-48), bs 2 = 
(9-)10(-12), F 3 = 83(-90), bs 3 = 13(-14). F x 4(-4.4) times as long as tergal branch t, F 2 
and F 3 (1.6-)1.7 and 3.4(-3.5) times as long as sternal branch s. Calyces subhemi- 
spherical, those of Fj largest, those of F 2 smallest. 

Trunk: Setae of collum segment (Fig. 139) short, furcate, primary branch cylin- 
drical, blunt, striate, secondary branch rudimentary, glabrous, submedian seta 1 = 6 urn, 
somewhat shorter than sublateral seta. Sternite process broad, anteriorly rounded, no 
incision. Appendages short, subcylindrical, caps flat. Process and appendages glabrous. 

Tergites: Anterior and lateral margins of tergite I (Figs 140-141) with a single 
row of campanulate, glabrous protuberances; lateral margins of other tergites with 
similar type of protuberances, but longish. Number of marginal protuberances: I. 
(23-)24(-25); H, 1 small -T x -\ small + 7(-9) + 1 small; III, 4 - T 2 - 1 small + 4(-5) 
+ 1 small; IV, (4-)5 - T 3 - 1 small + 4; V, (4-)5 - T A - 1 small + 3; VI, (1 small + 3)4 - 
T 5 - l(-2) + 2 small. Cuticle (Figs 142-143) thick, with coarse surface set with small 
cones and several fungus- or umbrella-like organs consisting of a davate or almost 
cylindrical stalk covered by a circular, transparent plate similar an umbrella or a hat of 
a fungus (Figs 140, 145). 

Bothriotricha: All but F 3 with very thin axes; these glabrous except for a minute 
pubescence on their distal third. F 3 (Fig. 146) glabrous and with thicker axes and dis- 
tal swelling. Relative lengths of bothriotricha (for holotype only): Fj = 100, T 2 = 114, 
T 3 = 68, T 4 = 90, T 5 = 81. 

Genital papillae (Fig. 147): Base segments long and wide, in the shape of a 
truncated cone, papilla conical, with strongly narrowing distal part. Length of papillae 
= 20 um, greatest = 12 urn, length of seta = 30 urn; papilla 1.7 times as long as 
greatest diameter, seta 1.5 times as long as length of papilla. Cuticle glabrous. Coxal 
seta of leg 2 with short and thin but not rudimentary secondary branch, length of seta 
= 13 urn. 

Legs: All legs 5-segmented. Seta on coxa (Fig. 148) and trochanter of leg 9 
similar to each other, furcate, striate; length of secondary branch 0.6 of primary one. 
These setae on more anterior legs with rudimentary pointed glabrous secondary 
branches, but not so in leg 2 in males. Tarsi short, tapering, length in leg 9 (19-)20 urn, 
in leg 1 (14-)15(-16) urn. Tarsus of leg 9 (Fig. 149) (1.9-)2.0 times as long as greatest 
diameter, with two tergal setae, both pointed, proximal seta, 1 = 5 urn, glabrous, 0.3 of 
length of tarsus and 1 .7 times as long as distal striate seta, its length = 3 um. Cuticle of 
tarsi glabrous. No proximal seta on tarsus of leg 1. All legs with large main claw and 
small setose anterior secondary claw; in leg 9 the former reaching 0.7 of length of 
tarsus. A blunt glabrous appendage on anterior side of femur of leg 1 , its length = 3 urn. 



566 U. SCHELLER 

Pygidium (Fig. 150): Tergum. Posterior margin with two narrowly triangular, 
posteriorly directed jags protruding from level of setae a 2 , small semicircular lobe pro- 
truding backwards above anal plate. Setae glabrous, flj and a 2 cylindrical, blunt, the 
former protruding backward and the latter converging, a 3 tapering, curved inward, 
converging, st somewhat davate, directed posteriorly. Lengths of setae: a^ = a 2 = 5, a 3 
= st = (6-)7 um. Distance aj -a^ =9(-10) urn, a 2 -a 2 = (25-)27 urn, 03-03 = (30- 

)32(-33) urn, a^ - a 2 - 6 urn, a 2 - 03 = (3-)4 pm, st - st = (6-)7 urn. Distance ay - a^ 
1.8(-2.0) times as long as aj, distance öj - a 2 1.5(-2.0) times as long as distance a 2 - 
a 3 , distance st - st as long as st and 0.7 of distance a^ - aj. Cuticle glabrous. 

Sternum. Posterior margin between b\ with shallow indentation. Setae thin, 
tapering, striate, their lengths: b x = 24(-26) pm, b 2 = 14(-15) urn, b 3 = (15-)16 pm. 
Distance b 1 -b l = 22(-23) urn, b 2 -b 2 = (37-)40(-41) pm, b 3 -b 3 = (15-)16 urn, b x - 
b 2 = 16 pm, b 2 - b 3 = 13(-14) pm. b x 0.9 of interdistance, b 2 about as long as distance 
bi - b 2 , b 3 as long as interdistance. Sternum glabrous. 

Anal plate 1.3 times as long as broad, broadest in the middle, lateral margins 
convex anteriorly, concave posteriorly; distal part of plate cleft by a U-shaped incision, 
depth about 1/3 of length of plate, incision forming two posterior branches with sub- 
paralleli sides. Each of the posterior branches with a davate appendage protruding 
backward, length of appendage 0.3 of length of plate. Plate glabrous, distal appendages 
with somewhat granular surface. 

26. Samarangopus sarawakensis sp. n. Figs 151-162 

Type material: Holotype: ad 9( 9 ), Malaysia, Sarawak, Kuching-Matang road, Gunung 
Serapi, soil sample (extraction at Kuching) from forest along road to the TV-station, alt. 670 m, 
9.XII.1987 (loc. Sar-87/64, leg. Hauser). Paratypes: 8 ad. 9(3 0* ,5 9), 1 juv. 5, same data as for 
holotype. -Altogether 10 specimens. 

Etymology: A latinized adjective of the name Sarawak. 

Diagnosis: S. sarawakensis sp. n. is most closely related to S. ternarius Serieller 
from Sabah (Serieller, 2001). They are similar, e. g. in the shape of the antennae, the 
protuberances of the tergites and the appendages of the collum segment. The following 
characters are useful for distinguishing these two species: the shape of the collum 
process (triangular and without anterior incision in S. sarawakensis sp. n., broadly 
rounded with anterior incision in S. ternarius), the shape of the genital papillae (distal 
part rounded without processes in S. sarawakensis sp. n., with three processes around 
the seta in S. ternarius), the distal part of bothriotricha T 3 (rounded distally in S. 
sarawakensis sp. n., cut squarely distally in S. ternarius), the shape of the posterior 
margin of the median subtergal plate (with point in S. sarawakensis sp. n., rounded in 
S. ternarius) and the setae b 2 of the pygidial sternum (thin, tapering in S. sarawaken- 
sis sp. n., lanceolate and with wings in S. ternarius). 

Description: Length = (0.64-)1.07 mm. 

Antennae (Fig. 151): Glabrous; segment 4 with 3 tapering, indistinctly striate 
setae, their lengths (for holotype only): p = 18 pm, p' = 20 pm, p" 10 pm. Sternal 
branch s, anterior margin = 14 pm, posterior margin =18 urn, of base = 7 pm, 
maximum = 10(-11) urn, q thin, tapering, indistinctly striate, 1 = 25 pm. Posterior 



RECORDS OF PAUROPODA 



567 




Figs 151-162 
Samarangopus sarawakensis sp. n., holotype, ad. 9(9). (151) right antenna, sternal view; (152) 
collum segment, median and left part, sternal view; (153) tergite I, anterior margin, sternal view; 
(154) tergite I, left posterolateral corner, sternal view; (155) tergite II, posterior margin; (156) 
tergite II, fungiform structure, lateral view; (157) T 3 ; (158) genital papillae, anterior view; (159) 
seta on coxa of leg 9; (160) tarsus of leg 9; (161) tarsus of leg 1; (162) pygidium, median and 
left part, sternal view. Scale a: Figs 152, 155; 158-161; b: Figs 151, 153-154, 157; c: Figs 156, 
162. 



568 U. SCHELLER 

margin/length of g (2.3-)2.6, posterior margin/maximum = (1.8-)1.9, maximum 0/0 
of base = 1.4(-1.6). Tergal branch subcylindrical, 1 = (25-)27 urn, of base = 5 pm, 
maximum = 8(-9) urn; pore not ascertained; length of //maximum = (2.8-)3.4. 
Globulus g, 1 = 7(-8) urn, maximum = 5 urn; length/maximum = 1.4(-1.6); number 
of bracts (7-)8, their length = 4 urn, capsule subspherical, bottom somewhat flattened, 
= 4 urn. Relative lengths of flagella (base segments included) and base segments: F x 
= 100, bs x = (14-)15, F 2 = (43-)46, bs 2 = (7-)8, F 3 = (89-) 100, bs 3 = (15-)16. Fj 
3.3(-3.5) times as long as tergal branch t, F 2 and F 3 (2.1-)2.2 times as long as and 
(0.8-)0.9 of length of sternal branch s, respectively. Calyces of Fj largest, conical, those 
of F 2 and F 3 , small, hemispherical. 

Trunk: Submedian setae of collum segment (Fig. 152) furcate, main branch 
tapering, striate, secondary branch rudimentary, pointed, glabrous, submedian seta 
1=13 um, sublateral seta 1=15 pm. Sternite process triangular, anteriorly rounded, no 
anterior incision. Appendages cylindrical, caps flat, somewhat larger than distal part of 
appendages. Process and appendages glabrous. 

Tergites: Tergite I with a single marginal row of campanulate, glabrous protu- 
berances (Figs 153-154); protuberances of anterior and lateral margins with tergal side 
somewhat triangularly lengthened, protuberances of anterior margin with median out- 
ward directed spine and protuberances of lateral margins with lengthened part folio- 
form, pointed, turned downwards distally. Number of marginal protuberances: I, 
(27-)31; II, 1 small -T x -\ small + (8-)9; III, 1 small + 5(-6) - T 2 - 1 small + 6; IV, 1 
small + 6 - T 3 - (4-)5(-6); V, (6-)8 - T 4 - 4; VI, 6 - T 5 - 1. Cuticular surface coarse, 
near margins and especially on posterior parts of tergites with many fungiform 
structures (Figs 155-156); the latter with the top covered by a small transparent cover 
in the shape of a mushroom hat or of an umbrella. 

Length/width ratio of tergites: I = 0.5(-0.6), II = 0.6, III = (4-)0.5, IV = V = 
VI = 0.5. 

Bothriotricha: All but T 3 with very thin glabrous axes. T 3 (Fig. 157) with thicker 
axes and distal endswelling with weak pubescence. Relative lengths of bothriotricha 
(for holotype only): T x = 100, T 2 = 105, T 3 = 55, T 4 = ?, T 5 = 126. 

Genital papillae (Fig. 158): (As long as -)1.2 times as long as greatest width. 
Base segments almost half of length of papillae. Length of papillae = 20 pm, greatest 
= 17(-20) pm, length of seta = 35(-38) pm. Papilla slowly tapering, rounded distally, 
opening of inserting cavity of seta circular, seta thick, (1.9-)2.1 times as long as length 
of papilla. Cuticle glabrous. Coxal seta of leg 2 as on leg 1, length = 20 pm. 

Legs: All legs 5-segmented. Seta on coxa (Fig. 159) and trochanter of leg 9 thin, 
furcate, striate, with glabrous base; length of secondary branch 0.4 of primary one. 
More anterior setae with rudimentary pointed glabrous secondary branches. Tarsi, 
tapering, those of leg 9 (Fig. 160) 1.6(-1.7) times as long as greatest diameter, with two 
tergal setae, both pointed glabrous. Proximal seta 9(-10) pm, distal one 4(-5) pm. 
Proximal setae (0.2-)0.3 of length of tarsus and 2.0(-2 2) times as long as distal seta. 
Cuticle of tarsus glabrous. No proximal seta on tarsus of leg 1 (Fig. 161). All legs with 
large main claw and small setose anterior secondary claw; in leg 9 the former reaching 
0.6 of length of tarsus. A triangular glabrous appendage on anterior side of femur of 
leg 1 , its length = 5 urn. 



RECORDS OF PAUROPODA 569 

Pygidium (Fig. 162): Tergum. Posterior margin with two narrowly triangular, 
pointed jags protruding backward from between setae a 2 and a 3 \ a shield-like lobe, 
posteriorly with triangular point, protruding backward above anal plate. Setae 
glabrous, a x and a 2 cylindrical, blunt, somewhat curved inward, a 3 straight, tapering, 
pointed, diverging; st cylindrical in proximal 1/4, more outward lanceolate. Lengths of 
setae: a x = (5-)6 urn, a 2 = (4-)6 urn, a 3 = (14-)17 urn, st = (12-)14 urn. Distance a x - 
a x = (10-)1 1 um, a 2 -a 2 = (25-)27(-28) um, a 3 -a 3 = (38-)39(-42) um, a l -a 2 = 9 um, 
a 2 -a 3 = (10-)1 1(-12) urn, st - st= 11(-13) urn. Distance a x -a x 1.8(-2.0) times as long 
as fl ls distance a x - a 2 1.3 times as long as distance a 2 - a 3 ; distance st - st 0.8(-1.0) 
of length of st and 1.1 (-1.2) times as long as distance a x - a x . Cuticle glabrous except 
for two small areas outside st. 

Sternum. Posterior margin between b\ with low and broad bulge. Setae b x lost, 
b 2 tapering, pointed, distal half with very short pubescence, b 3 thin, tapering, pointed, 
glabrous. Lengths of setae: b 2 = (25-)27(-30) urn, b 3 - (14-)15 urn. Distance b ì -b ì = 
(34-)36 urn, b 2 - b 2 »60 um, b 3 -b 3 = 22(-28) um, b l -b 2 = 25(-29) um, b 2 - b 3 «20 
urn. b 2 (as long as -)1.1 times as long as distance b± - b 2 , b 3 (0.5-)0.6 of interdistance. 

Anal plate 1.4(-1.5) times as long as wide, broadest at level of lateral 
appendages, these thin, cylindrical, blunt, glabrous, protruding obliquely backward; 
appendages 0.3 of length of plate; posterior half of plate divided into two tapering 
branches by a V-shaped incision, each branch provided with an almost glabrous stalked 
bladder, in sternal view similar to a knife-blade. These appendages 0.8 of length of 
plate. Plate and sternum glabrous. 

Sphaeropauropodinae 

Genus Sphaeropauropus Silvestri, 1930. 

27. Sphaeropauropus arcuatus Serieller 

Sphaeropauropus arcuatus Serieller, 2001: 981-984, figs 105-117. 

Material examined: Indonesia, Java, Cibodas, Lithocarpus-Castanopsis forest above 
the Botanical Garden, tourist trail to the waterfall, soil sample (extraction at Kuching, Sarawak) 
from between buttresses of large trees, alt. 1380 m, lad. 9(c?), 26.XI.1987 (loc. Sar-87/21, leg. 
Hauser). - Malaysia, Sarawak, Kuching-Matang road, Gunung Serapi, soil sample (extraction at 
Kuching) from forest along road to the TV-station, alt. 670 m, 3 ad. 9(9), 9.XII.1987 (loc. Sar- 
87/64, leg. B. Hauser); road Kuching-Serian, 18 km from Kuching, near Kampong Kuap, 
secondary forest, soil sample (extraction in Geneva) from between buttresses of a large tree, alt. 
30 m, 1 ad. 9(sex?), 1 juv. 6, 13.XII.1987 (loc. Sar-87/86, leg. B. Hauser). - Altogether 6 
specimens. 

General distribution: The species is here reported for the first time from Java. 
It was previously known from Sabah only (Serieller, 2001). 

REMARKS 

Pauropods have been almost unknown from the Indo-Australian area. 
Disregarding from Sabah in East Malaysia, only a few accidentally found species have 
been reported from Java (one species, Remy, 1933), Mindanao, the Palau Islands and 
Guam (five species, Remy, 1957d) and Papua New Guinea (one species, Serieller, 
1996). The collections from Sabah, 99 specimens in all (Scheller et al, 1994; Scheller, 
2001), contain 18 identified species, only three of which distributed elsewhere. 



570 U. SCHELLER 

The collection studied here, with 104 specimens from Singapore, Java, Bali and 
Sarawak, is the first one of importance from the Indonesian island arch and is one of 
the largest collections brought together from the Indo- Australian region. In comparison 
to the number of specimens, the number of species, 27, is considerable. Many of them, 
18 (67%), are new to science and only nine were known earlier from other areas. Part 
of the material comes from botanical gardens and the like, but these figures are similar 
to those earlier presented for New Caledonia (Serieller, 1993). 127 specimens from 
there were studied and 18 new species were identified (82% of total number of species 
found), only four widespread species could be found. A similar picture appears when 
comparing the faunas of north-western Thailand, 54 specimens, 10 new species (71% 
of total number of species found) (Serieller, 1995), south Vietnam, 45 specimens, 7 new 
species (87% of total number of species found) (Serieller, 2004) and Sabah, 99 
specimens, 14 new species (78% of total number of species found) (Scheller et al, 
1994; Scheller, 2001). 

Nine of the studied species studied above have been met with before. Four of 
them were known from Sabah: Hemipauropus dispar Serieller, Brachypauropoides 
penanorum Scheller, Samarangopus longipenes Serieller and Sphaeropauropus 
arcuatus Scheller. The three former can now be included in the fauna of Sarawak too, 
and the last mentioned also occurs on Java. They may all belong to an old Indo- 
Australian faunal element. Of the five remaining species described earlier, 
Allopauropus manjakotompensis Remy & Bello and A. pumilio Remy have been 
reported from Madagascar and Réunion, respectively, and may be a part of an old 
Gondwanan fauna. This means that only three of the species in this material have 
ranges extending outside the Oriental region: Allopauropus proximus Remy, A. pulcher 
Remy and A. mortensenii (Hansen). The first mentioned is widely distributed in the 
tropics but the whole distributional range of the other two is currently not known. A. 
pulcher was described from South Africa and has later been found in the USA, and 
records of A. mortensenii also show a very discontinuous picture including Egypt, Sri 
Lanka, Réunion, Mauritius, Koh Chang in the Gulf of Thailand and New Caledonia. 
There is also a doubtful rcord from Australia (Harrison, 1914). Accordingly only three 
species of 27 reported here have ranges extending outside the Oriental region. The wide 
range element is poor. 

The high portion of new species and the presence of only a few widespread 
species indicate a strong endemism in the Southeast Asian pauropod fauna. All 
available collections indicate a poorly known fauna with a high species diversity. 

ACKNOWLEDGEMENTS 

I am indebted to the Drs Bernd Hauser, Charles Lienhard and Pierre Strinati for 
sending me the material, which formed the basis of this study. 

REFERENCES 

Attems, C. 1938. Die von Dr. C. Dawydoff in französisch Indochina gesammelten Myriopoden. 

Mémoires du Muséum nationale d'Histoire naturelle (Paris) (N.S.) 6(2): 187-353. 
Hansen, H.J. 1902. On the genera and species of the order Pauropoda. Videnskabelige Meddel- 

elserfra Dansk Naturhistorisk Forening i Kj0benha\m 1901: 323-424, plates 1-6. 



RECORDS OF PAUROPODA 571 



Harrison, L. 1914. On some Pauropoda from New South Wales. Proceedings of the Linnean 

Society of New South Wales 39: 615-634, plates 70-71. 
Remy, P.A. 1933. Un Pauropode nouveau d'Indochine: Pauropus Dawydojfi n. sp. Faune des 

Colonies françaises (Paris) 5(3): 190-193. 

Remy, P.A. 1945. Nouvelle contribution à la faune des Myriapodes de Corse. Mémoires du 

Muséum national d'Histoire naturelle (Paris) (N.S.)21(4): 1 17-152. 
Remy, P.A. 1948a. Pauropodes de l'Afrique orientale britannique. Proceedings of the Zoological 

Society of London 118: 568-574. 
Remy, P.A. 1948b. Pauropodes de la Côte d'Ivoire, Afrique occidentale française. Mémoires du 

Muséum national d'Histoire naturelle (Paris) (N.S.) 7(5): 1 15-151. 
Remy, P.A. 1950. Pauropodes de Basse Egypte (récoltés de B. Condé). Bulletin de l'Institut 

d'Egypte 32: 35-48. 
Remy, P.A. 1952. Pauropodes de Nouvelle-Zélande. Record of the Canterbury Museum Christ- 
church) 6(2): 167-179. 
Remy, P.A. 1956a. Pauropodes de Madagascar. Mémoires de l'Institut scientifique de 

Madagascar (A)10: 101-229. 
Remy, P.A. 1956b. Pauropodes d'Afrique du Sud. Bulletin du Muséum national d'Histoire 

naturelle (Paris) (2) 28(3): 296-299. 
Remy, P.A. 1956c. Allopauropus pulcher, nouveau Pauropode d'Afrique du Sud. Annals of the 

Natal Natal Museum 13(3): 445-446. 
Remy, P.A. 1956d. Pauropodes des États-Unis d'Amérique. Mémoires de la Société nationale des 

Sciences naturelles et mathématiques de Cherbourg 47: 1-48. 
Remy, P.A. 1956e. Pauropodes de l'île de la Réunion. Mémoires de l'Institut scientifique de 

Madagascar (A)ll: 127-166. 
Remy, P.A. 1957a. Pauropodes des États-Unis (Tennessee, North Carolina). Revue française 

d'Entomologie 24(1): 82-87. 
Remy, P.A. 1957b. Description d'un nouveau Pauropode de Nouvelle-Calédonie. Bulletin du 

Muséum national d'Histoire naturelle (Paris) (2) 28(6): 519-523. 
Remy, P.A. 1957c. Additions à la faune pauropodienne de Côte d'Ivoire (récoltes de Stéphane et 

Blandine Bricka). Bulletin de l'Institut française d'Afrique noire (A)19(3): 751-762. 
Remy, P.A. 1957d. Pauropoda. In: Bernice P. Bishop Museum (ed.). Insects of Micronesia 

(Honolulu) 4(1): 1-12. 
Remy, P.A. 1959. Pauropodes de l'île Maurice. The Mauritius Institute Bulletin 5: 149-194. 
Remy, P.A. 1961. Pauropodes de la région de Pondichéry (Inde). Mémoires de la Société 

nationale des Sciences naturelles et mathématiques de Cherbourg (5)9: 1-24. 
Remy, P.A. 1962. Quelques Pauropodes de Ceylan. Izdanija. Zavoda za ribarstvo na N.R. 

Makedonija 3(2): 131-136. 
Remy, P.A. & Bello, J. 1960. Pauropodes du massif de l'Ankaratra (Madagascar). Mémoires de 

l'Institut scientifique de Madagascar (A)14: 71-93. 
Scheller, U. 1970. The Pauropoda of Ceylon. Entomologica scandinavica, Supplementum 1: 

5-97. 

Scheller, U. 1988. The Pauropoda (Myriapoda) of the Savannah River Plant, Aiken, South 
Carolina. Savannah River Plant and National Environmental Research Park Program 
(SRO-NERP) 17: 1-99. 

Scheller, U. 1993. Pauropoda (Myriapoda) from New Caledonia, (pp. 25-71). In: Matile, L., 
Najt, J. & S. Tillier (eds). Zoologia neocaledonica 3(34). Mémoires du Muséum na- 
tional d'Histoire naturelle (Paris) 157. 

Scheller, U. 1995. Pauropoda (Pauropodidae Eurypauropodidae) from north-western Thailand. 
Tropical Zoology 8(1): 7-41. 

Scheller, U. 1996. A new species of Samarangopus (Myriapoda: Pauropoda) from Papua New 
Guinea. Records of the Western Australian Museum 18: 87-91. 



572 U. SCHELLER 



Scheller, U. 2001. Pauropoda (Myriapoda) from Sabah (East Malaysia) (Pauropoda and 

Symphyla of the Geneva Museum XL) Revue suisse de Zoologie 108: 949-986. 
Scheller, U. 2004. Pauropoda (Myriapoda) from Vietnam (Pauropoda and Symphyla of the 

Geneva Museum XIII.) Revue suisse de Zoologie 111: 77-101. 
Scheller, U., Brtnck, P. & Enckell, PH. 1994. First record of Pauropoda (Myriapoda) on 

Borneo. Stobaeana (Lund) 1: 1-14. 
Scheller, U. & Bernard, E.C. 2005. New species of Pauropoda. (Pauropodidae, Brachy- 

pauropodidae) from Great Smoky Mountains National Park (Tennessee, North Carolina, 

USA), and a list of known species. Proceedings of the Biological Society of Washington 

118(4): 815-854. 
Silvestri, F. 1902. Ordo Pauropoda. In: Berlese, A. (ed.). Acari, Myriopoda et Scorpiones 

hucusque in Italia reperta. Portici. 
Silvestri, P. 1930. Descrizione di due nuovi Pauropodi (Myriapoda) di Giava. Bollettino del 

Laboratorio di Zoologia generale e agraria della Facoltà agraria in Portici 23: 

227-231. 
Verhoeff, K.W. 1934. Symphyla und Pauropoda. Pauropoda. (pp. 120-200). In: Bronns, H.G. 

(ed.). Klassen und Ordnungen des Tierreichs, 5 Arthropoda, 2 Myriapoda, 3 Symphyla 

und Pauropoda, 2 Pauropoda. Akademische Verlagsgesellschaft, Leipzig- 



Revue suisse de Zoologie 1 14 (3): 573-590; septembre 2007 



New species of larupea Martinez and morphological specializations 
among related taxa associated with ants and termites 
(Coleoptera: Scarabaeidae: Eupariini) 

Zdzislawa T. STEBNICKA 

Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, 

Slawkowska 17, 31-016 Krakow, Poland. Email: stebnicka@isez.pan.krakow.pl 

New species of larupea Martinez and morphological specializations 
among related taxa associated with ants and termites (Coleoptera: 
Scarabaeidae: Eupariini). - larupea Martinez is one of the South 
American, myrmecophilous genera of Eupariini with five species, including 
two species from Brazil described herein as new: /. goias sp. n. and /. luisae 
sp. n. The genus is redefined, a key for species, distribution, habitus photo- 
graphs and illustrations are given. Adaptive modifications in external 
morphology of larupea and of 14 related myrmecophilous genera are sum- 
marized. Of the total number of 33 species known to occur with social 
insects, most numerous species are recorded in association with leaf-cutting 
ants Atta F. (18 species) and Acromyrmex Mayr (5 species); the second 
group includes species associated with fire ants Solenopsis Westw. (7 

species) and Iridomyrmex Mayr (1 species) (Formicidae, Myrmicinae). Two 
highly derived species are recorded from termitaria of unknown hosts. 
Known or suspected behavioural aspects among the guests and hosts are dis- 
cussed. 

Keywords: Coleoptera, - Eupariini - larupea - new species - myrme- 
cophilous genera - morphological specializations - New World. 

INTRODUCTION 

Myrmecophily or myrmecophoby is wide-spread among Coleoptera and has, in 
some groups, led to remarkable modifications in the external morphology. 
Myrmecophilous tendencies among the Neotropical Scarabaeinae and Coprinae were 
documented in the literature several times (Vaz-de-Mello et al, 1998) while only few 
studies focused on the association of the Eupariini with social insects (Woodruff & 
Cartwright, 1967; Wojcik et al, 1977, 1978). This paper represents an initial attempt 
to bring together all available information on the New World myrmecophilous 
Eupariini and their morphological specializations. larupea Martinez (1952) is a small 
genus with five South American species, including two new species from Brazil 
described herein. This originally monospecific genus was diagnosed by Stebnicka 
(1999a) who added to it two distinctive species transferred from Euparia Le Peletier 
Saint Fargeau & Serville. The discovery of the following two species of larupea and 



Manuscript accepted 01.05.2007 



574 z - T - STEBNICKA 

examination of additional material enabled to emphasize the most striking morpho- 
logical modifications in external morphology of 15 genera containing 33 myrme- 
cophilous and termitophilous species. Members of the genera presented in Table 1 are 
poorly represented in the majority of the entomological collections, almost certainly 
due to their inquiline habits. Of the total number of 33 species, 23 species are asso- 
ciated with Attini, 8 species with Solenopsidini (Formicidae, Myrmicinae) and two 
species are recorded from termitaria. 

MATERIALS AND METHODS 

The nomenclature used in this paper to describe external features follows that 
of Chapin (1940), Cartwright (1974) and somewhat modified terminology of Stebnicka 
&Howden(1996, 1997). 

This study is based on the primary types of all known species and on material 
collected in South America and submitted for identification. The type material and 
other specimens studied may be found in the following repositories: Canadian Museum 
of Nature, Ottawa, Canada (CMNO); Florida State Collection of Arthropods, 
Gainesville, USA (FSCA); Hungarian Natural History Museum, Budapest, Hungary 
(HNHM); Institute of Systematics & Evolution of Animals, Krakow, Poland (ISEA); 
Muséum d'histoire naturelle, Geneva, Switzerland (MHNG); Muséum National 
d'Histoire Naturelle, Paris, France (MNHN); Staatliches Museum für Tierkunde, 
Dresden, Germany (SMTD); National Museum of Natural History, Smithsonian 
Institution, Washington DC, USA (NMNH); Zoologisches Museum für Naturkunde 
der Humboldt Universität, Berlin, Germany (ZMHB). 

The most numerous specimens, particularly those from Antonio Martinez col- 
lection are deposited in the Canadian Museum of Nature, Ottawa (CMNO). 

TAXONOMY 

Genus Iarupea Martinez, 1953 

Iarupea Martinez, 1953: 75-77.- Stebnicka 1999a: 290-291. 
Euparia Le Peletier de Saint Fargeau & Serville, 1828: 357 (in part) 
Iarupoides Chalumeau & Howden, 1984: 87 (nomen nudum). 

Type species: Iarupea lopeteguii Martinez, by monotypy. 

Diagnosis: morphological modifications. Head with coarse longitudinal 
wrinkles, genae sharply carinate in front of eyes. Pronotum furrowed basally with wide 
marginal collar and postero-lateral cavities to receive elytral lobe, anterior angles 
concave and slightly transparent. Elytra with humeral angles prolonged anteriorly into 
dentiform lobes (Fig. 6), apices of lobes with microtrichomes. Legs long; middle and 
hind tibiae slender, very narrow at base and weakly expanded apically. 

Redescription: length 4.5- 5.0 mm. Body (Figs 1, 3-5) elongate, alutaceous, 
usually covered with argillaceous coating. Head moderate in size, convex medially, 
genae right-angled. Eyes invisible from above; antennae 9-segmented, club ovoid, 3- 
segmented; mouthparts adapted to soft saprophagy. Pronotum strongly convex on disc, 
sides narrowly explanate and sinuate to obtuse posterior angles, side margin with 
fringe of short, very close, club-shaped setae. Scutellum subtriangular, depressed. 



NEW SPECIES OF IARUPEA 575 

Elytra narrower than pronotum, base margined and swollen, usually with large 
tubercles terminating intervals 3, 5, 7; intervals convex, crenate by strial punctures; epi- 
pleura oblique, reduced anteriorly, narrowed toward apex. Metathoracic wings well de- 
veloped. Prosternum with triangular process; mesosternum declivous toward metaster- 
num; mesocoxae separated, space between mesocoxae equal to width of mesofemur, 
meso-metasternal carina inconspicuous or lacking; metasternum long, convex; abdom- 
inal sternites fused; pygidium large, disc eroded. Profemur wide, anterior margin 
fringed with row of truncate setae, posterior margin widely flattened; middle and hind 
femora long, parallel-sided, without postfemoral lines; fore tibiae narrow, relatively 
short, third lateral tooth very small; middle and hind tibiae slender, apical spurs thin, 
slightly arcuate; tarsi shorter than tibiae, tarsomeres slender, claws fine, hornlike. 
Epipharynx specifically not differentiated. 

Male: terminal spur of fore tibia bent inwardly at the tip; penultimate abdo- 
minal sternite shorter and less arcuate than in female; genitalia specifically weakly 
differentiated, aedeagus elongate, parameres slender, as long as phallobase or slightly 
longer (Figs 9, 10). 

Female: pronotum more convex than in male; terminal spur of fore tibia 
straight; penultimate abdominal sternite longer than in male and slightly arcuate 
medially. 

Distribution: South America (Fig. 7); includes all previously published 
localities. 

Remarks: the unique combination of autapomorphic character states distin- 
guishes Iarupea from all other genera of Eupariini. The genus is most close to Arupaia 
Stebnicka, sharing with that genus a similar overall appearance, sculpture of the head 
and pronotum and the elytra with humeral lobes. Iarupea-species are known to occur 
with leaf-cutting ants of the genus Atta F. (Table 1). 

KEY TO SPECIES OF IARUPEA 

1 Pronotum with fine, uniformly distributed granules (Fig. 6), basal collar 

smooth; elytral intervals 3, 5, 7 carinate basally. Brazil goias sp. n. 

Pronotum with coarse pits or irregular wrinkles, basal collar longitu- 
dinally strigose; elytral intervals 3, 5 tuberculate or carinate basally 2 

2 Pronotum with coarse, longitudinally confluent wrinkles (Fig. 2); elytral 
striae with punctures strongly crenating intervals on each side. Bolivia, 
Brazil, Paraguay serratipennis (Petrovitz) 

_ Pronotum with coarse punctures or pits; elytral striae with punctures 

crenating inner margins of intervals 3 

3 Base of pronotum on each side of collar with prominent tubercle, preno- 
tai surface with coarse, rough, nearly contiguous pits; lateral intervals of 
elytra finely closely punctate. Argentina, Bolivia, Paraguay . lopeteguii Martinez 
Base of pronotum on each side of collar slightly convex without 
prominent tubercle, pronotal surface differently sculptured; lateral 
intervals of elytra impunctate 4 



576 Z. T. STEBNICKA 

4 Pronotai punctures or pits round or slightly elongate, separated by about 
one their diameter; elytral intervals 5, 7 carinate basally. Brazil 
attenuata (Harold) 

_ Prenotai punctures or pits elongate, nearly contiguous, tending to coale- 
sce into longitudinal lines; elytral intervals 3, 5 with tubercle basally. 
Brazil luisae sp. n. 

Iarupea lopeteguii Martinez Figs 7, 10 

larupea lopeteguii Martinez, 1953: 77-80, fig. 9; Stebnicka 1999a: 292, fig. 5. 

Material examined: Holotype and 4 paratypes (Argentina, Prov. Formosa, Puerto 
Irigoyen, Rio Pilcomayo) in CMNO. Other specimens (28): ARGENTINA - (14 ex) Prov. Salta, 
Dpto Gen. Ballivian, 1927, leg. Harrington; (2 ex) Dpto Gen. San Martin, Politos, XI. 1950; 
(1 ex) Dpto Anta Las Lajitas, XII. 1984; (2 ex) NS Telavera, XI. 1957, coll. Martinez (CMNO, 
MHNG); (1 ex) Dpto Tartagal, 12-19.XII.1990, M. Archangelsky (ISEA, NMNH). BOLIVIA - 
(1 ex) Villa Montes at Rio Pilcomayo, 1-29.XI.1930, S.G. Eisenstraut (ZMHB); (4 ex) Santa 
Cruz, P. Cordillera, Cabezas, 11.1971, coll. Martinez (CMNO). PARAGUAY - (3 ex) Dpto 
Boqueron, Gran Chaco, XI. 1956, coll. Martinez (CMNO). 

Diagnostic characters: colour dark reddish brown; clypeal margin slightly re- 
flexed, truncate anteriorly; surface of head with longitudinal wrinkles broken into 
irregular segments and with extremely short setae visible under high magnification. 
Pronotum convex medially, basal collar longitudinally strigose with sharp tubercle on 
each side; surface roughly sculptured, with large, very close, sharply edged pits bearing 
minute setae. Elytra parallel-sided, humeral lobes moderately long; striae impressed, 
strial punctures transversely crenate margins of intervals; intervals slightly convex, 
strongly microreticulate and minutely punctured, punctures on lateral intervals very 
close, 5 th interval at base with large, conical tubercle. Ventral sclerites alutaceous, 
minutely punctate, covered with extremely short, upright setae; abdominal sternites 
1-4 finely fluted along sutures, sternite 5 with longer and coarser fluting; eroded disc 
of pygidium longitudinally wrinkled, margin upturned. Profemur wide, perimarginal 
groove lacking, surface finely granulate and setigerous. Basal tarsomere of hind tarsus 
slightly longer than upper spur of tibia and longer than following three tarsomeres 
combined. 

Remarks: the species is most closely related to Iarupea serratipennis but may 
be easily recognized by the characters given in the key. Several specimens were 
collected to black light traps and found in the nest gallery of Atta vollenweìderi Forel 
(Martinez, 1953). 

Iarupea serratipennis (Petrovitz) Figs 1-2, 7, 8-9 

Euparia serratipennis Petrovitz, 1973: 185-186. 
lEuparia serratipennis: Chalumeau & Howden, 1984: 88. 
Iarupea serratipennis: Stebnicka 1999a: 292, fig. 6. 

Material examined: Holotype (Brazil, Minas Gerais) in MHNG. Other specimens 
(106): ARGENTINA - (2 ex) Prov. Misiones, Iguaçu, XII. 1957, coll. Martinez (CMNO). 
BOLIVIA - (1 ex) Prov. Sara, coll. Steinbach (ZMHB); (1 ex) Prov. Santa Cruz, XII. 1971, 
IX. 1972, F. Plaumann (CMNO); (2 ex) Guayaramirim (Beni), 23. XI. 1966, Hungarian Zool. Soil 
Exp, leg. Balogh & Mahunka & Zicsi (HNHM, ISEA). BRAZIL - (3 ex) (MS) Mato Grosso do 
Sul, Selviria, UNESP Farm, 18.m.l999, C. Hechtmann; (2 ex) (Ro) Rondonia, 62 km SW 
Ariquemes, near Faz. Rancho Grande, 25.LX.1992, U. Schmitz (FSCA); (1 ex) (Ma) Maranhào, 



NEW SPECIES OF IARUPEA 



511 




Figs 1-6 
(1) Iarupea serratipennis (Petrovitz): habitus of female; (2) same, sculpture of pronotum; (3) I. 
attenuata (Harold): habitus of male; (4) /. luisae sp. n: habitus of paratype female; (5) /. goias 
sp. n.: habitus of holotype female; (6) same, detail of pronotum and elytra. 



Pedrinhas, 26.VI.1984, C. Flechtmann (ISEA); (1 ex) (MT) Mato Grosso, Xingu, XL 1963; (1 
ex) (ES) Espiritu Santo, Linhares, XL 1962, coll. Martinez (CMNO); (90 ex) (Sc) Santa Catarina, 
Nova Teutonia, XII. 1972, F Plaumann (CMNO). PARAGUAY - (1 ex) Puerto P. Stroessner 
[=Ciudad del Este], 5-6.1.1966, leg. Mahunka (MHNG); (1 ex) Villarica, 25 km E Independentia, 
21.1.1991, S. Endrödi-Younga (ISEA). 



578 z - T - STEBNICKA 

Diagnostic characters: body (Fig. 1) rusty brown to dark brown, weakly 
shining. Clypeal margin broadly rounded on each side of shallow median emargina- 
tion; surface of head with coarse, contiguous longitudinal wrinkles. Pronotum convex 
medially, basal collar longitudinally strigose, anterior angles concave and slightly 
transparent; surface strongly swollen with irregular coarse wrinkles (Fig. 2). Elytra par- 
allel-sided, humeral lobes moderately long; striae impressed, strial punctures coarse, 
transversely crenate margins of intervals; intervals slightly convex or flat, intervals 3, 
5 with tubercle basally, surface minutely punctate. Abdominal sternites equally, dis- 
tinctly fluted along sutures; eroded disc of pygidium longitudinally strigose, margin 
upturned. Profemur wide, perimarginal groove fine, surface finely granulate. Basal tar- 
somere of hind tarsus longer than upper spur of tibia and longer than following three 
tarsomeres combined. Epipharynx as in Fig. 8. Male genitalia as in Fig. 9. 

Remarks: the species shows an advanced variation in the proportions and sculp- 
ture of the body. It is closest to larupea lopeteguii, but differs from that species by its 
more robust body and sculpture of the pronotum. Even though /. serratipennis is rep- 
resented by the greatest number of specimens, most of them were collected to black 
light traps; several specimens were taken from the nests of Atta sexdens (L.). 

larupea attenuata (Harold) Figs 3, 7 

Euparia attenuata Harold, 1870: 23-28; Schmidt 1922: 397. 
Iarupoides attenuatus: Chalumeau & Howden, 1984: 87. 
larupea attenuata: Stebnicka, 1999a: 291-292, fig. 11. 

Material examined: Lectotype "Brazil" (designated by Cartwright, 1973) in MNHN. 
Other specimens (17): BRAZIL - (2 ex) (Pe) Pernambuco (SMTD); (9 ex) (Pa) Para, 
Jacareacanga, X.1959, coll. Martinez (CMNO, MHNG); (2 ex) (MG) Minas Gerais, 
Cordisburgo, Faz. Pontinha, VII. 1974, F. Vaz-de-Mello (ISEA); (3 ex) (Sc) Santa Catarina, Nova 
Teutonia, X.1963, F Plaumann; (1 ex) (ES) Espiritu Santo, Linhares, X.1968, M. Alvarenga 
(NMNH). 

Diagnostic characters: body (Fig. 3) piceous, in some specimens with slight 
blue lustre, weakly shining. Clypeal margin broadly rounded on each side of shallow 
median emargination; surface of head with coarse, contiguous longitudinal wrinkles. 
Pronotum convex medially, basal collar longitudinally strigose; surface with irregularly 
spaced, coarse, circular or slightly elongate punctures or pits separated by one to three 
times their diameters. Elytra parallel-sided, humeral lobes moderately long, ended by 
microtrichomes; striae impressed, strial punctures coarse, transversely crenate only in- 
ner margins of intervals; intervals slightly convex or flat, minutely setigerous apically, 
intervals 5, 7 carinate basally. Abdominal sternites equally, finely fluted along sutures, 
eroded disc of pygidium longitudinally strigose, margin upturned. Profemur wide, per- 
imarginal groove fine, surface finely granulate. Basal tarsomere of hind tarsus longer 
than upper tibial spur and longer than following three tarsomeres combined. 

Remarks: the species is most closely related to larupea luisae sp. n. (see 
Remarks under that species). As indicated on the labels, some specimens of /. attenu- 
ata were taken from detritus cavities of the nests of Atta sexdens (L.). 



NEW SPECIES OF IARUPEA 579 



Iarupea luisae sp. n. Figs 4, 7 

Type material: Holotype female, Brazil, (Ma) Maranhäo, Pedrinhas, Saö Luis Island, 
29. VIII. 1999, black light trap in mangrove area, E.C. Bergmann, in ISEA. Paratype female, same 
locality as holotype, "ex Atta sp. dump", in ISEA. 

Description of females: length 4.8-5.0 mm. Body (Fig. 4) piceous to black, 
weakly shining. Clypeal margin broadly rounded on each side of shallow median emar- 
gination; surface of head from anterior margin to frons with coarse, contiguous longi- 
tudinal wrinkles blending into vertical band of round, close punctures separated by 
about one their diameter. Pronotum convex medially, diverging posteriorly, anterior 
angles slightly transparent, sides widely grooved and upturned, fringed with very close, 
short, club-shaped setae, basal collar longitudinally strigose; prenotai surface 
excluding lateral groove with coarse punctures or pits tending to coalesce into longi- 
tudinal lines. Elytra parallel- sided, humeral lobes rather long, ended by micro- 
trichomes; striae impressed, strial punctures coarse, transversely crenate margins of 
intervals; intervals slightly convex on disc, deplanate laterally, intervals 3, 5 with 
tubercle basally, surface microreticulate, impunctate. Ventral surface alutaceous; 
prosternai process widely triangular; mesosternum lower than metasternum with 
shallow, fine punctures separated by about one their diameter, posteriorly punctures 
scattered toward metasternum; meso-metasternal carina lacking; metasternum convex, 
midline impressed, discal punctures minute, scattered, lateral metasternal triangle 
broad, shallow, lateral area opaque, impunctate; abdominal sternites glabrous, 
impunctate with extremely fine fluting along sutures; pygidium broad, disc eroded, 
longitudinally strigose with upturned margin. Profemur with perimarginal groove and 
fringe of truncate setae, surface everywhere finely granular; middle and hind femora 
long, parallel-sided, smooth; middle and hind tibiae as long as femora, very slender and 
thin at base, apex fringed with 6-7 short setae, apical spurs slender, arcuate; basal 
tarsomere of hind tarsus subequal in length to upper spur of tibia and longer than 
following three tarsomeres combined. 

Male: unknown. 

Remarks: Iarupea luisae is most similar to /. attenuata from which it differs by 
having the pronotum broader with coarser and closer pits forming longitudinal lines. 

Etymology: named after the type-locality. 

Iarupea goias sp. n. Figs 5-6, 7 

Type material: Holotype female: Brazil, (Go) Goias, Mun. Bela Vista de Goias, 
Cristianopolis, Fazenda Arapuca Velha, 21. IX. 1993, Atta sp. nest gallery, A. Bankovics, in 
HNHM. 

Description of female: length 5.0 mm. Body (Fig. 5) castaneous, glabrous, 
weakly shining. Clypeal margin broadly rounded on each side of shallow median emar- 
gination; clypeal surface along anterior margin slightly concave and finely punctate, 
median convexity to frons with coarse longitudinal wrinkles blending into vertical 
band of round, fine, close punctures separated by less than one their diameter. 
Pronotum convex medially, diverging posteriorly, anterior angles slightly transparent, 
sides widely grooved and upturned, fringed with very close, short, club-shaped setae, 



580 



Z. T. STEBNICKA 



^=S^>j^^°"« 





* © \ 

*/ 
* ( 




A© 


© x J 




^0^— ^ 


O) 




* 


"~\ * / 




• r 


"V© / 




\ 3X 


tß\ d 


Dt attenuata 




^o? 


y m goias 






• ìopeteguii 






g liiisae 






serratipennis / 







Fig. 7 
Distribution of larupea species 

basal collar wide, smooth like as lateral groove, pronotai surface finely granulate, 
granules separated by one their diameter or nearly so. Elytra slightly arcuate, humeral 
lobes rather long (Fig. 6) ended by microtrichomes; striae impressed, strial punctures 
coarse, transversely crenate margins of intervals; intervals slightly convex on disc, 
deplanate laterally, intervals 3, 5, 7 obtusely carinate basally, surface microreticulate, 
impunctate. Ventral surface alutaceous; prosternai process triangular; mesosternum 
lower than metasternum, finely punctate in anterior third, punctures posteriorly become 
longitudinal, shallow, tending to coalesce into lines; meso-metasternal carina lacking; 
metasternum convex, midline impressed, discal punctures minute, scattered, lateral 
metasternal triangle broad, shallow, lateral area opaque, impunctate; abdominal 
sternites glabrous, impunctate with extremely fine fluting along sutures; pygidium 
broad, disc eroded, longitudinally strigose with upturned margin. Profemur with 
perimarginal groove and fringe of truncate setae, surface finely punctate; middle and 
hind femora long, parallel-sided, smooth; middle and hind tibiae as long as femora, 
very slender and thin at base, apex fringed with 5-6 short setae, apical spurs slender, 



NEW SPECIES OF IARUPEA 



581 













8 






10 



Figs 8-10 
(8) Iarupea serratipennis (Petrovitz): epipharynx; (9) same, male genitalia in lateral view; (10) 
/. lopeteguii Martinez: male genitalia in lateral view. 



arcuate; basal tarsomere of hind tarsus longer than upper tibial spur and equal in length 
to following three tarsomeres combined. 

Male: unknown. 

Remarks: I decided to describe this species on the basis of a single specimen, 
because its remarkably distinctive features are diagnostic. Iarupea goias sp. n. differs 
from all other species in the genus by its unusual sculpture of the pronotum (Fig. 6). 

Etymology: named after the type-locality. 



RELATED GENERA AND THEIR MORPHOLOGICAL SPECIALIZATIONS 



Euparia Le Peletier de Saint Fargeau & Serville, 1828: 357, et Auctt. 
Howden 1984: 85-87. 



Chalumeau & 



Distribution: Southern United States, Central America, Venezuela, West 
Indies. 

The numerous, world-wide species originally assigned to this genus have been 
transferred to the other existing or newly described genera (Stebnicka & Howden, 
1996). Euparia includes presently three species (Chalumeau & Howden, 1984) 
characterized by following morphological modifications: head broad; clypeal and pro- 
thoracic indentations forming a cavity to receive appendages and fore legs; pronotum 
with sides explanate, setaceous; elytra with humeral angles prolonged anteriorly into 
dentiform lobe; middle and hind legs long, femora longer than tibiae; tibiae thin, 
narrowed at base. Hosts are known for two species: 

Euparia castanea Le Peletier de Saint Fargeau & Serville, 1828 and E. mirabilis 
(Balthasar, 1945). These species are sympatric, very similar externally and difficult to 



582 Z. T. STEBNICKA 

distinguish. Most likely both species were studied in USA by Wojcik et al. (1977) and 
taken from excavated nests of tropical fire ants Solenopsis geminata F. and southern fire 
ants S. xyloni McCook. 

Arupaia Stebnicka, 1999a: 289. 

Distribution: Brazil. 

Monospecific genus, closely related to Euparia and to Iarupea, sharing with 
both these genera some morphological modifications: head broad, surface longitu- 
dinally wrinkled; presence of clypeal and prothoracic cavities to receive appendages 
and fore legs; pronotum with basal collar and explanate sides; elytra with humeral 
angles prolonged anteriorly into dentiform lobe; middle and hind legs long and thin. 

Arupaia friedenreichi (Harold, 1870) (Brazil) - collected in the nest of 
Solenopsis geminata F. 

Selviria Stebnicka, 1999a: 287.- 2005a: 23-25. 

Distribution: Brazil. 

The genus includes two species and seems to be most closely allied to Euparia. 
The unusual morphological modifications are: cuticle orange, strongly shining, 
smooth; head broad; sides of pronotum very widely explanate with upturned margin; 
elytral striae very fine; elytral margin narrowly explanate and upturned; mesosternum 
flattened, mesocoxae separate; legs very long, middle and hind tibiae thin, tarsal 
segments thick. Host is known for one species: 

Selviria matogrossoensis Stebnicka, 1999a (Brazil) - one specimen found in the 
nest of Solenopsis invida Buren. 

Euparixia Brown, 1927: 288.- Woodruff & Cartwright, 1967: 6; Gordon & McCleve 
2003: 685-686. 

Distribution: USA (Arizona, Louisiana), Cuba, Guatemala, Mexico, Panama. 

The genus includes presently 10 species with following morphological modi- 
fications: clypeal margin inflexed; pronotum strongly constricted basally with sides 
explanate; elytral epipleurae covering episternum, epimeron and apices of elongated 
and widely separated mesocoxae; legs long, slender. Euparixia is most closely related 
to Euparixoides Hinton containing five cryptic species of unusual appearance 
(Stebnicka & Skelley 2005), however, nothing is known on their hosts associations. Six 
species have been recorded in association with ants: 

Euparixia duncani Brown, 1927 (Arizona) - most likely associated with 
Acromyrmex versicolor Pergande and/or Atta mexicana Smith. 

Euparixia moseri Woodruff & Cartwright, 1967 (Louisiana) - specimens taken 
from nest excavation (fungus garden central cavity and detritus cavity) of Atta texana 
(Buck). 

Euparixia bruneri Chapin, 1940 (Cuba) - collected in the nest of Atta insularis 
Guérin known only from Cuba. 

Euparixia formica Hinton, 1934 (Mexico).- collected in the nest of Atta mexi- 
cana Smith. 



NEW SPECIES OF 1ARUPEA 583 

Euparixia costaricensis Hinton, 1936 (Costa Rica).- species associated with 
Atta colombica Guér. and Atta cephalotes (L.). 

Euparixia campbelli Gordon & McCleve, 2003 (Guatemala) - collected in large 
detritus cavity of the nest of Atta sp. 

Lomanoxia Martinez, 1951: 23-29.- Krikken, 1972: 70; Stebnicka, 1999c: 2; Skelley 
& Howden, 2003: 186. 

Distribution: Argentina, Brazil, Bolivia, Paraguay, Surinam, Costa Rica, 
Trinidad. 

The genus includes five species with following morphological modifications: 
body egg-shaped; prenotai and elytral margins with row of long setae; elytra flattened 
and sharply inflexed at 8th interval forming a broad false epipleural fold, surface with 
prominent setation and with scale-like setae; mesocoxae widely separated; mesepi- 
sternum, mesepimeron and elongated mesocoxae hidden; legs long. Hosts are known 
for three species. 

Lomanoxia costulata (Harold, 1867) (Brazil) - collected in Surinam from 
detritus cavities of the excavated nest of Atta sexdens (L.). 

Lomanoxia alternata Krikken, 1972 (Surinam) - collected from detritus cavities 
of the excavated nest of Atta cephalotes (L.). 

Lomanoxia canthonopsis Skelley & Howden, 2003 (Costa Rica) - found in 
refuse piles of Atta cephalotes (L.). 

Flechtmanniella Stebnicka, 1999b: 287. 

Distribution: Brazil. 

Monospecific genus closely related to Lomanoxia, sharing with that genus some 
morphological modifications: body shape oval; presence of clypeal and prothoracic 
indentations to receive appendages and fore legs; elytra flattened and sharply inflexed 
at 8th interval forming a broad false epipleural fold; mesocoxae widely separated; 
middle and hind tibiae short, expanded apically, tarsal segments short, thick. 

Flechtmanniella laticollis (Petrovitz, 1973) (Brazil) - found in the nest of 
fungus growing ants Acromyrmex lobicornis (Emery). 

Lomanoxoides Stebnicka, 1999a: 293-294.- Stebnicka & Skelley, 2005: 29. 

Distribution: Argentina, Bolivia, Brazil, Paraguay, Peru, Panama, Honduras. 

The genus includes seven species having most of the morphological characters 
unmodified like as in the other free-living taxa of the tribe: body oblong oval, convex; 
pronotum usually with swellings; elytral intervals subcarinate or carinate at middle 
with row of fine setae. Hosts are known for two species: 

Lomanoxoides nigrolineatus (Hinton, 1938) (Panama) - found in debris piles of 
Atta sp. and in refuse of Atta colombica Guér. 

Lomanoxoides tesari (Balthasar, 1963) (Paraguay) - found in the nests of Atta 
sexdens (L.). 

Paraplesiataenius Chalumeau, 1992: 194.- Stebnicka 2003a: 444. 
Distribution: Southeastern Brazil, Uruguay, Argentina. 



584 Z. T. STEBNICKA 

The genus is closely related to Lomanoxoides and includes three Aphodius-ap- 
pearing species with slightly modified morphological characters: body oblong oval, 
convex, shining; prenotai and elytral margins with row of setae; lateral elytral intervals 
at apex with short scarce setae; hind tibiae distinctly expanded apically. Hosts are 
known for all species of the genus: 

Paraplesiatcienius tremolerasi (Schmidt, 1911) (Argentina) - collected in 
Argentina in the nest of Acromyrmex lobicornis (Emery), in Uruguay found together 
with A. lundi Sant. 

Paraplesiataenius catarinaensis Stebnicka, 2003a (Brazil) - collected in the 
nest of Acromyrmex sp. 

Paraplesiataenius genieri Stebnicka, 2003a (Brazil) - found together with 
Acromyrmex sp. 

Martineziella Chalumeau, 1986: 386. 

Distribution: Southern USA, Argentina, Paraguay, Uruguay, Bolivia, Brazil, 
Guiana, Mexico, Cuba, Dominican Republic. 

The genus includes six species having most of the morphological characters un- 
modified like as in the free-living species of the tribe: body elongate, convex; prono- 
tum with sides explanate, fringed with very close, truncate setae; legs long. Hosts have 
been recorded for two species: 

Martineziella vandykei (Hinton, 1936) (Mexico) - paratypes were taken from 
the nests of "a small, red, bitting ants" (Hinton, 1936); other specimens collected in de- 
tritus remnants of the nests of Solenopsis invicta Buren. 

Martineziella dutertrei (Chalumeau, 1983) (Cuba) = Myrmecaphodius excava- 
ticollis (non Blanchard, 1843): Woodruff, 1973: 101-102, fig. 212; Wojcik et al, 1977: 
329-334. 

This species is widely distributed from southern USA to Argentina and occa- 
sionally very abundant. Collins & Markin (1971) reported this species from 98% of the 
fire ant nests they examined in the United States and they observed in the laboratory 
that the beetles in proximity to ants released a strong musky odour. A complete life cy- 
cle of the beetles apparently occurs in the nests where their preimaginal stages also 
have been found. M. dutertrei is known to occur in numerous nests of Solenopsis 
invicta Buren (red fire ant), S. richten Forel (black fire ant), S. geminata F. (tropical fire 
ant), S. xyloni McCook (southern fire ant) and Iridomyrmex humilis (Mayr) (argentine 
ant). 

Myrmecaphodius Martinez. 1952: 85.- Stebnicka 1999a: 292. 

Distribution: Argentina. 

Monospecific genus related to Martineziella. Its morphological modifications 
are as follows: head coarsely longitudinally wrinkled; pronotum strongly convex, sides 
explanate in anterior half, basal margin strong, wide at middle with longitudinal 
costulae, prenotai surface with deep, large pits. 

Myrmecaphodius proseni Martinez, 1952 (Argentina).- Stebnicka 1999a: 293 - 
type series (29 specimens) and other individuals examined were collected in the nest 
gallery of fire ants Solenopsis saevissima Smith. 



NEW SPECIES OF IARUPEA 585 

Haroldiataenius Chalumeau, 1981: 137. 

Distribution: Southern United States, Mexico, Guatemala, Honduras. 

About seven species are assigned to this genus (revision in preparation by 
Stebnicka). The morphological characters reveal a rather weak modifications: body 
oval, strongly convex; abdominal sternites and pygidium with row of blunt setae. Host 
is known for two species: 

Haroldiataenius hintoni (Saylor, 1933) (Mexico) - collected in Guatemala in 
detritus remnants of the nest of Atta mexicana Smith. 

Haroldiataenius limbatus (Bates, 1887) (Mexico) - found in Guatemala in "Atta 
sp. dump" (label data by J. Baster). 

Ataenius Harold, 1867: 82, et Auctt. 

Distribution: Pantropical. 

The genus contains presently about 190 species distributed throughout 
American continent and West Indies. The morphological characters of a number of 
species show some insignificant modifications, such as: head larger than usual; surface 
of body with rows of thick setae or cuticle coarsely, roughly sculptured (usually smooth 
or punctate); prenotai lateral margin fringed with close, truncate or club-shaped setae 
(usually setae are scarcer and slender); legs longer than usual; tarsal segments thick 
(usually slender). From among 190 species only two species are indicated in associa- 
tion with ants: 

Ataenius holopubescens Hinton, 1938 (Mexico) - collected from detritus 
remnants in the nests of Atta mexicana Smith. 

Ataenius variopunctatus Schmidt, 1922 (Argentina) - rare species known only 
from Buenos Aires. From among 6 specimens available to study one specimen was 
glued on card together with a specimen of ant and labeled "Solenopsis sp." by J. 
Daguerre. 

Batesiana Chalumeau, 1983: 143-144. 

Distribution: Panama, Costa Rica, Brazil. 

Monospecific genus with advanced morphological modifications: head very 
large; pronotum short and very broad; presence of clypeal and prothoracic indentations 
to receive appendages and fore legs; elytra strongly elevated medially, lateral and 
apical intervals with row of irregular tubercles (glands?); mesosternum deplanate, 
mesocoxae widely separated; fore tibiae short and narrow with two reduced lateral 
teeth; middle and hind tibiae short, strongly expanded apically, covered with dense 
hair; tarsi short and thick with long hair. 

Batesiana tuberculata (Bates, 1887) (Panama) - collected in Brazil with 
"termites". 

Napoa Stebnicka, 1999d: 290-291. 

Distribution: Ecuador. 

Monospecific genus closely related to Batesiana with advanced morphological 
specializations: head very large; presence of clypeal and prothoracic indentations to 



586 Z. T. STEBNICKA 

receive appendages and fore legs; elytra globular, lateral and apical intervals with row 
of irregular tubercles (glands?); mesosternum deplanate and calloused, mesocoxae 
widely separated; prosternum, mesosternum and mesocoxae with distinct pockets or 
cavities of possible mycangial function. 

Napoa peckorum Stebnicka, 1999d (Ecuador) - specimens found in broken 
termite nest. 

Remark: the genus Cartwrightia Cartwright, was recently assigned to the tribe 
Eupariini by Skelley & Howden (2005) on the basis of some character states shared 
with Lomanoxia. Apart from the general appearance of Cartwrightia strongly resem- 
bling various genera of Rhyparinae, the shape of head and characters of mouthparts of 
bitting type exclude this genus from the Eupariini. The bitting type of mouthparts is 
one of the principal diagnostic characters for Aegialiinae, Aulonocneminae and 
Rhyparinae as well. 

DISCUSSION 

The members of fifteen euparine genera here considered (Table 1) are charac- 
terized by more or less advanced morphological specializations. Although they have no 
distinct trichomes, exudatoria or exocrine glands attractive to the ants, their external 
features are frequently unusual. Some species have retained a generalized appearance 
and differ little from their free-living relatives, while others seem to be highly derived. 
These species, like as all other members of the tribe, have the mouthparts of filtering 
type (adapted for soft saprophagy) and they are unable to consume any hard particles 
of food. It means, that food preferences may include liquid or subliquid organic 
contents of specific enzymatic qualifications (Stebnicka, 1985). 

The morphological changes are always connected with function which a part of 
the body concerned has to perform. These modifications can be grouped as follows: 1/ 
rudimentation or disappearance of sclerites, 2/ displacement of sclerites, 3/ deve- 
lopment of new parts in preexisting sclerites, 4/ transformation of organs or their 
portions, 5/ abnormal (hypertelic) development of organs. It is unclear whether these 
characteristics are important to the integration of the guests into host colonies since the 
supporting behavioural information is not available. Species of Iarupea possess 
humeral microtrichomes, however, their function may be suspected only. Long legs 
with relatively thick tarsi probably allow guests to keep up with their hosts. The ability 
to pull appendages and fore legs into ventral cavities most certainly has a defensive 
significance and allows the beetles to survive occasional aggressive reactions by their 
hosts. The functions of other specializations, for example the presence of elytral 
swellings or tubercles (glands?) are not at all clear. 

The species of Eupariini listed in Table 1 have been observed in association 
with leaf-cutting (fungus growing) ants Atta Fabricius and Acromyrmex Mayr and with 
fire ants Solenopsis Westwood and Iridomyrmex Mayr (Myrmicinae). The distribution 
and biology of the gardening ants has been summarized by Weber (1972). Attini are 
widely distributed in America and have the largest colonies. Their fungus gardens 
consist mostly of vegetable material and insect fecal material to which the ants add 
saliva and liquid fecal droplets. The ants either cast out exhaused substrate and dead 
ants or store it in separate chambers of the nest, and these large volumes of decaying 



NEW SPECIES OF 1ARUPEA 



587 



Table 1: Eupariini species associated with termites and with ants Atta F., Acromynnex Mayr, 
Solenopsis Westw. and Iridomyrmex Mayr (Myrmicinae) 



Eupariini species 



Host species 



Locality 



References 



Iampea lopeteguii 

I. attenuata 

I. serratipennis 

I. goias sp. n. 

/. luisae sp. n. 

Euparia castanea/E. mirabilis 

Arupaia friedenreichi 

Selviria matogrossoensis 

Euparixia duncani 

E. moseri 

E. bruneri 

E. formica 

E. costaricensis 

E. campbelli 

Lomanoxia costulata 
L. alternata 
L. canthonopsis 

Flechtmanniella laticollis 

Lomanoxoides nigrolineatus 

L. te sari 

Paraplesiataenius tremolerasi 



P. catarinaensis 

P. genieri 
Martineziella vandykei 

Martineziella dutertrei 

(=Myrmecaphodius 

excavaticollis) 

Myrmecaphodius proseni 

Haroldiataenius hintoni 
H. limbatus 

Ataenius holopubescens 
A. variopunctatus 
Batesiana tuberculata 

Napoa peckorum 



Atta vollenw eider i 
nest gallery 
Atta sexdens 

Atta sexdens 

Atta sp. 
nest gallery 
Atta sp. dump, 
mangrove area 
Solenopsis xyloni, 
S. geminata 
Solenopsis geminata 

Solenopsis invicta 

Acromynnex versicolor 

Atta texana 

Atta insularìs 

Atta mexicana 

Atta colombica, 
A. cephalotes 
Atta sp. 

Atta sexdens 
Atta cephalotes 
Atta cephalotes 

Acromyrmex lobicornis 

Atta colombica 

Atta sexdens 
Acromyrmex lobicornis 
A. lundi 

Acromyrmex sp. 

Acromyrmex sp. 
Solenopsis invicta 

Solenopsis invicta, 

S. richten, S. geminata, 

S. xyloni, Iridomyrmex 

humilis 

Solenopsis saevissima 

Atta mexicana 
Atta sp. 
Atta mexicana 
Solenopsis sp. 
"termites" 

"broken termite nest" 



Argentina, 
Prov. Formosa 
Brazil, Minas 
Gerais 
Brazil, Mato 
Grosso do Sul 
Brazil, Goi'as 

Brazil, Saö Luis 
Island 
Southern USA 

Brazil, Santa 
Catarina 
Brazil, Goias 

USA (Arizona) 

USA (Louisiana) 

Cuba 

Mexico 

Costa Rica 

Guatemala 

Surinam 
Surinam 
Costa Rica 

Brazil, Sao Paulo 

Panama 

Paraguay 
Argentina 
Uruguay 

Brazil, Santa 
Catarina 
Brazil, Bahia 
Mexico 

USA (Alabama, 

Florida, Georgia, 

Louisiana, 

Mississippi, Texas) 

Argentina, 

Buenos Aires 

Guatemala 

Guatemala 

Mexico 

Argentina" 

Brazil 

Ecuador 



Martinez, 1953 

label data by 

Martinez 

label data by 

Steinbach 

label data by 

Bancovics 

label data by 

Bergmann 

Wojcik et al, 1977 

label data by 
Triplehorn 
label data by 
Morgante & Silva 
Woodruff & 
Cartwright, 1967 
Woodruff & 
Cartwright, 1967 
Woodruff & 
Cartwright, 1967 
Woodruff & 
Cartwright, 1967 
Woodruff & 
Cartwright, 1967 
Gordon & 
McCleve, 2003 
Krikken, 1972 
Krikken, 1972 
Skelley & Howden, 
2003 

label data by 
Martinez 
Stebnicka & 
Skelley, 2005 
Stebnicka, 1999a 
Chalumeau, 1992 
label data by 
Fernandez 
Stebnicka, 2003a 

Stebnicka, 2003a 

label data by 

Stinger 

Wojcik et al, 1977 



Martinez, 1952 

Saylor, 1933 
label data by Baster 
Stebnicka, 2003b 
Stebnicka, 2005 
label data by 
Degallier 
Stebnicka, 1999d 



588 z - T - STEBNICKA 

refuse may attract the beetles. The fire-ants Solenopsis form populous colonies usually 
in open soil localities, often on pastures under cow dung and often in close proximity 
to the nests of large ants from whom they steal brood and other food (Moody & Franke, 
1982). 

Judging from the rather scarce collection data, the beetles exploit either aban- 
doned mounds of ants and the active colonies. The small euparine beetles visiting or 
living in the colonies of ants are practically defenceless, in particular in the nests of 
Solenopsis, the workers of which are extremely aggressive and possess a painful sting. 
Kistner (1979) has identified a number of mechanisms that enable the integration of 
guests into host colonies, for example the guests may use chemical means and mimic 
chemical communication systems of the hosts. Collins & Markin (1971) recorded 
Martineziella dutertrei from 98% of the fire ant nests they examined in the United 
States and they observed in the laboratory that the beetles in nearness of ants released 
a strong musky odour. It seems most likely that either the ants and beetles may pro- 
duce, in some conditions, the integrative pheromones, for example, the beetles may be 
attracted by odorous secretion, a glandular substantion skatole/indole (Brown et al, 
1979; Keegans et al, 1993) used by hosts to mark their trails. The integrative mecha- 
nisms are complicated and very diffcult to explain, however, the most important factor 
which lures beetles to the nests of ants is that the large host colonies have a higher 
diversity of microhabitats and sources of food. The vast amounts of decomposing 
vegetable matter collected and discarded by ants constitute nutritionally richest 
resource for saprophagous beetles of Eupariini, therefore, their possible role of the 
nests cleaners may be tolerated or accepted by hosts. 

The development of fungus-gardens of both ants and termites was probably one 
of the factors which led to extensive invasion of their nests by various beetles in later 
times. 

ACKNOWLEDGEMENTS 

I am grateful to Henry F. Howden from the Canadian Museum of Nature for his 
invaluable cooperation as well as to the following persons for their assistance: 
F. Génier (Ottawa), P.E. Skelley (Florida), O. Merkl (Budapest), I. Lobi, G. Cuccodoro 
and B. Merz (Geneva), Y. Cambefort (Paris), D. Ahrens (Dresden), fN. Adams, D. 
Fürth and G. House (Washington), H. Wendt (Berlin). I wish to thank Marek Kopec 
from ISEA Krakow for making the photographs of beetles. 

REFERENCES 

Balthasar, V. 1945. De uno Genere et nonnullis Aphodiinarum speciebus novis. Casopis 

Ceskoslovenske Spolecnosti Entomologicke 42: 104-115. 
Balthasar, V. 1963. Eine neue Gattung, Untergattung und neue Arten der Familie Aphodiidae. 

Reichenbachia 1: 277-290. 
Bates, H.W. 1887. Col.:Lam. (Copridae, Aphodiidae, Orphnidae, Hybosoridae, Geotrupidae, 

Trogidae, Aclopidae, Chasmatopteridae, Melolonthidae). Biologia Centrali-Americana. 

Insecta 2: 26-160. 
Brown, CA., Watktns, J.F. & Eldridge, D.W. 1979. Repression of bacteria and fungi by the 

army ant secretion: skatole. Journal of the Kansas Entomological Society 52: 119-122. 
Brown, W.J. 1927. Two New North American Genera of the Tribe Eupariini. Canadian 

Entomologist 59: 288-290. 



NEW SPECIES OF I ARU PEA 589 



Cartwright, O. L. 1973. Additional lectotype designations in the Aphodiinae. The Coleopterists 

Bulletin 27: 41-43. 
Cartwright, O. L. 1974. Ataenius, Aphotaenius and Pseudataenius of the United States and 

Canada. Smithsonian Contributions to Zoology 154: 1-106. 
Chalumeau, F. 1981. Nouveau genre et sous-genres d'Eupariini d'Amérique tropicale. Nouvelle 

Revue d'Entomologie 11: 137-142. 
Chalumeau, F. 1983. Batesiana et Martinezia, nouveaux genres d'Eupariini du nouveau monde. 

Bulletin mensuel de la Société linnéenne de Lyon 52: 142-153. 
Chalumeau, F. 1986. Martineziella nomen novo pour Martinezia Chalumeau. Nouvelle Revue 

d'Entomologie 3: 386. 
Chalumeau, F. 1992. Eupariini du nouveau monde: un mise au point. l re Partie. Nouvelle Revue 

d'Entomologie 9: 189-206. 
Chalumeau, F. & Howden, H.F 1984. Le genre Euparia Saint-Fargeau & Serville, 1828. 

Mitteilungen der Schweizerischen Entomologischen Gesellschaft 57: 85-94. 
Chapin, E.A. 1940. A revision of the West Indian beetles of the Scarabaeid subfamily 

Aphodiinae. Proceedings of the United States national Museum 89: 1-41. 
Collins, H.L. & Markin, G.P. 1971. Inquilines and other arthropods collected from nests of the 

imported fire ant, Solenopsis saevissima richten. Annals of the entomological Society of 

America 64: 1376-1380. 
Gordon, R.D. & McCleve, S. 2003. Five new species of Euparixia Brown (Coleoptera: Scara- 

baeidae: Eupariinae), with a revised key to species. Proceedings of the entomological 

Society of Washington 105: 685-697. 
Harold, E. 1867. Diagnosen neuer Coprophagen. Coleopterologische Hefte 1: 76-83. 
Harold, E. 1870. Die Arten der Gattung Euparia. Coleopterologische Hefte 6: 19-30. 
Hinton, H. E. 1934. Two genera of Aphodiinae new to Mexico. The Pan-Pacific Entomologist 10: 

27-30. 
Hinton, H.E. 1936. A new species of Ataenius from Mexico. Revista de Entomologia 6: 

472-474. 
Hinton, H.E. 1938. New species of Neotropical Aphodiinae. Revista Entomologia 8: 122-129. 
Keegans, S. J., Billen, J., Morgan, E.D. & Gökeen, O.A. 1993. Volatile glandular secretions 

of three species of new world army ants: Eciton burchelli, Labidus coecus and Labidus 

praedator. Journal of Chemical Ecology 19: 2705-2719. 
Kistner, D.H. 1979. Social and evolutionary significance of social insects symbionts (pp. 340- 

413). In: Herman, H.R. (ed.). Social insects. Academic Press: New York & London. 
Krikken, J. 1972. Species of the South American genus Lomanoxia (Coleoptera: Aphodiidae). 

Studies on the fauna of Suriname and other Guyanas 13: 68-83. 
Le Peletier de Saint-Fargeau, A.L.M. & Audinet- Serville, J. G. A. 1828. Scarabé (pp. 346- 

382). In: Latreille, P. A. (ed.). Histoire Naturelle. Entomologie, ou Histoire Naturelle 

des Crustacés, des Arachnides et des Insectes. Encyclopédie Méthodique. Paris 

(Agasse) 10. 
Martinez, A. 1951. Scarabaeidae nuevos o poco conocidos. II. Publicaciones Misión de 

Estudios de Patologia regional argentina 22: 23-36. 
Martinez, A. 1952. Scarabaeidae nuevos o poco conocidos. III. Publicaciones Misión de 

Estudios de Patologìa regional argentina 23: 53-118. 
Martinez, A. 1953. Scarabaeidae nuevos o poco conocidos. IV. Publicaciones Misión de 

Estudios de Patologia regional argentina 24: 69-85. 
Moody, J.V. & Francke, O.F. 1982. The Ants (Hymenoptera, Formicidae) of Western Texas, 

Part I: Myrmicinae. Graduate Studies Texas Tech University 27: 1-80. 
Petrovitz, R. 1973. Neue Aphodiinae, Hybosorinae und Acanthocerinae aus der Neotropischen 

Region. Studia entomologica 16: 141-202. 
Saylor, L.W. 1933. Two new Scarabaeidae. Canadian Entomologist 65: 158-159. 
Schmidt, A. 1911. Eine Serie neuer Aphodiinen. Stettiner entomologische Zeitung 72: 1-51. 



590 Z. T. STEBNICKA 

Schmidt, A. 1922. Coleoptera, Aphodiinae. Das Tierreich, Vol. 45. Walter de Gruyter and Co., 
Berlin und Leipzig, 614 pp. 

Skelley, P.E. & Howden, H.F. 2003. A new species of Lomanoxia Martinez from Costa Rica 
(Coleoptera: Scarabaeidae: Aphodiinae). Insecta Mundi 17: 185-190. 

Stebnicka, Z. 1985. A new genus and species of Aulonocneminae from India with notes on 
comparative morphology (Coleoptera, Scarabaeidae). Revue suisse de Zoologie 93: 
649-658. 

Stebnicka, Z. 1999a. Neotropical Eupariini: New and little known genera and species 
(Coleoptera: Scarabaeoidea: Aphodiinae). Revue suisse de Zoologie 106: 285-300. 

Stebnicka, Z. 1999b. Flechtmanniella, a new name for Flechtmannia Stebnicka, 1999. Acta zo- 
ologica cracoviensia 42: 287. 

Stebnicka, Z. 1999c. Lomanoxia Martinez, 1951, and a new tribe Lomanoxiini with notes on 
comparative morphology (Coleoptera: Scarabaeoidea: Aphodiinae). Acta zoologica 
cracoviensia 42: 279-286. 

Stebnicka, Z. 1999d. A new genus and species of termitophilous Eupariini from Ecuador with 
checklist of the Neotropical genera (Coleoptera: Scarabaeoidea: Aphodiinae). Acta zoo- 
logica cracoviensia 42: 289-295. 

Stebnicka, Z. T. 2001. Aphodiinae (Insecta: Coleoptera: Scarabaeidae). In: Fauna of New 
Zealand vol. 42, 64 pp, Manaaki Whenua Press, Lincoln Canterbury. 

Stebnicka, Z. 2003a. The genus Paraplesiataenius Chalumeau, with descriptions of two new 
species from Brazil (Coleoptera: Scarabaeidae: Eupariini). The Coleopterists Bulletin 
57: 444-450. 

Stebnicka, Z. 2003b. The New World species of Ataenius Harold, 1867. III. Revision of the A. 
imbricatus-group sensu lato (Coleoptera: Scarabaeidae: Aphodiinae: Eupariini). Acta 
zoologica cracoviensia 46: 219-249. 

Stebnicka, Z. 2005a. A Second Species of Selviria Stebnicka from Brazil (Coleoptera: 
Scarabaeidae: Eupariini). The Coleopterists Bulletin 59: 23-25. 

Stebnicka, Z. 2005b. The New World species of Ataenius Harold, 1867. VI. Revision of the A. 
aequalis-platensis-group (Coleoptera: Scarabaeidae: Eupariini). Acta zoologica cra- 
coviensia 48B: 99-138. 

Stebnicka, Z., & Howden, H.F. 1996. Australian Genera and Species in the Tribes Odonto- 
lochini, Psammodiini, Rhyparini, Stereomerini and Part of the Eupariini (Coleoptera: 
Scarabaeoidea: Aphodiinae). Invertebrate Taxonomy 10: 97-170. 

Stebnicka, Z. & Skelley, P.E. 2005. Review of some New World aphodiine genera and de- 
scriptions of new species (Coleoptera: Scarabaeidae: Aphodiinae). Acta zoologica cra- 
coviensia 48B: 23-42. 

Vaz de Mello, F.Z., Louzada, J.N.C. & Schroeder, J.H. 1998. New Data and Comments on 
Scarabaeidae (Coleoptera: Scarabaeoidea) Associated with Attini (Hymenoptera, 
Formicidae). The Coleopterists Bulletin 52: 209-216. 

Weber, W.A. 1972. Gardening Ants. The Attines. Memoirs of the American Philosophical 
Society 92: 1-146. 

Wojcik, D.P, Banks, W.A. & Habeck, D.H. 1978. Fire ant myrmecophiles: flight periods of 
Myrmecaphodius excavaticollis (Blanchard) and Euparia castanea Serville (Col. 
Scarabaeidae). The Coleopterists Bulletin 32: 59-64. 

Woicik, D.P, Banks, W.A., Hicks, D.M. & Summerlin, J.W 1977. Fire ant myrmecophile: new 
hosts and distribution (A Myrmecaphodius excavaticollis (Blanchard) and Euparia cas- 
tanea Serville. The Coleopterists Bulletin 31: 329-334. 

Woodruff, R.E. 1973. The Scarab Beetles of Florida. Part.I. The Laparosticti. Subfamilies: 
Scarabaeinae, Aphodiinae, Hybosorinae, Ochofdaeinae, Geotrupinae, Acanthocerinae. 
Arthropods of Florida and Neighboring Land Areas 8: 1-220. 

Woodruff, R.E., Cartwright, O.L. 1967. A Review of the Genus Euparixia With Description 
of a New Species From Nests of Leaf-Cutting Ants in Louisiana (Coleoptera: 
Scarabaeidae). Proceedings of the United States National Museum 123: 1-21. 



Revue suisse de Zoologie 1 14 (3): 591-598; septembre 2007 



Hisonotus hungy sp. n. (Siluriformes, Loricariidae) a new species 
from arroyo Tirica, Misiones, Argentina 

M. de las Mercedes AZPELICUETA 1 , Adriana E. ALMIRÓN 1 , 

Jorge R. C ASCIUTTA 1 & Stefan KOERBER 2 

1 Division Zoologia Vertebrados, Facultad de Ciencias Naturales y Museo, Paseo del 

Bosque s/n, 1900 La Plata, Argentina. E-mail: azpeli@ museo.fcnym.unlp.edu. ar 
2 Friesenstr. 11, 45476 Muelheim, Germany. 

E-mail: pecescriollos@koerber-germany.de 



Hisonotus hungy sp. n. (Siluriformes, Loricariidae) a new species from 
arroyo Tirica, Misiones, Argentina. - Hisonotus hungy sp. n. is described 
from the arroyo Tirica, rio Parana basin in Misiones province, Argentina. 
Hisonotus hungy sp. n. is diagnosed by the following combination of 
characters: 20-22 lateral plates, 5 branched anal-fin rays, absence of pad on 
snout tip, blunt and deep snout, snout 45.5-50.9 (% in HL), eye 13.4-17.1 
(% in HL), predorsal unpaired plates absent; vent completely covered by 
two rows of large lateral platelets and one median series with irregular 
plates. 

Keywords: Freshwaters fish - loricariids - Hypoptopomatinae - Hisonotus - 
South America - rio Parana basin. 

INTRODUCTION 

The arroyo Urugua-i together with the rio Iguaztì are the main tributaries of the 
rio Parana in the province of Misiones, Argentina. Eight kilometers from the 
confluence with the rio Parana, a hydroelectric dam was built at the end of the eighties. 
Before the construction of the dam, several collecting trips had been done which 
resulted in the description of new species such as Oligosarcus menezesi Miquelarena 
& Protogino, 1996; Bryconamericus sylvicola Braga, 1988; Australoheros tembe 
(Casciotta, Gomez & Toresani, 1995); Gymnogeophagus che Casciotta, Gomez & 
Toresani, 2000, and Astyanax leonidas Azpelicueta, Casciotta & Almirón, 2002. 

The lake formed by the dam receives several streams; the arroyo Tirica is one 
of the smaller ones flowing into the lake from the south. 

The aim of this paper is to describe a new species of Hisonotus recently col- 
lected from the arroyo Tirica, arroyo Urugua-i basin. 

MATERIAL AND METHODS 

Specimens were cleared and counterstained following Taylor & Van Dyke 
(1985). Measurements were taken as straight line distances using digital calliper to the 
nearest 0.1 mm. Values of the holotype are indicated by an asterisk. Institutional 



Manuscript accepted 01.05.2007 



592 M - M - AZPELICUETA ET AL. 

abbreviations are as listed in Leviton et al. (1985) with the addition of Asociación 

Ictiológica, La Plata, Argentina (AI) and Zoologia Vertebrados, Facultad de Ciencias, 

Universidad de la Repüblica, Montevideo, Uruguay (ZVC-P). 

Comparative material examined (SL in mm): Hisonotus candombe Casciotta, 
Azpelicueta, Almirón & Litz, 2006: ZVC-P 5595, holotype, 29.9, Repüblica Oriental del 
Uruguay, Departamento Salto, rio Uruguay basin, arroyo Palomas. AI 177, 1 ex. (C&S) 29.7, sa- 
me collecting data. - Hisonotus charma Almirón, Azpelicueta, Casciotta & Litz 2006: ZVC-P 
5639, holotype, 50.7, Repüblica Oriental del Uruguay, Departamento Tacuarembó, rio Uruguay 
basin, Canada de Los Pena. AI 171, 3 ex., 21.0-32.2 (C&S), Repüblica Oriental del Uruguay, Rio 
de la Plata basin, arroyo Tropa Vieja, Departamento Canelones. - Hisonotus maculipinnis 
(Regan, 1912): AI 122, 1 ex., 27.5 (C&S), Argentina, Corrientes province, rio Parana, Ita Ibaté. 
AI 123, 5 ex., 23.4-27.0, Argentina, Corrientes province, rio Parana basin, Esteras del Iberâ, 
Rincón del Diablo, Laguna Yac are. - Hisonotus cf. laevior Cope, 1894: AI 178, 6 ex., 30.0-38.0, 
Brasil, Rio Grande do Sul, Sào Leopoldo, Rio lacuî basin, rio dos Sinos, 29°45'S - 51°10'W. - 
Hisonotus paulinus (Regan, 1908): BMNH 1907.7.6.9, holotype, Brasil, rio Piracicaba, Säo 
Paulo (examined by J. Maclaine, Natural History Museum, London). -Hisonotus sp. A, AI 120, 
1 ex., 23.3, Argentina, Misiones, rio Uruguay basin, arroyo Oveja Negra. - Hisonotus sp. B: 
MHNG 2408.025, 10 ex., 17.8-29.0, Paraguay, route 2, arroyo Pirayü. Hisonotus ringueleti 
Aquino, Schaefer & Miquelarena, 2001: AI 179, 1 ex., 36.4, Repüblica Oriental del Uruguay, 
Departamento Artigas, rio Uruguay basin, arroyo Lenguazo. - Hisonotus taimensis (Buckup, 
1981): AI 216, 7 ex., 38.6-55.0, Repüblica Oriental del Uruguay, Rocha, arroyo Las Conchas, 
34°24.05'S-54° 17.05 'W. - Epactionotus aky Azpelicueta, Almirón, Casciotta & Koerber, 2004: 
AI 124, holotype, 30.5, Argentina, Misiones province, rio Uruguay basin, arroyo Garibaldi. - 
Epactionotus yasi Almirón, Azpelicueta & Casciotta, 2004: MACN-ict 8649, holotype, 1 ex. 
32.0, Argentina, Misiones province, rio Iguazü basin, arroyo Lobo. 

RESULTS 

Hisonotus hungy sp. n. Fig 1, Table 1 

Holotype: MACN-ict 8860, 35.7 mm SL, Argentina, Province of Misiones, rio Parana 
basin, arroyo Tirica, affluent of arroyo Urugua-f (26°01'S - 55°22'W), coll: S. Koerber, 
R. Filiberto, J. O. Fernandez Santos, 5 January 2001. 

Paratypes: same collecting data as holotype: AI 189, 5 ex., (1 C&S) 31.5-37.6 mm SL; 
ZFMK 39472-75, 4 ex., 30.4-42.0 mm SL; MHNG 2664.79, 2 ex., 31.9-37.0 mm SL; ZSM 
33313, 2 ex., 33.3-36.7 mm SL. 

Diagnosis: Hisonotus hungy sp. n. is diagnosed by the following combination 
of characters: 20-22 lateral plates, 5 branched anal-fin rays, absence of pad on snout 
tip, blunt and deep snout, snout 45.5-50.9 (% in HL), eye 13.4-17.1 (% in HL), 
predorsal unpaired plates absent; vent completely covered by two rows of large lateral 
platelets and one median series with irregular plates. 

Description: Morphometries of holotype and 13 paratypes are presented in 
Table 1. Body elongated, head deep and short (Fig. 1). Greatest body depth at dorsal- 
fin origin. Head as wide as trunk. Dorsal profile of head convex from snout tip to 
dorsal-fin origin. Snout tip rounded in dorsal view, without enlarged odontodes. One 
pair of rostral median plates without notch. Several plates placed in anterior area of 
nares, leaving a narrow naked surface. Eyes placed dorsolaterally, horizontal eye 
diameter shorter than nare diameter. Iris diverticulum present, about half of pupil 
diameter. Three infraorbitals surrounding orbit, fourth infraorbital expanded ventrally. 
Margins and surface of lips covered with papillae. Maxillary barbels short. Jaw teeth 
bifid, tooth slender with major cusp expanded distally and a very minor one pointed. 



NEW HÌSONOTUS FROM ARGENTINA 



593 




MHHHMM 




WÊÊÊÊBKL 



Fig. 1 
Hisonotus hungy sp. n., holotype, 35.7 mm SL, Argentina, Province of Misiones, rio Parana 
basin, arroyo Tinca (26°01'S - 55°22'W), lateral, dorsal, and ventral views. 



Absence of accessory teeth on premaxilla and dentary. One series of teeth, 10-17 
(mode 12) on premaxilla and 8-15 (mode 11) on dentary. Pterotic-supracleithrum 
bearing large openings. Preopercular sensory canal directed towards pterotic-supra- 
cleithrum. 

Body covered by dermal plates except for a naked area around base of pelvic 
fins, lateral opening of swimbladder capsule, and area between pectoral girdle and 
lower lip. Posrostral and anterior rostral plates slightly reflected ventrally. Five lateral 
series of plates on trunk. Plates of dorsal series continuous; mid-dorsal series 



594 M. M. AZPELICUETA ETAL. 



Table 1. Morphometric data of the holotype and 13 paratypes of Hisonotus hungy sp. n. SD: 
standard deviation. 

Holotype Range Mean SD 

Standard length (mm) 35.7 30.4-42.3 

Percents of SL 

Predorsal distance 
Head length 
Cleithral width 
Dorsal-fin spine length 
Trunk length 
Pectoral-fin spine length 
First pelvic-fin ray length 
Abdominal length 
Caudal peduncle length 
Caudal peduncle depth 
Head depth 
Snout length 
Horizontal eye diameter 
Interorbital width 

Percents of HL 

Head depth 
Snout length 
Horizontal eye diameter 
Interorbital width 
Cleithral width 



incomplete, and continuous. Median series 20 (5), 21 (4), 22 (5*). Mid- ventral series 
incomplete and continuous; plates of ventral series continuous. Plates bearing lateral- 
line canal incomplete and discontinuous, anterior portion of median series with 4 (5), 
5 (7*), 6 (2) perforated plates; posterior portion with 7 (7*), 8 (3), 9 (1), 11(1), 12 (1), 
and 13 (1) perforated plates. First two lateral line plates small, second one placed on 
rib of sixth vertebra. Abdomen almost completely covered by plates, arranged in two 
lateral rows of two or three plates each and a median row with several plates irregularly 
arranged (Fig. 1). Three pairs of post anal plates, first one meeting at midline and last 
third separated by small external portion of first anal-fin proximal radial. Coracoid and 
cleithrum exposed ventrally, excluded very small arrector fossae area. Absence of 
unpaired predorsal plates. 

Odontodes covering head, trunk, and fin rays. Head and trunk odontodes 
uniformly distributed. Odontodes usually small on body, except for somewhat enlarged 
odontodes on ventral margin of snout, on pectoral and pelvic spines, and a tuft formed 
by a few odontodes in some specimens. Odontodes along anterior margin of snout bi- 
serially arranged, dorsad and ventrad series not separated by a naked area. 

Dorsal fin with one spine and 7 branched rays, its origin posterior to vertical 
through pelvic-fin origin. Dorsal fin moved posteriorly behind seventh vertebra. First 
dorsal-fin proximal radial articulated with eighth vertebra. Adipose fin absent. Pectoral 
fin with one spine and 6 branched rays, reaching nearly half of pelvic-fin length. Very 
large pectoral-fin axillary slit present Pelvic fin with one unbranched and 5 branched 



42.9 


42.2-46.8 


44.6 1.69 


32.8 


31.1-35.8 


33.8 1.62 


23.5 


22.5-26.9 


24.9 1.32 


22.4 


21.1-25.0 


23.0 1.29 


17.6 


15.1-18.4 


17.2 0.85 


24.6 


22.2-27.8 


25.2 1.66 


20.2 


15.4-20.5 


18.5 1.63 


19.6 


19.6-22.2 


20.9 0.92 


34.7 


32.0-39.2 


35.5 2.34 


12.0 


10.6-13.3 


12.1 0.71 


18.5 


16.3-19.4 


18.2 0.91 


15.1 


15.0-17.8 


16.5 0.95 


5.3 


4.7-5.7 


5.2 0.29 


13.4 


13.0-14.9 


14.0 0.55 


56.4 


50.5-56.9 


54.0 2.13 


46.2 


45.5-50.9 


48.8 1.90 


16.2 


13.4-17.1 


15.5 0.88 


41.0 


38.4-45.5 


41.4 2.21 


71.8 


70.9-77.6 


73.7 2.22 



NEW HISONOTUS FROM ARGENTINA 



595 




Fig. 2 

Hisonotus paulinus, holotype BMNH 1907.7.6, Brasil, Sào Paulo, rio Piracicaba, dorsal, lateral 
and ventral views. 



rays, reaching anal-fin origin in females and surpassing it only in males. Fleshy flaps 
on pelvic fins of males. Caudal fin with fourteen branched rays. 

Neural spine of seventh vertebra contacting nuchal plate partially. Neural arch 
of seventh vertebra without expansion. 

Color in life: Background brown-chocolate, with clear lines between snout 
and eyes, extending backward to postemporo-supracleithrum. Ventral margin of snout, 
area around pectoral and pelvic fin insertions yellowish. Two light dots on caudal fin, 
upper most and lower most caudal-fin rays light. Dorsal, pectoral, pelvic, and anal fins 
with dots forming bands. 

Color in alcohol: Ground color of dorsolateral body surface brownish, ventral 
surface of head and body pale brown. In some specimens a narrow light stripe from 
snout tip to anterior nare, continuing posteriorly, very faint until supraoccipital level. 
Ventrolateral margin of snout and head light brown. Pectoral, pelvic, dorsal, and anal 
fins pale brown with dots forming series of darker bands. Caudal fin dark brown 



596 M - M - AZPELICUETA ETAL. 

excluded two light triangular dots near half way. Both light dots placed on first three 
or five dorsal and ventral branched rays. Tip of lower and upper caudal lobes usually 
light. 

Sexual dimorphism: Pelvic-fin spines of males longer than that of females 
(17.7-20.5 vs. 15.4-17.1% SL; 4 females and 10 males). Distal tip of pelvic fins 
surpassing anal-fin origin in males. Males have fleshy flap on pelvic fins and triangular 
genital papilla. Females bear rounded and broad genital papilla. 

Etymology: The specific epithet hungy is a guarani word that means brown. A 
noun in apposition. 

Distribution and habitat: This species is only known from the arroyo Tirica, 
a tributary of the arroyo Urugua-i. The arroyo Tirica, at bridge of provincial road 237- 
km 29, is a stream with two different environments; one of them had the natural vege- 
tation of the region, whereas in the other one coniferous trees were introduced. 
Hisonotus hungy was collected in the portion of the stream where a plantation of small 
coniferous let the brook to be exposed completely to sunlight. Photographs of live 
specimens, the environment, and a map with the geographical distribution are provided 
by Koerber (in prep.) and Evers & Seidel (2005, sub H. cf. ringueleti). 

The following species were caught together with H. hungy from the arroyo 
Tirica: Astyanax eigenmanniorum, A. cf. fasciatus, A. cf. troya, Australoheros tembe, 
Bryconamericus iheringii, Characidium sp., Corydoras carlae, Gymnogeophagus che, 
Heptapterus mustelinus, Oligosarcus jenynsii, O. paranensis, and Schizodon nasutus. 

DISCUSSION 

The genus Hisonotus includes 16 nominal species (Aquino et ah, 2001 and 
Britski & Garavello, 2003) from different basins in Brasil, Argentina, and Uruguay. 
Recently, two new species, H. charma and H. candombe, have been described from no 
Uruguay basin (Almirón et al., 2006; Casciotta et al., 2006). 

The species described from the upper rio Parana basin are H. insperatus Britski 
& Garavello, 2003, H. depressicauda (Miranda Ribeiro, 1918), H. depressinotus 
(Miranda Ribeiro, 1918), H. paulinus (Regan, 1908), and H. francirochai (Ihering, 
1928). Hisonotus maculipinnis was recorded in the lower Parana basin, from "La 
Plata" without precise locality. 

Among the species of Hisonotus distributed in the Rio de la Plata basin, and 
Lagoa dos Patos system, H. hungy sp. n. differs from all of them - except H. paulinus - 
by the number of lateral plates (20-22 vs. 23-31). 

Hisonotus hungy also differs from H. ringueleti, H. charrua, and H. candombe 
by the absence of soft pad in the snout tip. The other species living in the basin is 
H. maculipinnis which has a large free area in the anterior margin of the snout that is 
not present in H. hungy and the profile of the snout strongly depressed whereas 
H. hungy has a blunt and deep snout. 

In spite of papers by Schaefer (1991, 1998) and Aquino et al. (2001), the genus 
Hisonotus has not been clearly defined phylogenetically. In addition, the tribe 
Otothyrini diagnosed by Schaefer (1991, 1998) is considered a paraphyletic group at 



NEW HISONOTUS FROM ARGENTINA 



597 




Fig. 3 
Hisonotus hungy sp. n. from arroyo Tirica in aquarium, specimen not preserved. 



present (Gauger & Buckup, 2005). Gauger & Buckup (2005) included some new osteo- 
logical characters to define the genera of Hypoptopomatinae. Considering those char- 
acters, H. hungy has the mesethmoid covered by prenasal plates, rostral plates wider 
than long, lateral ethmoid not exposed on dorsal surface of head, infraorbital canal en- 
tering in the neurocranium via sphenotic, supraoccipital without crest, pectoral girdle 
completely exposed, arrector fossae reduced to a small area close to midline, mid-dor- 
sal series with about 16 plates, dermal plates of thorax and abdomen large, regularly 
distributed, thorax and abdomen completely covered by dermal plates, first post anal 
plate meeting at midline, and neither traces of modifications nor unpaired platelets lo- 
cated in adipose-fin region. 

Eschmeyer (2006) indicated Argentina with doubts in the distribution of 
Hisonotus paulinus (Regan, 1908) described from rio Piracicaba, Säo Paulo, Brasil 
(Fig. 2). This citation is taken from Lopez et al. (2003) that recorded H. cf. paulinus 
from arroyo Tirica, Misiones. The specimens mentioned by those authors were 
collected by one of the present paper authors (SK) together with the specimens 
examined in this study. The holotype of H. paulinus has the same number of lateral 
plates, nonetheless it differs from Hisonotus hungy in having 4 branched anal-fin rays 
vs. 5 (we do not know if it is an anomalous specimen); the snout is long and depressed, 
38.2% vs. 30.0-32.3% of predorsal length (compare dorsal and lateral views of both 
species in the figures). 



598 M. M. AZPELICUETA ETAL. 



ACKNOWLEDGEMENTS 

The authors thank H. Britski (Museu de Zoologia, Säo Paulo, Brasil) and 
G. Garcia (Facultad de Ciencias, Montevideo, Uruguay) for gift of material, S. Fisch- 
Müller (Muséum d'histoire naturelle de Genève, Switzerland) and K. Busse 
(Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany) for loan of ma- 
terial under their care, J. Maclaine (Natural History Museum, London, United 
Kingdom) for photographs of and valuable information on H. paulinus, an anonymous 
reviewer for critical revision of the paper and comments, M. Rinas (Ministerio de 
Ecologia de Misiones) for collecting permission, the board and members of Gfl 
(Ichthyological Society, German language area) for partial support in the collection in 
arroyo Tirica, J. O. Fernandez Santos and R. Filiberto for help in the field, and I. Seidel 
for providing the photo of the specimen in aquarium. 

REFERENCES 

Almirón, A. E., Azpelicueta. M. de las M., Casciotta, J. R. & Lrrz, T. 2006. A new species 

of Hisonotus (Siluriformes, Loricariidae, Otothyrini) from the Repüblica Oriental del 

Uruguay. Revue suisse de Zoologie 113: 87-94. 
Aquino, A. E, Schaefer, S. A. & Miquelarena, A. M. 2001. A new species of Hisonotus 

(Siluriformes, Loricariidae) of the upper Rio Uruguay Basin. American Museum 

Novitates 3333: 1-12. 
Britskj, H. A. & Garavello, J. C. 2003. Hisonotus insperatus: New species, from the Upper rio 

Parana basin (Pisces: Ostariophysi: Loricariidae). Copeia 2003: 588-593. 
Casciotta, J. R., Azpelicueta, M. de las M., Almirón, A. E. & Lrrz, T. 2006. Hisonotus can- 

dombe, a new species from the rio Uruguay basin in the Repüblica Oriental del Uruguay 

(Siluriformes, Loricariidae, Otothyrini). Spixiana 29: 147-152. 
Eschmeyer, W. N. 2006. Catalog of fishes. Electronic version available at 

http://www.calacademy.org 
Evers, H. G. & Seidel, I. 2005. Catfish- Atlas Vol. 1. South American Catfishes of the families 

Loricariidae, Cetopsidae, Nematogenyidae, and Trichomycteridae. Mergus, Melle, 

Germany, 944 pp. 
Gauger, M. F. W. & Buckup, P. A. 2005. Two new species of Hypoptopomatinae from the rio 

Parafba do Sul basin, with comments on the monophyly of Parotocinclus and the 

Otothyrini (Siluriformes: Loricariidae). Neotropical Ichthyology 3: 509-518. 
Koerber, S. (in prep.) Colour, habitat and distribution of Hisonotus hungy. File available at 

http://www.pecescriollos.de/go/hisonotus-hungy 
Levtton, A. E., Gibbs Jr., R. H., Heal, E. & Dawson, C. E. 1985. Standards in herpetology and 

ichthyology: Part I. Standard symbolic codes for institutional resource collections in 

herpetology and ichthyology. Copeia 1985: 802-832. 
Lopez, H. L., Miquelarena, A. M. & Menni, R. C. 2003. Lista comentada de los peces conti- 
nentales de la Argentina. Probiota, Serie Tècnica y Didâctica 5, 85 pp. 
Schaefer, S. A. 1991. Phylogenetic analysis of the loricariid subfamily Hypoptopomatinae 

(Pisces: Siluroidei: Loricariidae), with comments on generic diagnoses and geographic 

distribution. Zoological Journal of the Linnean Society 102: 1-41. 
Schaefer, S. A. 1998. Conflict and resolution: impact of new taxa on phylogenetic studies of the 

Neotropical cascudinhos (Siluroidei: Loricariidae). (pp. 375-400). In: Malabarba, L. 

R., Reis, R. E., Vari, R. P., Lucena, Z. M. & Lucena, C. A. S. (eds). Phylogeny and 

classification of neotropical fishes. EDIPUCRS, Porto Alegre, 603 pp. 
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 1 14 (3): 599-614; septembre 2007 



Phylogenetic position, morphology and natural history of the 
Vietnamese water skink Tropidophorus noggei Ziegler, 
Vu & Bui, 2005 (Sauria: Scincidae) 

Thomas ZIEGLER 1 , Astrid HEIDRICH 12 , Andreas SCHMITZ 3 
& Wolfgang BÖHME 2 

1 AG Zoologischer Garten Köln, Riehler Straße 173, D-50735 Köln, Germany. 
E-mail: tziegler@zoo-koeln.de 

2 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 
D-531 13 Bonn, Germany. 

3 Muséum d'histoire naturelle, Department of Herpetology and Ichthyology, 
C.P. 6434, CH-121 1 Geneva 6, Switzerland. 



Phylogenetic position, morphology and natural history of the 
Vietnamese water skink Tropidophorus noggei Ziegler, Vu & Bui, 2005 
(Sauria: Scincidae). - The depressed-bodied water skink species 
Tropidophorus noggei was recently described from Phong Nha - Ke Bang 
National Park, a karst forest region in the Truong Son, central Vietnam. 
Subsequent field research at the type locality led to the finding of additional 
specimens which allowed to extend the knowledge about the species' 
morphology including hemipenis morphology. The molecular positioning of 
the species within Tropidophorus supports that the body depression 
recognized in several Indochinese taxa is likely to have occurred at least 
twice in parallel as an adaptation to saxicolous habitats. New insights into 
the natural history of Tropidophorus noggei are given, including habitat 
choice, abundance, activity and habits, as well as the species' feeding and 
reproductive ecology. 

Keywords: Sauria - Scincidae - Lygosominae - Tropidophorus - T noggei - 
mitochondrial DNA sequence data - systematics - morphology - natural 
history - Vietnam. 

INTRODUCTION 

Recently, Honda et al. (2005) presented a molecular phylogeny of the oriental 
lygosomine genus Tropidophorus Duméril & Bibron, 1839, including the recently 
described T. latiscutatus, T. matsuii and T. murphy i (Hikida et al, 2002). These water 
skink species exclusively share distinctly depressed bodies, as well as saxicolous 
habitats. Hikida et al. (2005) assumed that the strongly depressed body shape reflects 
an adaptation to narrow rock crevices as shelters, as known also from few other lizard 
families, such as Cordylidae Gekkonidae, Iguanidae, Lacertidae, and Xenosauridae 



Manuscript accepted 22.06.2007 



600 T - ZIEGLER ET AL. 

(Vitt, 1981; Doughty & Shine, 1995; Ballinger et al, 2000). Subsequently, the analyses 
of Honda et al. (2005) showed that the characteristic body shape has independently 
evolved in T. murphyi and in the common ancestor of T. latiscutatus and T. matsuii. 
However, another depressed-bodied Tropidophorus species was not included in Honda 
et fl/.'s (2005) molecular approach: T noggei, which was described recently from a 
karst forest region in the Truong Son mountain range, Vietnam (Ziegler et ai, 2005). It 
is the aim of this study to clarify the phylogenetic position of this further depressed- 
bodied Tropidophorus by molecular means as well as to add new data to the 
morphology and natural history of the barely known species. 

MATERIAL AND METHODS 

Specimens collected by local people subsequently to the original description of 
T. noggei (Ziegler et al, 2005) in the region of Phong Nha - Ke Bang, Quang Binh 
Province, Vietnam, were deposited in the following collections: Department of 
Herpetology and Ichthyology, Muséum d'histoire naturelle, Geneva (MHNG), 
Switzerland (MHNG 2683.99); Science Research Centre of the Phong Nha - Ke Bang 
National Park, Quang Binh Province, Vietnam (PNNP 111, PNNP 112, PNNP 174, 
PNNP 175). 

To examine the phylogenetic position of Tropidophorus noggei in the general 
framework of related species of the genus we sequenced a portion of the mitochondrial 
16S rRNA gene of MHNG 2693.99 (GenBank accession number: EF611186) and 
compared it with the following species of Tropidophorus (sequences were taken from 
the GenBank): T berdmorei (AB028823), T grayi (AB222957), T partelloi 
(AB222962), T baconi (AB222953), T. sinicus (AB222954), T. cocincinensis 
(AY308323 and AB222959), T baviensis (AB222958), T hainanus (AB222960), T 
murphyi (AB222961), T. robinsoni (AB222955), T. thai (AB222956), T. latiscutatus 
(AB222950), T. matsuii (AB222952), T. brookei (AB222949), T beccami (AB222951), 
and Tropidophorus sp. (AY308322). Eutropis longicaudata (AF153572) was used as 
outgroup. 

DNA was extracted using a modified Chelex-Protocol (Walsh et al, 1991; 
Schmitz, 2003). The primers 16sar-L (light chain; 5' - CGC CTG TTT ATC AAA AAC 
AT - 3') and 16sbr-H (heavy chain; 5' - CCG GTC TGA ACT CAG ATC ACG T - 3') 
of Palumbi et al. (1991) were used to amplify a section of the mitochondrial 16S 
ribosomal RNA gene. PCR cycling procedure followed Schmitz et al. (2005). PCR 
products were purified using Qiaquick purification kits (Qiagen). Sequences (including 
complimentary strands for assuring the accuracy of the sequences) were obtained using 
an automatic sequencer (ABI 377). The obtained sequences (lengths referring to the 
aligned sequences including gaps) comprised 593 bp. Sequences were aligned using 
ClustalX (Thompson et al. 1997; default parameters) and manually checked using the 
original Chromatograph data in the program BioEdit (Hall, 1999). We used PAUP* 
4.0b 10 (Swofford, 2002) to compute the uncorrected pairwise distances for all 
sequences. 

We performed maximum parsimony (MP), Neighbor-joining (NJ) and Bayesian 
(PP) reconstructions. For Bayesian analysis parameters of the model were estimated 



SYSTEM ATICS AND NATURAL HISTORY OF TROPIDOPHORUS NOGGEI 601 

from the data set using MrModeltest 2.2 (Nylander, 2005). The NJ-analysis used the 
uncorrected 'p-distances\ Additionally, we used bootstrap analyses with 2000 (MP) 
and 20000 (NJ) pseudoreplicates to evaluate the relative branch support in phylo- 
genetic analysis. For the MP analysis, we used the "heuristic search" with the "random 
addition" option of PAUP* (Swofford, 2002) with 10 replicates, using the TBR (tree 
bisection-reconnection) branch swapping option. All Bayesian analyses were 
performed with MrBayes, version 3.0b4 (Huelsenbeck & Ronquist, 2001). We ran two 
MCMC analyses for 10 6 generations each. The initial 100000 (10%) trees were 
disregarded as "burn-in". We consider posterior probabilities (PP) of 95% or greater to 
be significantly supported. The exact parameters used for the Bayesian analyses 
followed those described in detail by Reeder (2003). 

Specimens recorded during ecological studies in Phong Nha - Ke Bang from 17 
June to 27 August 2006 by two of us (AH and TZ) were only studied in the field or 
captured for taking most important measurements (to the nearest milimeter, using a 
digital vernier caliper) and scalation characters and were subsequently released. 
Capture took place at night, beyond the skink's activity phase. We used a long pair of 
tweezers to reach and grasp the skinks in their hiding places within narrow karst rock 
crevices. Prior to their release, specimens were marked by a green-coloured deco mark- 
er (label Edding 4000) for enabling the recognition of eventual recaptures. 

Abbreviations are as follows: SVL - snout-vent length; TaL - tail length; TL - 
total length; HL - head length (distance from tip of snout to posterior margin of inter- 
parietal); HW - maximum head width; BB - dorsal body bands between limbs; MS - 
transversal midbody scale count; MD - middorsal (paravertebral) scales (from the 
posterior end of parietals to posterior margin of thigh); D - transversal dorsal scales at 
midbody; V - transversal ventral scales at midbody; L - keeled rows of lateral scales at 
midbody. 

The invertedly fixed hemipenes of the adult male specimen MHNG 2683.99 
were removed from the tail base and brought to subsequent eversion based on a method 
described by Pesantes (1994) for snakes and then successfully applied to lizards by 
Ziegler & Böhme (1997). After the temporarily storation in 2% potassium hydroxide 
solution (KOH) at 25°C, the subsequently everted outer genital organs were stored in 
70% ethanol (see also Ziegler et ah, 2005). 

For measuring the temperature and humidity we used a digital thermo- and 
hygrometer; weighing of the skinks (in cotton bags) was carried out with a spring 
balance (maximum 100g, in lg steps). Skink photographs in the field were taken with 
a Canon IXUS 55 digital camera. 

RESULTS AND DISCUSSION 

Molecular Systematics 

All molecular analyses produced an almost identical tree topology shown in 
Fig. 1. We excluded 106 bp from all analyses which were too variable to be aligned 
unambiguously. The heuristic search of the MP analysis produced 4 most-parsimonious 
trees (tree length = 261; CI = 0.475; RI = 0.557; RC = 0.264). The comparison between 
the different likelihood scores for each model showed that the GTR+I+G model (Yang 



602 



T. ZIEGLER ET AL. 



100/ 
100/ 
1 oo 



721- 



100/100 
1.00/0.99 



100/100 
1.00 



98/89 



94/80 



Eutropis longicaudata AF1 53572 
Tropidophorus berdmorei AB028823 
Tropidophorus grayi AB222957 
Tropidophorus sp. AY308322 
Tropidophorus baconi AB222953 
Tropidophorus sinicus AB222954 
Tropidophorus cocincinensis AY308323 
Tropidophorus cocincinensis AB222959 
Tropidophorus baviensis AB222958 
Tropidophorus hainanus AB222960 
Tropidophorus murphyi AB222961 
Tropidophorus robinsoni AB222955 
Tropidophorus thai AB222956 
Tropidophorus noggei EF61 1 1 86 



I Tropidophorus latiscutatus AB222950 



■— Tropidophorus matsuii AB222952 



100/98 



1.00 



r— — Tropi 

90/7/1 „. . 
0.96 *— Tr °P" 



Tropidophorus brookei AB222949 
Tropidophorus becca rii AB222951 
Tropidophorus partelloi AB222962 



FlG. 1 
Cladogram based on 487 bp of the mitochondrial 16S ribosomal RNA gene sequences. Values 
above the nodes represent bootstrap values in percent for neighbor-joining (NJ) and maximum 
parsimony (MP) analyses, respectively; lower values are Bayesian posterior probabilities (PP). 
Only significant values (a70 for NJ and MP; >0.95 for PP) are shown. Depressed-bodied species 
of Tropidophorus are written in bold. 



SYSTEMATICS AND NATURAL HISTORY OF TROPIDOPHORUS NOGGE1 603 

et al, 1994) was determined to be the optimal ML model for data set. This model in- 
corporates unequal base frequencies [tt {A) = 0.3274, tt^ = 0.2145, ttvq = 0.2593, tt (G) 
= 0.1987], a proportion of invariable sites (I = 0.4742), and a gamma distribution shape 
parameter (a = 0.4343). The optimal ML tree had a log-likelihood of -InL = 2183.2847. 

As expected for the comparatively short sequences compared, none of the trees 
showed any basal resolution but all analyses identified the same 7 terminal groups, all 
of which received significant bootstrap values to strongly support them. Therefore, we 
cannot specifically comment on the valitidy of the results of Honda et al. (2005) which 
indicated a basal split within Tropidophorus and separated a clade of continental 
Indochinese species exclusive of T. cocincinensis and T. microlepis from one 
comprising T. cocincinensis, T. microlepis and species from Borneo, Sulawesi and the 
Philippines. But in all our analyses Tropidophorus noggei clustered .well supported by 
bootstrap values (NJ: 94/MP: 83/PP: 1.00) in a clade with T. latiscutatus and T. mat- 
suii and therefore fits well into the principal "Indochina clade" of Honda et al. (2005). 

Within Tropidophorus four more or less depressed-bodied species had been 
known before the description of T noggei (Hikida et ai, 2002; Honda et ai, 2005; 
Ziegler et ai, 2005). While two moderately depressed species were collected from 
small areas in northeastern and eastern Thailand (T latiscutatus, T matsuii), one futher 
species with an extremely depressed head and body was found from one limited area 
in northern Vietnam (T. murphyi). All three species most resemble T baviensis Bourret, 
1939 from northern Vietnam in body size, body shape, and scutellation. However, the 
body depression in T. baviensis is by far not so prominent as in the other three species. 
In all our phylogenetic trees T baviensis clusters with very significant bootstrap 
support in one of the terminal clades together with T hainanus and T. murphyi (NJ: 
98/MP: 89/PP: 0.99). The newly described Tropidophorus noggei is a further species 
with a moderately depressed body and head, and its phylogenetic position as a member 
of a clade together with T latiscutatus and T matsuii (which are clearly separated from 
the former clade) confirms that the body depression recognized in several Indochinese 
species is likely to have occurred convergently at least twice as an adaptation to 
saxicolous habitats, as was presumed by Honda et al. (2005). Also the direct 
comparison of the uncorrected distance data confirms both the species status of T. 
noggei from its closest relatives T matsuii and T latiscutatus (3.29% and 2.68%, 
respectively), as well as the distinctness and separation of the two above metioned 
terminal clades (internal distances: 2.68%-3.29% ("noggd-clade") and 2.88%-3.91% 
("baviensis-clade"); distances between the two clades: 5.76%-6.78%). 

Morphology 

Beside the male holotype ZFMK 83668, the specimen MHNG 2683.99 
represents the second collected male specimen of Tropidophorus noggei. Hence, we 
herein provide a detailed description in the following: 

The measurements of MHNG 2683.99 are as follows: snout-vent length 101.8 
mm; tail length 104.5 mm; total length 206.3 mm; head length (distance from tip of 
snout to posterior margin of interparietal) 17.1 mm; head width 16.7 mm; head height 
10.3 mm; snout length (from tip to anterior margin of eye) 7.6 (left) to 8.0 (right) mm; 
eye to tympanum length (from hind margin of eye to anterior border of tympanum) 



604 T. ZIEGLER ET AL. 

8.9 mm; tympanum width 2.6 mm; tympanum height 3.7 mm; snout to forelimb length 
36.8 mm; axilla to groin length 51.4 mm; minimum neck width 13.6 mm; midbody 
width 21.6 mm; midbody height 12.1 mm; forelimb length (from body insertion to base 
of claw of fourth finger) 29.0 mm; hindlimb length (from body insertion to base of 
claw of fourth toe) 38.7 mm; length of fourth toe (without claw) 13.4 mm. 

This second preserved adult male known of T. noggei largely corresponds with 
the description of the male holotype provided by Ziegler et al. (2005), except for the 
following characters: 1) frontonasal being not only in contact with rostral, nasals, 
anterior loreals, and prefrontals, but also with the frontal (because the prefrontals are 
separated, and not in contact, as it is the case in the male holotype); 2) frontal in contact 
with frontonasal; 3) supraoculars laterally bordered by two (right) to three (left) supra- 
ciliaries (versus five supraciliaries each in the holotype); 4) left supraciliary row 
complete along the entire length of the lateral edge of the supraoculars, as it is also the 
case in the holotype, but right supraciliary row incomplete (ending at third supra- 
ocular); 5) each parietal is followed by two enlarged nuchals (three in the holotype); 6) 
43 middorsal scales (instead of 44 in the holotype) from posterior end of pari étais to 
posterior margin of thigh (paravertebral scales); 7) subdigital lamellae smooth, 
numbering 20 (versus 18-20 in the holotype) on the fourth digit of the pes; 8) seven 
pale brown transverse bands (opposite to eight to nine in the holotype) are discernible 
on the dorsum between the limbs and ten or more light bands (opposite to 17 in the 
holotype) on the dorsal tail. 

Some important measurements, the number of dorsal body bands and selected 
scalation characters of four female specimens that were collected in Phong Nha - Ke 
Bang subsequent to the original description (Ziegler et ah, 2005) are listed in Tab. 1. 
Respective data of additional 14 specimens that were captured and subsequently 
released between 16 July and 27 August 2006 are summarized in Tab. 2. Due to this 
new data gathered from the afore mentioned 19 specimens in addition to the type 
series, the diagnosis of T. noggei given in Ziegler et al. (2005) must be partially 
modified as follows: 1) existence of six to nine (mean 7.5) transverse body bands 
between the limbs; 2) the supraoculars are bordered by two to five supraciliaries; 3) the 
number of midbody scale rows ranges from 22-24 (mean 22.4), with 5-6 (mean 5.9) 
dorsals, each 5-6 (mean 5.2) laterals, and 6-7 (mean 6.1) ventral scales; 4) 43-49 (mean 
47.1) middorsal (paravertebral) scales (see also Tab. 3). 

Hemipenis Morphology 

A description of the partly everted hemipenes of the holotype of Tropidophorus 
noggei was provided by Ziegler et al. (2005). Meanwhile, with the subsequently 
everted and ca. 15 mm long left hemipenis of the specimen MHNG 2683.99 (Fig. 2) 
we have a better prepared outer genital organ at hand. It corresponds very well with the 
description of the hemipenes of the holotype, viz. being elongate, unpigmented and 
having a smooth pedicel without any further ornamentation, too. The sperm groove of 
the left hemipenis of the specimen MHNG 2683.99 is bordered with lips, apically 
forked and each running as well along the apical lobes. Furthermore, the truncus 
laterally bears each a thin-skinned tissue bulge. Asulcally, the transitory area between 
the truncus and apex is at a turgid state marked by two inflated, thin-skinned rises; 



SYSTEMATICS AND NATURAL HISTORY OF TROPIDOPHORUS NOGGEl 



605 



Table 1. Selected measurements (in mm), number of dorsal body bands between limbs and some 
scalation characters of the four female Tropidophorus noggei deposited in the collection of the 
Science Research Centre of the Phong Nha - Ke Bang National Park, Quang Binh Province; for 
abbreviations see Material and Methods. 



PNNP 1 1 1 



PNNP 112 



PNNP 174 



PNNP 175 



Min - max 
(x±s) 



SVL 


101.4 


HL 


20.3 


HW 


16.1 


BB 


8 


MS 


22 


D 


5 


V 


6 


L 


5/6 



104.2 
20.6 
16.0 

7 
22 

5 

6 
5/6 



101.2 
19.6 
15.7 

7 

24 

6 

6 
6/6 



110.2 


101.2- 110.2 


21.1 


(104.3 ±4.2) 
19.6-21.1 


16.9 


(20.4 ± 0.6) 
15.7 - 16.9 


6 


(16.2 ±0.5) 
6-8 


23 


(7 ± 0.8) 

22-24 


5 


(22.8 ±1.0) 
5-6 


6 


(5.3 ± 0.5) 
6 


6/6 


5-6 




(5.8 ± 0.5) 



along each of the outer margins of the two rises, few plicae (petala sensu Savage, 1997; 
terminology used herein after Ziegler & Böhme, 2004) are discernible. Although the 
apical lobes of the everted left hemipenis of the specimen MHNG 2683.99 are termi- 
nally not completely everted, they allow a better genital morphological analysis than it 
was the case with the only partially protruded hemipenes of the holotype of T. noggei. 
Asulcally, above the two inflated, thin-skinned rises the actual plica ornamentation 
stretches more or less horizontally along the sulcal, lateral and asulcal sides of the 
apex, below the apical lobes. These plicae, which are relatively difficult to discern and 
therefore hardly countable, are differentiated from the truncus by a distinct tissue seam. 
In addition, the "deeply bifurcated hemipenis" corroborates placement of the genital 
morphologically poorly known genus Tropidophorus in the Sphenomorphus group 
(Greer, 1979; Greer & Biswas, 2004). 

Natural History 
Habitat and abundance 

Despite recent field research from June to August 2006 within different areas of 
the karst forests of Phong Nha - Ke Bang National Park, we only could record 
T. noggei in a single mountainous region of Cha Noi area (Figs. 3, 8), from where also 
the type series originated (Ziegler et al, 2005). However, this is not astonishing, as the 
genus Tropidophorus exhibits considerable local endemism (e. g. Greer & Biswas, 
2004). All specimens of T. noggei subsequently seen by us were found in the primary 
forest in altitudes between 300-400 m above sea level. Within seven field nights at such 
higher elevations we recognized 17 different specimens, of which 14 could be captured 
for marking and taking selected measurements as well as scalation characters and sub- 



606 



T. ZIEGLER ET AL. 




Fig. 2 
Sulcal (a) and asulcal (b) view of the formerly invertedly fixed and subsequently everted left 
hemipenis of Tropidophorus noggei (MHNG 2683.99). 



Table 2. Sex, selected measurements (in mm), transversal dorsal body bands between limbs, and 
selected scalation characters of the 14 captured and subsequently released specimens of 
Tropidophorus noggei from Phong Nha - Ke Bang (for abbreviations see material and methods); 
* - with regenerated tail tip. 



Nr. 


Sex 


SVL 


TaL 


HL 


FIW 


BB 


MD 


MS 


D 


V 


L 


1 


ad. 


104.2 


124.6 


22.5 


17.2 


7 




22 


6 


6 


5/5 


2 


ad. 


90.0 


106.0 


20.2 


14.1 


6 


48 


22 


6 


6 


5/5 


3 


ad. 


68.8 


93.0 


16.2 


12.1 


8 


47 


22 


6 


6 


5/5 


4 


ad. 


94.1 


91.0* 


22.5 


17.8 


7 


43 


22 


6 


6 


5/5 


5 


f pre 


96.2 


101.5 


21.2 


17.0 


7 


49 


22 


6 


6 


5/5 


6 


ad. 


106.5 


123.5 


21.0 


16.3 


8 


48 


23 


6 


7 


5/5 


7 


f pre 


93.6 


97.0* 


20.0 


15.3 


8 


48 


22 


6 


6 


5/5 


8 


juv. 


71.5 


89.2 


16.3 


12.0 


7 


48 


22 


6 


6 


5/5 


9 


juv. 


84.9 


113.8 


19.9 


14.2 


8 


48 


22 


6 


6 


5/5 


10 


ad. 


94.2 


109.2 


22.2 


17.1 


8 


47 


23 


6 


6 


6/5 


11 


f pre 


103.3 


97.4 


22.1 


16.2 


6 


49 


22 


6 


6 


5/5 


12 


f pre 


102.1 


93.8 


21.3 


17.1 


7 


48 


23 


6 


6 


5/6 


13 


ad. 


84.4 


111.3 


21.2 


17.0 


7 






6 


6 




14 


juv. 


84.6 


119.2 


19.8 


15.3 


7 


45 


24 


6 


6 


6/6 



SYSTEMATICS AND NATURAL HISTORY OF TROPIDOPHORUS NOGGEl 607 

CHINA 




Fig. 3 
Map showing the single known locality of Tropidopiwrus noggei. 

sequently were released at the capture site. Eleven different specimens of T. noggei 
were recorded at a transect of 0.5 km within a single night only, from 19: 14h to 23:37h, 
pointing to a regionally high abundance. Because eight of the 14 marked individuals 
have been captured at prominent karst formations, we were able to subsequently 
relocate the individual sites. After a period of five weeks we found six of the eight 
skinks at the same location anyway. This recapture rate of 75% explicitly argues for a 
seldom relocation, thus the territorial habits of T. noggei. 

All our records of Twpidophorus noggei took place in heights of 0.3 to 1.5 m 
above the ground. With a piece of shed skin found on 21 June 2006 in a chink of a karst 
rock projection together with an observation of a specimen dwelling inside a karst 
crevice we also could prove that the species is at least able to climb up to 1.8 m height 
above the ground. The vertical diameter of the karst rock crevices in which we found 
live specimens, measured seven to 50 mm (mean ca. 20 mm). With the dorso- ventrally 
depressed body as well as the robust and resistant scalation the species is exceedingly 
adapted to such narrow rock crevices (Figs. 4-5). All specimens recognized by us were 
found solitarily, however, nine (i. e. ca. 64%) of the 14 captured specimens lingered in 
crevices of immediate vicinity, within distances of 0.3 to 1 m to each other. This 
suggests, that T. noggei occupies certain territories and that adults as well as juveniles 
together with adults occur in close vicinity. 



608 



T. ZIEGLER ET AL. 




Figs 4-7 
(4) Tropidophorus noggei active during daytime in front of its hiding place, a narrow karst rock 
crevice. (5) Such narrow karst rock crevices represent the microhabitat of Tropidophorus noggei. 
(6) Tropidophorus noggei at night in its hiding place in typical sleeping as well as defending 
position. (7) Tropidophorus noggei during daytime lurking at the entrace of a karst crevice. 



Further lizard species that were observed by us in syntopy with T. noggei were 
scincids of the genera Lygosoma and Scincella, the gekkonids Cyrtodactylus 
phongnhakebangensis, C. cryptus, Gekko gecko, G. scientiadventura, as well as the 
agamids Acanthosaura lepidogaster and Calotes emma (Heidrich et al, 2007; Ziegler 
et al, 2007). As potential predators of at least juvenile T. noggei we found large 
centipedes (Scolopendra sp., Scutigera sp.) crawling in and around the karst rock 
crevices. Furthermore, the snake species Psammodynastes pulverulentus and 
Protobothrops cornutus, as well as Lycodon representatives were observed occuring 
together with T. noggei. 

Activity and habits 

Ziegler et al. (2005) reported, that T. noggei specimens were found at night at 
the base of karst rock outcrops in front of horizontal and narrow rock crevices about 30 
to 50 cm above the forest floor. After being disturbed, they escaped into the narrow 
rock crevices, where they were well concealed and anchored up to 15 to 30 cm inside 
the karst rock crevices. During our recent ecological field work we could find out that 
most skinks spent the night resting or sleeping in narrow rock crevices. We further 



SYSTEMATICS AND NATURAL HISTORY OF TROP1DOPHORUS NOGGEI 



609 




Fig. 8 
Habitat of Tropidophorus noggei in Cha Noi area, Phong Nha - Ke Bang National Park, Quang 
Binh Province, Truong Son, central Vietnam. 



observed a typical defensive position (Fig. 6), with the tail, being laterally positioned 
in front of the body to protect the damageable trunk parts and the head. It seems that 
the species is also able to autotomize its tail once being heavily grasped at, which was 
once observed by us on 21 June 2006. Further, T. noggei was observed by us to be 
mainly active during daytime. However, also at daytime, the crevices were seldomly 
left. For example, the adult specimen 1 (see Tab. 2 and 4), that was observed at the 
afternoon of 7 August 2006 for a period of two hours (15:15 to 17:15), left its crevice 
only once for a short time and remained only in a range of about one meter in doing 
so. The skink rather spent most of the two hours of our observation lurking at its 
crevice entrance (Fig. 7). 

Reproduction 

Concerning reproductive biology, the testes of the male MHNG 2683.99 
measured 5.8-6.3 mm in length and 3.2-3.6 mm in width. The testes of the commen- 
surate male holotype measured 9.0 mm in length and 6.2 mm in width (Ziegler et al, 
2005). Whilst the specimen MHNG 2683.99 most probably was collected during the 
dry season, the larger testes of the holotype may be established by its collect at the end 
of the dry season. One of the two female paratypes, that was also collected at the end 



610 



T. ZIEGLER ET AL. 



Table 3. Maximum snout-vent length, head length, and head width of the three type specimens 
of Tropidophorus noggei (from Ziegler et al, 2005), of the subsequently collected male MHNG 
2683.99, of the four specimens deposited in the Science Research Centre of the Phong Nha - Ke 
Bang National Park (see Table 1), and of the 14 captured and subsequently released specimens 
(see Table 2), as well as minima, maxima, mean and standard deviation of the number of dorsal 
body bands between the limbs and of selected scalation characters (for abbreviations see material 
and methods). 







n 


Min-max 

x ± s 


SVL 


max. 


22 


110.2 mm 


HL max. 


22 


22.5 mm 


HW 


max. 


22 


17.8 mm 


BB 




22 


6-9 
(7.5 ± 0.9) 


MD 




16 


43-49 
(47.1 ±2.1) 


MS 




21 


22-24 
(22.4 ± 0.7) 


D 




22 


5-6 
(5.9 ± 0.4) 


V 




22 


6-7 
(6.1 ±0.2) 


L 




21 


5-6 
(5.2 ± 0.4) 



Table 4. Collecting date, sex, weight (in g), time of discovery, temperature, relative humidity 
and vertical karst crevice diameter (CD: in mm) as well as crevice height above ground (CH: in 
m) for the 14 captured and subsequently released specimens of Tropidophorus noggei from 
Phong Nha - Ke Bang. For abbreviations see Material and Methods; further abbreviations are as 
follows: Ad. - adult; f - female; pre - pregnant; juv. - subadult. 



Nr. 


Date 


Sex 


Weight 


Time 


Temp. 


Humidity 


CD 


CH 


1 


16.7. 


ad. 


25 


18:56 


24.9°C 


94% 


18 


1.0 


2 


17.7. 


ad. 


19 


19:14 


24.3°C 


94% 


13-27 


1.0 


3 


17.7. 


ad. 


6.5 


19:25 


24.3°C 


94% 


7-19 


0.3 


4 


17.7. 


ad. 


23 


19:49 


24.1°C 


93% 


22-34 


1.5 


5 


17.7. 


f pre 


24.5 


19:55 


24. rc 


93% 


33-50 


1.5 


6 


17.7. 


ad. 


30.5 


20:37 


24.3°C 


93% 


9-23 


0.4 


7 


17.7. 


f pre 


22 


21:14 


24.4°C 


92% 


14-22 


1.2 


8 


17.7. 


juv. 


6.5 


21:25 


24.4°C 


93% 


18 


1.2 


9 


17.7. 


juv. 


14 


21:45 


24.4°C 


92% 


11 


1.3 


10 


17.7. 


ad. 


19.5 


22:16 


24.4°C 


92% 


11-30 


1.4 


11 


17.7. 


f pre 


24 


22:27 


24.2°C 


88% 


14.5-16 


0.8 


12 


17.7. 


f pre 


30 


23:37 


24.2°C 


88% 


17-20 


0.7 


13 


27.8. 


ad. 


21 


21:10 


25.0°C 


86% 




1.0 


14 


27.8. 


juv. 




21:20 


25.0°C 


86% 


14-18 


1.1 



SYSTEM ATICS AND NATURAL HISTORY OF TROPIDOPHORUS NOGGEI 611 

of the dry season, contained three large eggs of about 13x10 mm diameter. The four 
female specimens deposited in the collection of the Science Research Centre of the 
Phong Nha - Ke Bang National Park most probably were collected during different 
seasons. The two specimens (PNNP 111, PNNP 112), that most probably were 
collected during the dry season, contained nine and 18 small eggs of 2-3 and 1-4 mm 
maximum diameter, respectively. In contrast, the specimen PNNP 1 74 contained five 
eggs of 12.5 x 8 mm maximum diameter, and in the dissected specimen PNNP 175 we 
found seven eggs of 10.6 x 8.9 mm size. Throughout July, at the end of the dry season, 
we found pregnant females in the field (see Tab. 4), well recognizable from their 
distinctly swollen bodies. It should be interesting to continue with studying the species' 
reproductive mode (see Hikida et al, 2002), because several authors have assumed that 
in lizards the physical constraint from the crevice-dwelling habits provides an evolu- 
tionary force to some reproductive traits, such as relative clutch mass and frequency of 
clutch production (Vitt, 1981, 1993; Doughty & Shine, 1995). 

Diet 

With respect to feeding ecology, we found remains of a worm-like invertebrate 
(cf. Annelida) in the stomach of MHNG 2683.99. The stomachs of the four females 
deposited in the collection of the Science Research Centre of the Phong Nha - Ke Bang 
National Park contained each remains of a worm (Annelida) in the specimens PNNP 
174 and PNNP 175, and three ants (Hymenoptera: Formicidae) in the specimen PNNP 
111. The stomach of the female PNNP 112 was empty, but its gut contained four ants, 
four termites (Isoptera) and one katydid (Ensifera). Whereas the guts of the specimens 
PNNP 174 and PNNP 175 were empty, the gut of PNNP 1 1 1 contained five ants. The 
contents of the gastro-intestinal tracs of the afore mentioned specimens together with 
the data obtained from the dissected holotype and one female paratype of T. noggei (see 
Ziegler et al, 2005) are summarized in Fig. 9. 

The most frequent prey items were ants and termites, followed by annelid 
worms (see Fig. 9). However, the higher prey item amount in the guts as can be seen 
in Fig. 9 is put down to an accumulation of prey fragments that are hard to digest to not 
digestible, as is the case with chitin fragments of ants and termites. For this reason, and 
because gut contents usually do not contain easily digestible prey like soft-skinned in- 
sect larvae or worms, and therefore have limited value in comparison with more diverse 
stomach contents, stomach and gut content data were presented separately in Fig. 9 
(see also Ziegler, 2003). Therefore the high number of ant and termite prey items must 
be carefully interpreted. In addition, only prey item numbers are presented in Fig. 9 
and not their respective masses. To visualize this in a direct comparison: three of the 
relatively large annelid worms (with at least 3 mm body diameter) were found in the 
stomachs of three (43%) of the altogether seven dissected skinks, but the much more 
smaller ants were only in the guts of two individuals (29%) and the 5 mm small ter- 
mites only in the gut of a single skink (14%). Based on the worms' high biomasses and 
the fact that 43 percent of the skinks had worm remains in their stomach, it can be as- 
sumed that worms play an important role in the feeding ecology of T. noggei. In 
addition, it is interesting to note, that ants only were found in the gastro-intestinal tracts 
of the specimens PNNP 1 1 1 and PNNP 1 12, the latter one of which contained as 



612 



T. ZIEGLER ET AL. 



Hymenoptera 
(Formicidae) 



Isoptera 



Annelida 



Ensifera 



Insecta indet. 



Chilopoda 



Fig. 9 
Number of prey items found in the stomachs (white columns) and guts (grey columns) of seven 
examined specimens of Tropidopiwrus noggei. 

single skink specimen also termites. Due to their small egg sizes, both skinks most 
probably were collected during another season than the females PNNP 174 and PNNP 
175, which both had worm remains in their stomachs, as well as the male MHNG 
2683.99, that most probably was collected during or at the end of the dry season. This 
further would allow the conclusion of a seasonal food supply and prey spectrum, 
respectively, because it can be expected that the worm abundance increases with 
beginning rains. However, during our field work in July and August 2006 we found 
both earth worms, as well as ants and termites in and around the karst crevices being 
inhabited by T. noggei. 



ACKNOWLEDGEMENTS 

We thank Dr V. T. Hoang and N. T. Vu (Centre for Natural Resources and 
Environmental Studies, Vietnam National University, Hanoi), the People's Committee 
of Quang Binh. and the Phong Nha - Ke Bang National Park directorate (T. H. Nguyen, 
X. C. Cao, and M. T Luu) for their support as well as for issuing respective permits. 
The National Park staff, especially Dang Ngoc Kien (Science Research Centre) helped 
again to make fieldwork successful. For their assistance during field work we would 
like to express our deepest thanks to our local guide Nguyen Van Hoan, as well as to 
Dr Martina Vogt (Cologne Zoo nature conservation project, Phong Nha - Ke Bang) and 
Bernhard Forster (Frankfurt Zoological Society / Cologne Zoo nature conservation 
project, Phong Nha - Ke Bang). Our field work was funded in large by the Zoological 
Garden Cologne, the Kölner Kulturstiftung der Kreissparkasse Köln, and BIOPAT 
(www.biopat.de). Current lizard research in the Phong Nha - Ke Bang region is 



SYSTEM ATICS AND NATURAL HISTORY OF TROPIDOPHORUS NOGGEI 613 

supported as well by the Alexander-Koenig-Gesellschaft (AKG), the Alexander 
Koenig Stiftung (AKS), the Deutsche Gesellschaft für Herpetologie und Terrarien- 
kunde (DGHT), and by the European Union of Aquarium Curators (E.U.A.C.). 

REFERENCES 

Ballinger, R.E., Lemos-Espinal, J.A. & Smith, G.R. 2000. Reproduction in females of three 

species of crevice-dwelling lizards (genus Xenosaurus) from Mexico. Studies on 

Neotropical Fauna and Environment 35: 179-183. 
Doughty, P. & Shine, R. 1995. Life in two dimensions: natural history of the southern leaf-toed 

gecko, Phyllurus platurus. Herpetologica 51: 193-201. 
Greer, A. E. 1979. A phylogenetic subdivision of Australian skinks. Records of the Australian 

Museum 32: 339-371. 
Greer, A.E. & Biswas, S. 2004. A generic diagnosis for the southeast Asian scincid lizard 

Tropidophorus Duméril and Bibron, 1839 with some additional comments on its 

morphology and distribution. Journal of Herpetology 38(3): 426-430. 
Hall, T.A. 1999. BioEdit: a user- friendly biological sequence alignment editor and analysis 

program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95-98. 
Heidrich, A., Rösler, H., Vu Ngoc Thanh, Böhme, W. & Ziegler, T 2007. Another new 

Cyrtodactylus (Squamata: Sauria: Gekkonidae) from Phong Nha - Ke Bang National 

Park, central Truong Son, Vietnam. Zootaxa 1445: 35-48. 
Hikida, T, Orlov, N.L., Nabhitabhata, J. & Ota, H. 2002. Three new depressed-bodied water 

skinks of the genus Tropidophorus (Lacertilia: Scincidae) from Thailand and Vietnam. 

Current Herpetology 21(1): 9-23. 
Honda, M., Ota, H, Murphy, R. W. & Hikida, T 2005. Phylogeny and biogeography of water 

skinks of the genus Tropidophorus (Reptilia: Scincidae): a molecular approach. 

Zoologica Scripta 35: 85-95. 
Huelsenbeck, J. P., RONQUIST, F. 2001. MRBAYES: Bayesian inference of phylogenetic trees. 

Bioinfonnatics 17: 754-755. 
Nylander, J. A. A. 2005. MrModeltest 1.1b, Uppsala, Sweden. 
Palumbi, S.R., Martin, A., Romano, S., McMillan, W.O., Stice, L., Grabowski, G. 1991. The 

simple fool's guide to PCR. Department of Zoology and Kewalo Marine Laboratory, 

Hawai: 47 pp. 
Pesantes, O.S. 1994. A method for preparing the hemipenis of preserved snakes. Journal of 

Herpetology 28: 93-95. 
Reeder, T.W. 2003. A phylogeny of the Australian Sphenomorphus group (Scincidae: 

Squamata) and the phylogenetic placement of the crocodile skinks (Tribolonotus): 

Bayesian approaches to assessing congruence and obtaining confidence in maximum 

likelihood inferred relationships. Molecular Phylogenetics and Evolution 27: 384-397. 
Savage, J.M. 1997. On terminology for the description of the hemipenes of squamate reptiles. 

Herpetological Journal 7: 23-25. 
Schmitz, A. 2003. Taxonomic and phylogenetic studies on scincid lizards (Reptilia: Scincidae). 

Unpubl. PhD thesis. University of Bonn, 262 pp. 
Schmitz, A., Ineich, I., Chirio, L. 2005. Molecular review of the genus Panaspis sensu lato in 

Cameroon, with special reference to the status of the proposed subgenera. Zootaxa 863: 

1-28. 
Swofford, D.L. 2002. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). 

Version 4.0M0. Sinauer Associates, Sunderland, Massachusetts, USA. 
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin F., Higgins, D. G. 1997. The ClustalX 

windows interface: flexible strategies for multiple sequence alignment aided by quality 

analysis tools. Nucleic Acids Research 24: 4876-4882. 
Vitt, L.J. 1981. Lizard reproduction: habitat specificity and constraints on relative clutchmass. 

American Naturalist 117: 506-514. 



614 T. ZŒGLER ET AL. 



Vitt, L.J. 1993. Ecology of isolated open-formation Tropidurus (Reptilia: Tropiduridae) in 

Amazonian lowland rain forest. Canadian Journal of Zoology 71: 2370-2390. 
Walsh, P.S., Metzger, D.A., Higuchi, R. 1991. Chelex 100 as a medium for simple extraction 

of DNA for PCR-based typing from forensic material. BioTechniques 10(4): 506-513. 
Yang, Z. H., Goldman, N. & Friday, A. 1994. Comparison of Models for Nucleotide 

Substitution Used in Maximum-Likelihood Phylogenetic Estimation. Molecular Biology 

and Evolution 1 1 : 3 16-324. 
Ziegler, T. 2003. Diet and natural history notes of Leptodactylus laticeps (Anura: 

Leptodactylidae) in the Gran Chaco of Paraguay. Salamandra 39(1): 39-48. 
Ziegler, T. & Böhme, W. 1997. Genitalstrukturen und Paarungsbiologie bei squamaten 

Reptilien, speziell den Platynota, mit Bemerkungen zur Systematik. Mertensiella 8: 

1-207. 
Ziegler, T. & Böhme, W. 2004. On the genital morphology of blue-tongued skinks (Scincidae: 

Lygosominae: Tiliqua). In: Hitz, R., Shea, G. M., Hauschild, A., Henle, K. & H. 

Werning (eds): Blue-tongued skinks. Contributions to the knowledge of Tiliqua and 

Cyclodomorphus. Mathias Schmidt Publications, Münster. 45-52. 
Ziegler, T., Gaulke, M. & Böhme, W. 2005. Genital morphology and systematics of Varanus 

mabitang Gaulke & Curio, 2001 (Squamata, Varanidae). Current Herpetology 24(1): 

13-17. 
Ziegler, T., Schmitz, A., Heidrich, A., Vu Ngoc Thanh & Nguyen Quang Truong. 2007. A 

new species of Lygosoma (Squamata: Sauria: Scincidae) from the Central Truong Son, 

Vietnam, with notes on its molecular phylogenetic position. Revue Suisse de Zoologie 

114(2): 397-415. 
Ziegler, T., Vu Ngoc Thanh & Bui Ngoc Thanh. 2005. A new water skink of the genus 

Tropidophorus from the Phong Nha - Ke Bang National Park, central Vietnam 

(Squamata: Sauria: Scincidae). Salamandra 41(3): 137-146. 



Revue suisse de Zoologie 1 14 (3): 615-653; septembre 2007 



Studies of the genus Anthelephila Hope (Coleoptera: Anthicidae) 
10. Species related to A. imperatrix 

Zbynëk KEJVAL 

Muzeum Chodska, Domazlice CZ-344 01, Czech Republic. 

E-mail: kejval@cmail.cz 



Studies of the genus Anthelephila Hope (Coleoptera: Anthicidae) - 10. 
Species related to A. imperatrix. - Anthelephila imperatrix LaFerté- 
Sénectère, 1849, A. subtruncata (Pic, 1899) and male characters of A. besu- 
chen Bonadona, 1989, A. congoana Uhmann, 1981 and A. ovipennis 
(Bonadona, 1984) are redescribed. Eight new species are described: A. car- 
damontis sp. n. (India), A. curvitarsis sp. nov. (South Africa), A. aratrix sp. 
n. (India), A. irula sp. n. (India), A. kresli sp. n. (Nepal), A. lobulicula sp. n. 
(Nepal), A. sculpta sp. n. (India, Bhutan) and A. vanhillei sp. n. (South 
Africa). New synonymy, A. imperatrix LaFerté-Sénectère, 1849 (= Formi- 
comus punctaticeps Pic, 1916 syn. n.), is proposed. Status oï F. cribriceps 
Marseul, 1876 is discussed and its previous placement in synonymy with A. 
imperatrix LaFerté-Sénectère, 1849 is supported. 

Keywords: Coleoptera - Anthicidae - Anthelephila - systematics - new 
species - new synonymy. 

INTRODUCTION 

An informal A. imperatrix species-group is here established for five known and 
eight newly described species of the genus Anthelephila Hope, 1833. Apart from their 
prevailing aptery and simple forelegs of the males, the included species are distinguis- 
hed by possession of the simpler structure of male abdominal segment VIII, nearly stra- 
ight lateral mesosternal margins, and the paired incisions on the posterior, exposed 
margin of the prosternum. All of these characters occur rarely within the genus, and so- 
me of them suggest, along with the discontinuous ranges, a rather basal position of this 
species-group, and also a closer relationship of Anthelephila to Stenidius LaFerté- 
Sénectère, 1847. 

ABBREVIATIONS AND EXPLANATIONS 

The following abbreviations of collections are used (in round brackets): 

ADBC collection Augusto Degiovanni, Bubano, Italy 
AMGS Albany Museum, Grahamstown, South Africa 
BMNH The Natural History Museum, London, England 



Manuscrit accepté le 09.03.2007 



616 Z. KEJVAL 

DCDC collection Donald S. Chandler, Durham, New Hampshire, U. S. A. 

GUPC collection Gerhard Uhmann, Pressath, Germany 

HNHM Hungarian Natural History Museum, Budapest, Hungaria 

MHNG Muséum d'histoire naturelle, Genève, Switzerland 

MNHB Museum für Naturkunde der Humboldt Universität, Berlin, Germany 

MNHN Muséum National d'Histoire Naturelle, Paris, France 

NHMB Naturhistorisches Museum, Basel, Switzerland 

NHMW Naturhistorisches Museum, Wien, Austria 

NKME Naturkundemuseum Erfurt, Erfurt, Germany 

NMPC National Museum, Prague, Czech Republic 

S ANC S. A. National Collection of Insects, ARC - Plant Protection Research 

Institute, Pretoria, South Africa 

SMF Senckenberg-Museum, Frankfurt am Main, Germany 

SMNS Staatliches Museum für Naturkunde, Stuttgart, Germany 

TMP Transvaal Museum, Pretoria, South Africa 

ZILS Universitets Zoologiska Institut, Lund, Sweden 

ZKDC collection Zbynëk Kejval, Domazlice, Czech Republic 

The author's comments on the type material are placed in square brackets: [p] 
= printed, [h] = handwritten. Exact label data are quoted only for the type specimens. 
Separate labels are indicated by back slashes ( \ ). The terminology of body setation 
follows Werner & Chandler (1995). 

SYSTEMATICS 

Anthelephila imperatrix species-group 

Diagnosis: Small to medium sized (2.6-4.6 mm), almost exclusively apterous 
species, very rarely with fully developed metathoracic wings (only six specimens of A. 
imperatrix and A. kresli sp. n.); posterior margin of prosternum with a pair of incisions 
(Fig. 7, best observed after removing the coxae); elytra ovoid, often rather strongly 
convex and subtruncate/truncate apically, with obsolete humeri and lacking postscutel- 
lar impression (characters related to aptery); forelegs in males simple; meso- and me- 
tatibiae often modified (six species); lateral margins of mesosternum nearly straight 
(Fig. 35); tergum VII lacking paired spinulose patches on dorsal side (a character pos- 
sibly related to aptery, patches may be used in folding of metathoracic wings); male 
sternite VIII less differentiated, median sclerite indistinct and paired prongs more or 
less tightly joined medially, latero-basal plates inconspicuous; male tergite VIII for- 
ming single sclerite; tegmen trilobed apically; median lobe of aedeagus terminating in 
a pair of narrow projections (Fig. 15). Species included: A. aratrix sp. n., A. besu- 
chen, A. cardamontis sp. n., A. cunntarsis sp. n., A. congoana, A. imperatrix, A. irula 
sp. n., A. kresli sp. n., A. lobulicula sp. n., A. ovipennis, A. sculpta sp. n., A. subtrun- 
cata and A. vanhillei sp. n. 

Distribution: One Asian species, A. imperatrix, exhibits a wide distributional 
range. All other species are known from limited regions in the following countries: 
South Africa (3); Zaire, Uganda and Kenya (2); south-western India and Sri Lanka (3); 
north-eastern India and Nepal (4). 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 617 

Comments: It should be emphasized, that A. imperatrix species-group is 
established as an informal group of convenience. Its polyphyletic origin is not 
excluded, as the following important character states appears to be primitive and/or 
variable within the genus: 

Mesosternal margins: In Anthelephila the lateral mesosternal margins are typi- 
cally moderately arcuate in their posterior half (see figs 1, 6 by Kejval, 2003). Straight 
margins, exhibited by members of A. imperatrix species-group, occur in more primitive 
Anthicinae and they are considered to be the ancestral state (Chandler, 1982). However, 
the form/width of lateral arms of mesosternum shows variation and some Anthelephila 
species appears to be intermediate in this character. 

Abdominal segment VIII: The modified male segment VIII is regarded as a 
synapomorphy for the genera of Formicomini Bonadona, 1875 (Kejval, 2003), and 
species of Anthelephila are characterized almost exclusively by the more complicated 
structure of this segment (sternite VIII differentiated into five separable parts, tergite 
VIII composed of two sclerites). Consequently, the rather simple structure of segment 
Vili in the A. imperatrix species-group represents either a more primitive condition 
within the genus, or a derived form characterized by simplification. 

Prosternai incisions: In all Anthelephila the portion of the prosternum beneath 
the coxae is sclerotized and extended posteriorly as a distinct sclerite. The posterior 
margin of this sclerite is typically simple and shallowly emarginate (see Fig. 6 by 
Kejval, 2003), very rarely with a pair of incisions (Fig. 7). The phylogenetic signi- 
ficance of these prosternai incisions is not clear. Moreover, this character is not 
restricted to the A. imperatrix species-group. Showing clear variation in prominence, it 
occurs also in A. kanheri Kejval, 2002 and several related species inhabiting the region 
of Western Ghat on the Indian subcontinent, which are rather typical Anthelephila with 
well-developed metathoracic wings, more complicated structure of segment VIII and 
modified forelegs in males. Other Indian species, more widely distributed and showing 
similarities in male characters to this small group, lack prosternai incisions completely. 

Remarkably, the related genus Stenidius resembles species of A. imperatrix 
group in having a less complicated structure of the modified abdominal segment VIII, 
and its species often display similar prosternai incisions. 

A KEY TO SPECIES OF A. IMPERATRIX SPECIES-GROUP 

1(4) Dorsal outline of pronotum rather strongly convex in anterior two thirds, 
impressed and then distinctly bulging before base in lateral view 
(Fig. 6); anterior portion of pronotai disc with conspicuous median 
longitudinal impression/groove. 

2(3) Ordinary punctation and setation of elytra very sparse, scattered; 
posterior band of contiguous punctures and thicker whitish setae 
directed postero-mediad from lateral margins (Fig. 8); male sternum VII 
with conspicuous postero-median process (Fig. 10) A. aratrix sp. n. 

3(2) Ordinary punctation and setation of elytra somewhat denser; posterior 
band of contiguous punctures and thicker whitish setae directed antero- 
mediad from lateral margins (Fig. 2); male sternum VII modified, but 
lacking any postero-median process A. imperatrix LaFerté-Sénectère 



618 Z. KEJVAL 

4(1) Entire dorsal outline of pronotum more or less convex in lateral view; 
pronotai disc rather evenly shaped, median longitudinal impression 
absent or at most moderately indicated and inconspicuous. 

5(10) Elytra rather sparsely and evenly punctured and setose, lacking any 
bands/patches of more densely spaced punctures and contrasting whitish 
setae, their setation pale, longer, with conspicuous erect setae; male 
metasternum simple. 

6(7) Elytra rufous, with paired dark spots at about midlength laterally and 
subapically (Fig. 1); surface of head and pronotum almost entirely 
smooth (excepting fine wrinkles in postero-lateral impressions of pro- 
notum), finely punctured and very glossy A. cardamontis sp. n. 

7(6) Elytra largely dark coloured, at most with paler base and suture; surface 
of head and/or pronotum coarsely corrugated, more distinctly punctured. 

8(9) Dorsal corrugation of head obsolete, punctation distinct; lateral margin 
of elytra moderately convex subapically, elytral apices rounded; male 
character as in Figs 22-25 A. irula sp. n. 

9(8) Dorsal corrugation of head conspicuous, punctation concealed by corru- 
gation and thus less distinct; lateral margin of elytra slightly concave 
subapically, elytral apices rather pointed; male characters as in Figs 
16-18 A. besuchen Bonadona 

10(5) Elytra unevenly punctured and setose, except ordinary punctures/setae 
with two, paired, transverse to oblique bands/patches of denser, mostly 
contiguous punctures and thicker whitish setae, if punctation nearly 
evenly sparse and setose bands/patches vaguely indicated (sparse) to 
indistinct, then setation of elytra uniformly short and inconspicuous, 
lacking longer, more raised setae; male metasternum with a pair of pro- 
trusions posteriorly, bordering laterally more or less prominent postero- 
median impression. 

1 1(20) Dorsal punctation of head distinct but comparatively fine and sparse; se- 
tation generally short and inconspicuous, whitish setose bands/patches 
sparser, vaguely indicated to indistinct; Afrotropical species. 

12(15) Species nearly uniformly dark in colour, including legs and antennae, at 
most basal 2-3 antennomeres slightly paler; head somewhat widely 
rounded posteriorly in dorsal view; basal protarsomere in males rather 
short and narrow. 

13(14) Pronotum wider, less narrowed and constricted posteriorly in dorsal 
view, its postero-lateral impressions rather shallow and situated close be- 
fore base; male middle legs uniformly short setose, mesofemora simple, 
mesotibiae subapically strongly excavated on inner side (Figs 75, 76); 
metatibiae rather simple apically; other male characters as in Figs 77-79 
A. vanhillei sp. n. 

14(13) Pronotum more narrowed and constricted posteriorly in dorsal view, its 
postero-lateral impressions deeper and more distant from base; male 
middle legs with long setae on inner side (Fig. 71), mesofemora slightly 
produced on inner side distally, mesotibiae simple; inner apical margin 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 619 

of metatibiae projecting into blunt process (Fig. 69); other male 
characters as in Figs 70, 72-74 A. subtruncata (Pic) 

15(12) Species with dark body and distinctly paler legs, antennae and palpi (at 
least partly); head posteriorly circular to oval in dorsal view; basal pro- 
tarsomere in males enlarged, conspicuously long. 

16(17) Postero-lateral impressions of pronotum conspicuous, more distant from 
base and thus pronotum clearly constricted in posterior half in dorsal 
view (Fig. 3); whitish setose bands/patches of elytra indistinct, at most 
slightly indicated; basal protarsomere in males narrow, curved (Fig. 39); 
other male characters as in Figs 38, 40-43 A. curvitarsis sp. n. 

17(16) Postero-lateral impressions of pronotum somewhat less conspicuous, si- 
tuated near base; whitish setose bands/patches rather sparse but distinct, 
especially posterior one; basal protarsomere in males wider, straight. 

18(19) Base of head somewhat unevenly rounded, slightly produced medially; 
male metatibiae uniformly, sparsely setose; male characters as in Figs 
26-29; female tergum VII triangular, strongly narrowed, bluntly pointed 

apically (Fig. 34) A. congoana 

Uhmann 

19(18) Base of head rather evenly rounded; male metatibiae subapically with 
small patch of brownish, short and dense setae on inner side (Fig. 36); 
other male characters as in Figs 30-33; female tergum VII subtriangular, 
moderately narrowed, rather rounded apically (Fig. 37) 
A. ovipennis (Bonadona) 

20(11) Dorsal punctation of head coarse and rather dense, punctures separated 
by about their diameter; setation much longer and more raised, whitish 
setose bands/patches dense, conspicuous; Oriental/Palaearctic species. 

2 1(22) Pronotum distinctly narrower than head including eyes; punctation of 
head and pronotum clearly different, punctures of pronotum, especially 
the postero-dorsal and lateral ones, much finer that those of head; male 
characters as in Figs 58-61, 63 A. lobulicula sp. n. 

22(21) Pronotum at most slightly narrower than head including eyes; punctati- 
on of head and pronotum nearly identical. 

23(24) Surface of head and pronotum less glossy, very coarsely and rather even- 
ly sculptured, punctures situated in nearly contiguous, rather deep, pit- 
like impressions; male characters as in Figs 64-68 A. sculpta sp. n. 

24(23) Surface of head and pronotum normally, coarsely and somewhat une- 
venly punctured, with interspaces smooth and glossy; male characters as 
in Figs 54-57, 62 A. kresli sp. n. 

Anthelephila aratrix sp. n. Figs 8-15 

Type material: Holotype o\ INDIA W. Bengal Darjeeling dist. Algarah 1800 m 9-X-78 
Besuchet Lobi \ Anthelephilus imperator (Laferté) det.G.Uhmann 1986 (MHNG). - Paratypes: 
1 6,1 9 , same data as holotype (ZKDC, MHNG). -19, same data as holotype, except: 250 m 
[see Remarks] (MHNG). - 1 3, Darjeeling Distr India Bhakta B. \ Gorco Bethan 840m 
21.IV.1979 \ Anthelephila imperator (LaF.) det. D. Telnov, 2000 (NHMB). - 1 o\ Darjeeling 
Distr India Bhakta B. \ Chuba 670m, 11. IV. 1979 \ Anthelephila imperator LaF. det. D. Telnov, 



620 



Z. KEJVAL 




Figs 1-7 
Habitus of Anthelephila: (1) A. cardamontis sp. n., male. (2) A. imperatrix LaFerté-Sénectère, 
female. (3) A. cunitarsis sp. n., male. (4) A. sculpta sp. n., male. (5) A. vanhillei sp. n., male. 
A. imperatrix LaFerté-Sénectère: (6) pronotum, lateral view. (7) pronotum, antero-ventral view. 
Scale (1 mm): A - Figs 1-5, B - Figs 6, 7. 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 62 1 

1999 (ZKDC).- 1 ?,DarjeelingDistrIndiaBhaktaB.\Monshong 1350m 23.XI. \ Anthelephila 
imperator (LaF.) det. D. Telnov, 2000 (NHMB). - 1 ? , Indien Darjeeling D Ch. J. Rai \ Kalimp. 
800m Upper Janake 17.IV. 1987 \ Anthelephila imperator LaF. det. D. Telnov, 1999 (NHMB). 

Etymology: From Latin arator (ploughman); named in reference to peculiar, 
plough-like shaped process of male sternum VII. 

Description: Body length 2.9-3.5 mm (holotype 3.3 mm). 

Male (holotype): Identical with A. imperatrix, except for the following 
characters: antennae somewhat more slender in distal third; ordinary setation of elytra 
sparser, posterior band of contiguous punctures and thicker whitish setae directing 
from lateral margin postero-mediad (Fig. 8); procoxae angulately produced, punctation 
of metafemora somewhat finer and less conspicuous. Sternum VII (Figs 9, 10) 
emarginate posteriorly and projecting ventrad into peculiar, conspicuously large 
median process, apical portion of this process abruptly curved posteriad, and strongly 
widened, trapezoidal in ventral view, and with small denticle on each side latero- 
basally. Tergum VII (Fig. 1 1 ) truncate posteriorly, its posterior margin moderately 
sinuous. Sternite VIII (Figs 12, 13); paired prongs robust, rather simple, arcuately 
curved ventrad in lateral view, their dorso-median margin dilated mediad at about 
midlength into small, rounded process; setation of prongs inconspicuous, short, fine 
and scattered. Tergite VIII (Fig. 12) nearly parallel-sided in dorsal view, truncate 
posteriorly, with rounded postero-lateral angles, finely and shortly setose. Aedeagus 
(Figs 14, 15); apical portion of tegmen 0.8 times as long as basal-piece, trilobed 
apically, middle lobe conspicuously wide, strongly and nearly evenly narrowing 
towards pointed apex, slightly longer than apically rounded, lateral lobes; apical paired 
projections of median lobe of aedeagus flattened, angulately dilated laterad and ventrad 
subapically, rounded apically. 

Female: Externally identical with male, except as follows: procoxae simple; 
sternum VII simple, evenly rounded posteriorly; tergum VII triangular, narrowed and 
rounded apically, its apical portion rather strongly vaulted, but evenly shaped, lacking 
a median edge or protrusion. 

Variability: Inconspicuous. 

Differential diagnosis: A. aratrix sp. n. is habitually very similar and 
undoubtedly closely related to A. imperatrix, but differs substantially in most male 
characters; see the above description and the key. 

Distribution: India (West Bengal). 

Remarks: With respect to the identical sample number, handwritten on the 
other side of label, all the type specimens from Algarah should bear the same locality 
data; the altitude "250 m" in a paratype from MHNG is definitely a labeling mistake 
(G. Cuccodoro, pers. comm.) 

Anthelephila besucheti Bonadona, 1989 Figs 16-18 

Anthelephilus besucheti Bonadona, 1989: 263, figs 6, 14, 16-18. 

Type material: Holotype o\ CEYLAN Southern Yala nat. park 24.1.1970 Mussard 
Besuchet Lobi [p+h] Holotype [p; red label] \ Anthelephilus besucheti nsp. RBonadona dét. 1976 
[p+h] (MHNG). - Paratypes: 1 o\ 2 ? 5, same data as holotype (MHNG, 1 9 ZKDC). -13, 



622 



Z. KEJVAL 




Figs 8-15 
Anthelephila aratrix sp. n., male: (8) elytron. (9) sternum Vu, ventral view. (10) the same, lateral 
view. (11) tergum VII. (12) segment VIII, dorsal view. (13) the same, lateral view. (14) tegmen. 
(15) median lobe of aedeagus. Scale (1 mm): A - Fig. 8; (0.5 mm): B - Figs 14, 15, C- Fig. 11, 
D- Figs 9, 10, 12, 13. 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 



623 




Figs 16-25 

Anthelephila besuchen Bonadona, male: (16) sternum VII. (17) sterilite VIII (half). (18) tegmen. 

A. cardamontis sp. n., male: (19) sternum VII. (20) sternite VIII (half). (21) tegmen. A. irula 

sp. n., male: (22) sternum VII. (23) sternite VIII (half). (24) tergite VIII. (25) tegmen. Scale 

(0.2 mm): A - Fig. 20, B - Fig. 16, C - Figs 17, 18, 23, 25, D - Figs 19, 21, 22, 24. 



624 z - KEJVAL 

1 9 , CEYLAN Northern, Mullaittivu 6.II.70 Mussard Besuchet Lobi [p+h] \ Paratype [p; red 
label] \ Anthelephilus besucheti nsp. P.Bonadona dét. 1976 [p+h] (MHNG). -lo", CEYLAN 
North western Rajakadaluwa 31.1.1970 Mussard Besuchet Lobi [p+h] \ Paratype [p; red label] \ 
Anthelephilus besucheti nsp. P.Bonadona dét. 1976 [p+h] (MHNG). According to Bonadona 
(1989), the three paratypes from Yala and Mullaittivu are deposited also in the coll. Bonadona 
(MNHN; not examined). 

Redescription: Body length 2.8-3.5 mm (holotype 2.8 mm). 

Male (paratype, Yala): Mesosternum, metasternum and all legs simple. 
Sternum VII (Fig. 16) nearly simple, slightly produced and bluntly pointed postero- 
medially, its apical margin with numerous short, stiff setae. Tergum VII moderately 
widely rounded posteriorly, with apical margin slightly emarginate medially in dorso- 
caudal view. Sternite VIII (Fig. 17); paired prongs rather simple, dorso-ventrally 
flattened, conspicuously wide, strongly narrowed, convergent and rounded apically; 
each prong with five, conspicuously long, thick setae dorsally and about eight shorter, 
thick setae ventrally, along lateral margin. Tergite VIII simple, its posterior margin 
rather widely rounded, sparsely setose. Aedeagus (Fig. 18); apical portion of tegmen 
0.6 times as long as basal-piece, trilobed apically, middle lobe wide, rounded apically, 
distinctly shorter and wider than narrow lateral lobes. 

Female (paratype, Yala): Externally identical with male, except as follows: both 
sternum and tergum VII evenly rounded posteriorly, sternum impressed ventro- 
medially and with apex moderately bent ventrad. 

Differential diagnosis: Anthelephila besucheti is closely related to A. irula 
sp. n. from southern India. For their separation see the key and the differential 
diagnosis of the latter species. 

Distribution: Sri Lanka. 

Anthelephila cardamontis sp. n. Figs 1, 19-21 

Type material: Holotype o\ S-INDIA, Kerala state, 10 km SW of Kumily, Vallakadavu 
vili, env., 77°07'E 9°31'N \ Cardamom hills, alt. ca 1000 m, 24.xii.1993, sifted, Z. Kejval & D. 
Boukal Igt. (NMPC). - Paratypes: 7 S3, same data as holotype (ZKDC, 1 specim. in MHNG). 

Etymology: Named after the type locality; composed of cardamom 
(Cardamom hills Mts.) and Latin montium/montis (mountain ridge). 

Description: Body length 3.4-3.8 mm (holotype 3.4 mm). 

Male (holotype): Body rufous, head moderately darker, elytra with two pairs of 
vaguely outlined, brown black spots (Fig. 1); anterior spots circular, situated at about 
midlength, distinctly separated from both suture and lateral margins, posterior spots 
situated apically, slightly touching latero-apical margins; antennae, legs and palpi 
rufous. 

Head: 1.3 times as long as wide, longitudinally oval, nearly evenly rounded 
posteriorly in dorsal view; posterior temporal angles entirely obsolete. Surface smooth, 
very glossy; dorsal punctation distinct but rather sparse and uneven. Setation short, 
mostly subdecumbent, with scattered suberect to erect, slightly longer setae. Eyes 
medium sized, moderately convex. Antennae rather long, clearly exceeding base of 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 625 

pronotum, moderately but distinctly enlarged in apical third; antennomere X 1 .2 times 
and antennomere XI 2. 1 times as long as wide. 

Thorax: Pronotum 1.2 times as long as wide, slightly narrower than head 
including eyes, unevenly, somewhat widely rounded anteriorly, strongly and nearly 
straightly narrowing posteriad, and shallowly impressed postero-laterally close before 
base (not constricted posteriorly in dorsal view); entire dorsal outline of pronotum 
convex, only slightly impressed close before basal margin; pronotai disc evenly 
shaped, without longitudinal impressions. Surface largely smooth and very glossy, 
postero-lateral impression shortly, distinctly wrinkled and this basal corrugation 
extending also somewhat dorso-laterally; dorsal punctation as on head, somewhat 
denser mesally. Setation as on head. Both mesosternum and metasternum simple. 

Elytra: 1.7 times as long as wide, longitudinally ovoid, strongly convex, 
narrowed and rather conjointly rounded apically; humeri entirely obsolete; 
postscutellar impression absent. Surface smooth, very glossy; punctation somewhat 
finer and distinctly sparser than on head. Setation distinctly longer than on head, and 
generally more raised, mostly decumbent, with numerous erect setae. Metathoracic 
wings nearly entirely reduced. 

Legs: Penultimate tarsomeres rather narrow, with terminal tarsomere articulated 
before midlength; all legs simple. Setation normally developed, inconspicuous. 

Abdomen: Sternum VII (Fig. 19) rather simple, somewhat truncate and with 
posterior margin slightly sinuous, sparsely setose, with some longer, mostly scattered 
stiff setae. Tergum VII narrowed and nearly evenly rounded posteriorly. Sternite VIII 
(Fig. 20); paired prongs wide and nearly parallel-sided in basal half, angulately 
protruding laterad at about mid-length and then strongly narrowed and converging, 
with apex laterally flattened and rounded; surface of prongs finely setose along ventral 
margin of their apical narrowed portion, and with tuft-like accumulation of longer, stiff 
setae on/near lateral protrusions. Tergite VIII simple, arcuate, with posterior margin 
evenly rounded, sparsely setose. 

Aedeagus (Fig. 21): Apical portion of tegmen 0.4 times as long as basal-piece, 
trilobed apically, middle lobe wide basally, strongly narrowing towards blunt apex, 
slightly shorter than evenly narrow, lateral lobes. 

Female: Unknown. 

Variability: Some of the examined specimens are generally more pale in 
colour and with very fine punctation on the surface (probably teneral specimens). 

Differential diagnosis: Anthelephila cardamontis sp. n. may resemble 
A. irula sp. n. and A. besucheti by the evenly sparse punctation of the elytra and the 
simple metasternum in males. It differs from these species especially by the paler 
colouration, the presence of dark spots on the elytra, the smooth and glossy surface of 
both the head and pronotum, and in the morphology and setation of male sternum VII 
and sternite VIII. 

Distribution: India (Kerala). 

Comments: The specimens were collected by sifting forest litter. 



626 Z. KEJVAL 

Anthelephila congoana Uhmann, 1981 Figs 26-29, 34 

Anthelephilus congoanus Uhmann, 1981: 193, figs 4-9. 

Type material: Holotype S, Soil-Zoological Exp. Congo-Brazzaville Sibiti Irho rain 
forest [p] \ 2.12.1963. No331 beaten in forest leg. Balogh & Ziczi [p] \ Holotypus 1980 
Anthelephilus congoanus Uhmann [p+h; red frame] \ Typus [p; red label] \ Anthelephilus 
congoanus n.sp. det.G.Uhmann 1980 [p] (HNHM). - Paratypes: 1 5, 331 [p; bluish label] \ 
Congo-Exped. d. Inst. Syst. Zool. Budapest 16.X.63. - 21.1.64. Fundort No.: 331 [p+h] \ Sibiti, 
Irho oil-palm plantat. 2. XII. 1963 leg. Balogh / Zicsi [p+h] \ Para-typus [p; red label] \ 
Anthelephilus congoanus n. sp. det. G. Uhmann 1980 [p] \ Paratypus 1980 Anthelephilus 
congoanus Uhmann [p+h; red frame] (DCDC). - 1 6, Congo-Exped. d. Inst. Syst. Zool. 
Budapest 16.X.63. - 21.1.64. Fundort No.: 261 [p+h] \ Sibiti, Irho rain forest K 27.1 1.1963 leg. 
Balogh / Zicsi [p+h] \ 261 [p; bluish label] \ Paratypus 1980 Anthelephilus congoanus Uhmann 
[p+h; red frame] \ Para-typus [p; red label] \ Anthelephilus congoanus n. sp. det. G. Uhmann 
1980 [p] (DCDC). - 1 â, Soil-Zoological Exp. Congo-Brazzaville Sibiti Irho rain forest \ 
26.1 1.1963. No 249 singled on fallen fruits leg. Balogh & Zicsi \ Paratypus 1980 Anthelephilus 
congoanus Uhmann \ Para-typus \ Anthelephilus congoanus n.sp. det.G.Uhmann 1980 [p] 
(HNHM). -19, Soil-Zoological Exp. Congo-Brazzaville Kindamba, Méya Bangou forest \ 
9.11.1963. No 141 singled fallen fruits in forest leg. Endrödy-Younga [the 3th to 5th label the 
same] (HNHM). - 2 ââ, 2 9 9, Soil-Zoological Exp. Congo-Brazzaville Lefinie reservation 
Nambouli river \ 1 1.1.1964. No 652 beaten in galery forest leg. Balogh & Ziczi [the 3rd to 5th 
label the same] (HNHM, ZKDC). - 1 9 , Congo-Exped. d. Inst. Syst. Zool. Budapest 16.X.63.- 
21.1.64. Fundort No.: 597 \ Lefinie, Reservat. Nambouli River, 1.7. K. bushes/forest 1964 leg. 
Balogh/Zicsi \ 597 [the 3th to 5th label the same] (HNHM). 

Other material examined: 2 6 o\ 2 9 9 , Uganda, Ruwenzori Mts., above Bundibugyo, 
1300m, 2 I.V. 1993, Cuccodoro & Erne leg. (MHNG, 1 specim. ZKDC). - 1 6, Uganda, 
Ruwenzori, above Kilembe, 1950 m, 4.V.1993, Cuccodoro & Erne leg. (MHNG). - 1 9, W- 
Uganda, SW of Hioma, Rwera env., 30.xi.2001, M. Snizek leg. (ZKDC). 

Redescription: Body length 3.5-4.3 mm. 

Male (paratype, ZKDC): Mesosternum simple; metasternum with a pair of 
strong, apically setose protrusions posteriorly, near median margin of metacoxae. All 
legs simple and normally setose; basal protarsomere enlarged, rather long and wide. 
Sternum VII (Fig. 26) with posterior margin distinctly sinuous, its middle lobe wider, 
slightly upturned apically and more lengthily setose. Tergum VII evenly rounded 
posteriorly. Sternite VIII (Figs 27, 28); prongs rather simple, narrowed and moderately 
converging distally in dorsal view, their ventral margin/edge coarsely denticulate at 
about midlength and subapically, apex of prongs somewhat obliquely truncate and 
pointed; setation of prongs less conspicuous, rather fine and scattered, with some 
longer setae apically. Tergite VIII with posterior margin moderately emarginate 
medially. Aedeagus (Fig. 29); apical portion of tegmen 0.8 times as long as basal-piece, 
trilobed apically, all three lobes narrowing towards apex, middle lobe slightly longer 
and more pointed. 

Female (paratype, DCDC): Externally identical with male, except as follows: 
metasternum simple, lacking paired protrusions; basal protarsomere of smaller size; 
sternum VII simple, its posterior margin slightly unevenly rounded and bearing a tuft 
of dense, longer setae medially; tergum VII (Fig. 34) strongly narrowed, tapering and 
longer, densely setose apically. 

Variability: Moderately variable in characters of corrugation and micro- 
sculpure of the head and pronotum. Some of the specimens from Uganda differ in the 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 



627 




c •- 



Figs 26-33 
Anthelephila congoana Uhmann, male: (26) sternum VII. (27) sternite VIII (half). (28) prong of 
sternite VIII (outline), lateral view. (29) tegmen. A. ovipennis (Bonadona), male: (30) sternum 
VII. (31) sternite VIII (half). (32) prong of sternite VIII (outline), lateral view. (33) tegmen. 
Scale (0.2 mm): A - Figs 27, 28, 31, 32, B - Fig. 29, C - Figs 26, 30, 33. 

more distinct corrugation of the frontal surface of the head, extending posteriad along 
median margins of eyes, and/or in the less glossy, finely microsculptured surface of the 
pronotum, especially on the lateral sides. 



628 Z. KEJVAL 

Differential diagnosis: Anthelephila congoana is most closely related to 
A. ovipennis, as suggested especially by the very similar form of male sterilite VIII, but 
differs by the head being somewhat unevenly rounded and slightly produced postero- 
medially, the simply setose male metatibiae, male sternum VII sinuous apically with a 
wide middle lobe, male tergite VIII moderately emarginate apically, female sternum 
VII tapering apically, as well as by some details in morphology of male sternite VIII 
and the tegmen. 

Distribution: Zaire, Uganda. 

Anthelephila curvitarsis sp. n. Figs 3, 38-43 

Type material: Holotype o\ S. Afr: Kruger Nat. Pk Pumbe sands 24.12 S - 31.55 E \ 
22.11.1994; E- Y: 3063 groundtraps Endrödy, Bellamy \ ground traps with meat bait (TMP). - 
Paratypes: 1 S , same data as holotype (ZKDC); 1 9, S. Afr: Little Karoo Gamkaberg, 1000 m 
33.44 S - 21.57 E \ 21.12.1993; E- Y: 3069 ground traps, 24 days leg. Endrödy- Younga \ 
groundtrap with banana bait (TMP). - 1 S, 4 9 9, SOUTH AFRICA: KZN Tembe Elephant 
Park, Sihangwane Area 27.02S 32.25E 100 m 03.ii.1996 R. Stals \ Habitat: Sand forest Sieved 
from forest litter \ National coll. of insects Pretoria, S. Afr. (SANC, 1 9 ZKDC). 

Etymology: Composed from Latin curvatus (curved) and tarsus; named in 
reference to the curved basal protarsomere of the males. 

Description: Body length 2.8-3.6 mm (holotype 3.6 mm). 

Male (holotype): Body uniformly black; legs dark brown, base of tibiae and 
tarsi paler, rufous brown, antennae and palpi dark brown, basal and apical antennome- 
res slightly paler. 

Head: 1.3 times as long as wide, longitudinally oval, with base rather evenly 
rounded in dorsal view; posterior temporal angles entirely obsolete. Surface with 
extremely fine, dense, net-like microsculpture and thus only moderately glossy; dorsal 
punctation distinct, but somewhat obscured by microsculpture, uneven, punctures 
rather shallow, separated by more than their diameter. Setation inconspicuous, very 
short, fine, mostly appressed, with few short, erect setae. Eyes small, slightly convex. 
Antennae moderately exceeding base of pronotum, distinctly enlarged in apical third; 
antennomere X slightly transverse, 0.95 times as long as wide, antennomere XI 1.4 
times as long as wide. 

Thorax: Pronotum 1.3 times as long as wide, moderately narrower than head 
including eyes, evenly round/globose anteriorly in dorsal view, strongly narrowing pos- 
teriad and distinctly impressed postero-laterally (constricted posteriorly in dorsal view, 
Fig. 3); entire dorsal outline of pronotum more or less convex in lateral view; prenotai 
disc with shallow median longitudinal furrow in anterior and posterior thirds. Surface 
only moderately glossy, with the same microsculpture as head, bottom of postero-lat- 
eral impressions very finely wrinkled; dorsal punctation slightly coarser than on head, 
especially mesally and dorso-laterally in posterior third. Setation as on head, 
inconspicuous. Mesosternum simple. Metasternum with a pair of distinct longitudinal 
protrusions posteriorly, near median margin of metacoxae, their edge covered with 
brownish, felt-like setation. 

Elytra: 1.9 times as long as wide, ovoid, strongly convex, narrowed and trun- 
cate apically; humeri entirely obsolete; postscutellar impression absent. Surface some- 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 



629 




Figs 34-43 
(34) Anthelephila congoana Uhmann, female tergum VII. A. ovipennis (Bonadona), male: (35) 
mesosternum and adjacent sclerites. (36) metatibia. (37) female tergum VII. A. curvitarsis sp. n., 
male: (38) metatibia. (39) protibia with tarsus. (40) sternum VII. (41) sternite VIII (half). (42) 
prong of sternite VIII, lateral view. (43) tegmen. Scale (0.2 mm): A - Figs 41, 42, B - Figs 
34-39, C - Fig. 43, D - Fig. 40. 

what uneven and less glossy; punctation similar, only moderately sparser than on head 
and pronotum. Setation similar to that on head, very short and inconspicuous, with very 
vague indication of two transverse bands of slightly thicker, whitish setae. 
Metathoracic wings almost entirely reduced. 



630 z - KEJVAL 

Legs: Penultimate tarsomeres rather narrow, terminal tarsomere subapically in 
metatarsi; metatibiae moderately sinuous, apically strongly angulately produced on in- 
ner side (Fig. 38); basal protarsomere conspicuously long, distinctly curved (Fig. 39). 
Setation normally developed, inconspicuous. 

Abdomen: Sternum III with shallow longitudinal impression postero-medially. 
Sternum VII (Fig. 40) deeply emarginate posteriorly, its postero-median margin long 
and densely setose. Tergum VII evenly rounded posteriorly. Sternite VIII (Figs 41, 42): 
paired prongs short and wide in dorsal view, strongly widened in apical half in lateral 
view, hollowed dorso-medially, their ventral median margin sinuous and bearing 
numerous densely spaced, stiff setae, wide apex of prongs conspicuously long and 
richly setose. Tergite VIII arcuate, nearly evenly rounded posteriorly. 

Aedeagus (Fig. 43): Apical portion of tegmen 0.5 times as long as basal-piece, 
trilobed apically, middle lobe moderately widening towards subtruncate apex and with 
small apical protuberance, as long as and apically wider than lateral lobes. 

Female: Externally identical with male, except as follows: antennae slender, 
less enlarged distally; metasternum simple, lacking paired protrusions; metatibiae 
simple, rather straight, basal protarsomere much shorter and straight; sternum III 
evenly convex, lacking impression; sternum VII simple, its posterior margin quite 
evenly rounded. 

Differential diagnosis: Of the related Afrotropical species, Anthelephila 
curvitarsis sp. n. may resemble A. congoana in having a more elongate and posteriorly 
rather oval head, but differs clearly by the pronotum being strongly impressed postero- 
laterally and distinctly constricted in dorsal view, and by all the male characters (e.g. 
long and curved basal protarsomere, moderately sinuous, angulately produced and and 
simply setose metatibiae apically on inner side, quite different forms of sternum VII 
and sternite VIII). 

Distribution: South Africa. 

Anthelephila imperatrix LaFerté-Sénectère, 1849 Figs 2, 6, 7, 44-53 

Anthelephilus imperator LaFerté-Sénectère, 1849a: 2, fig. 2. 

Anthelephilus imperator LaFerté-Sénectère, 1849b: 66, fig. 2; see Chandler (2000). 

Anthelephila imperatrix: Krekich-Strassoldo, 1931: 15, fig. 30. 

Anihicus formicarius Nietner, 1856: 533. 

Anihicus quisquiliarius Nietner, 1857b: 20. 

Anihicus myrmecodes Gemminger, 1870: 123. 

Formicomus (Anthelephilus) imperator var. ruficolor Pic, 1916a: 5. 

Formicomus cribriceps Marseul, 1876: 459. 

Formiconus [sic!] (Anthelephilus) cribriceps: Miwa, 1931: 180 (misspelling). 

Anthelephila cribraceps [sic!]: Hua, 2002: 131 (misspelling). 

Formicomus (Antelephilus [sic!]) punctaticeps Pic, 1916b: 11, syn. n. 

Type material: Anthelephilus imperator - Holotype $ [lacking distal antennomeres and 
foreleg]: 47394 [p] \ Linga Rottg. [h; grey label] \ imperator Laf.*typ. [h; grey label] \ Type [p; 
reddish label] \ Hist. -Coll. (Coleoptera) Nr. 47394 Anthelephilus imperator Laf. Linga. Roettger 
Zool. Mus. Berlin [p; yellow label, black frame] (MNHB). Formicomus punctaticeps - 
Syntypes: 1 o\ Kulu [p] \ type [h; yellowish label] \ TYPE [p; red label] \ Muséum Paris Coll. 
M. Pic [p] \ Anthelephilus punctaticeps Pic [h] (MNHN). - 1 6, Kulu [h] \ Museum Paris Coll. 
M. Pic [p]. F imperator var. ruficolor - Syntypes: 2 6 S , Manille ... [h; partly illegible] \ type 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 



631 




Figs 44-53 
Anthelephila imperatrix LaFerté-Sénectère (Indonesia): (44) male sternum VII. (45) male 
sternite VIII (half). (46) prong of male sternite VIII, lateral view. (47) male sternite VIII, antero- 
dorso-lateral view. (48) male tergite VIII (49) tegmen. (50) apical portion of female abdomen, 
lateral view. A. imperatrix, male (Japan): (51) sternum VII. (52) sternite VIII (half). (53) prong 
of sternite VIII, lateral view. Scale (0.2 mm): A - Fig. 49, B - Figs 44-46, 48, C - Figs 52, 53, 
D - Figs 47, 51; (1 mm): E - Fig. 50. 



632 z - KEJVAL 



[h] \ TYPE [p; red label] \ v. ruficolor Pic [h]; 4 specimens, bearing label "Manille" or "Manille 
Baer", but lacking the type labels (MNHN, coll. Pic). Formicomus cribriceps - Syntypes: 1 6 , 
1 9 [mounted on common label]: Formicomus cribriceps Japon ... [h; yellowish round label; 
partly illegible]; 5 specimens, mounted in the same way, but lacking the locality label (all 
MNHN). 

Other material examined: INDONESIA: 2 9 9, Sumatra, Aceh-Selatan Prov., 
Babahrot, 100 m, 7.VÜ.1983, J. Klapperich leg. (SMNS). - 1 9, C Sulawesi, 17 km E Pendolo, 
800 m, 120.45.49 E 2.06.33 S 4.-9.VÜ.1999, Bolm leg. (SMNS). - 3 66, 3 9 2, Lombok, 
Sembalun Lawang, Mt. Rinjani, 1700 m, 6.-8.Ü.1995, Bolm leg. (SMNS, ZKDC). - 1 9, 
Lombok, Senggigi Pemenang, 20. xi. 1999, E. Heiss leg. (ZKDC). -299, Ceram Island, Wahai 
env., 12.ii.1989, M. Jäch leg. (NHMW). - 5 6 6, 2 9 9, Sula Islands (E of Sulawesi), Mangole 
Island, vii.-xii.1977, V. & G. Wegener leg. (NHMB, ZKDC). PHILIPPINES: 1 6,2 9 9, 
Mindanao, Maramag Prov., Portulin, 750 and 1700 m, 3.-4.U991, Bolm leg. (NHMB). 
VIETNAM: 1 6, Hanoi, riverbank, 26.ix.1980, F. Hieke leg. (GUPC). - 1 6, SE of Hanoi, Yen 
So, 19.-23.iv. 1966, G. Topâl leg. (ZKDC). - 3 9 9, Da Nang, 2.-3.xi.l970, A. R. Gillogly leg. 
(GUPC, DCDC). USA (pacific territories): 2 6 6, 2 9 9, Mariana Islands, Saipan Island, 
28.viii.1951, R. M. Bohart leg. (DCDC, ZKDC). - 1 9, Saipan Island, vi. 1958, N.L.H. Krauss 
leg. (DCDC). - 1 ó\ 1 9, Mariana Islands, Guam Island, Piti, 18.vii.1936, O. H. Swezey leg. 
(BMNH). - 1 9, Guam Island, Yona, 21.iv.1936, E. H. Bryan leg. (BMNH). - 2 9 9, Guam 
Island, Pago Bay, 2.vi.l945, H. S. Dybas leg. (DCDC). - 1 9, Guam Island, ca 1.5 km SE of 
Asan, 180-250 m, 5.xi.l947, H. S. Dybas leg. (DCDC); 1 9, Mariana Islands, Tinian Island, 1.- 
14.iv.1945, H. S. Dybas leg. (DCDC). - 1 6, Palau Islands, Koror Island, 24.xi.1947, H. S. 
Dybas leg. (DCDC). - 2 6 6, 1 9 , Hawaii, Honolulu, Internat. Airport, 19.ii.1968, G. Funasaki 
leg. (DCDC, GUPC). JAPAN: 3 S 6 , 2 9 9 , G. Lewis leg. (BMNH). - 5 S 6 , 14 9 9 , Kyushu, 
Oita, Reitter (NMPC, ZKDC). -2 6 6,1 9 , Honshu, Kobe, Harada, 23.ix.1915, J. E. Lewis leg. 
(BMNH). - 1 9, Honshu, Kobe, Hyogo, 18.iv. 1931 (NMPC); series of specimens, Honshu, Idzu, 
vi.1910, S. Akiyama leg. (BMNH). - 1 6,2 9 9, Okinawa Island, Motobu Peninsula, W slope 
of Katsu Dake, ca 800 m, 28.ix.1945, F G. Werner leg. (DCDC, ZKDC, GUPC). - 1 6, 1 9, 
Okinawa, Yogi, Naha, lO.viii. 1951, F. G. Werner leg. (DCDC). CHINA: 1 ó\ NE Zhejiang Prov., 
Chusan [=Zhoushan] Archipelago, Entrance Island, Walker leg. (BMNH). - 1 9 , N Zhejiang 
Prov., ca 100 km SW of Shanghai, Haining, Walker leg. (BMNH). -19, Fukien Prov., Kuatun, 
10.V.1946, Tschung Sen. leg. (SMNS). -16, Guangxi Prov., Gul Lin, 30.xii.1981, Rougemont 
leg. (coll. Bonadona, MNHN). TAIWAN: 1 6, Akau, E. Csiki coll. (HNHM). NEPAL: 2 6 6, 
Kathmandu, Baneshwar, 1300 m, 21.-25.vi.1998, W. Schawaller leg. (SMNS, ZKDC). - 1 6, 
same data, except: 1350 m, 20.-2 l.v.2000 (SMNS). -16, Kathmandu valley, Pashupatinath, 
Bagmati River, 1300 m, 14.X.1992, A. Weigel leg. (NKME). - 1 o\ 1 9, NE of Kathmandu, 
Gorkana Park, near Bagmati river, 27°43'N 85°23'E, 29.ix.1996, M. Hartman leg. (NKME, 
ZKDC). -2 6 6, same data, except: 1300-1400 m, 28.V.1997, Grill/Hartmann leg. (NKME). - 
6 6 6, 3 9 9, same data, except: 1280 m, 24.xi.1998 (NKME). -166,1 9 9, same data, 
except: 1340 m, 17.vi.1999 (NKME). - 1 6,1 9, same data, except: Bagmati riverbank, 1350 
m, 15.6.2001, A. Kopetz leg. (NKME). -4ó*c?,2 99,6km SSW of Kathmandu, bank of 
Taudaha Lake, 27°39'N 85°09'E, 1300 m, 17.vii.2001, M. Hartmann leg. (NKME, ZKDC). - 1 
o\ Dhaulagiri Himal, Kali Gandaki valley, Jhi vili. N Beni, 1750 m, 16.vi.1998, Berndt & 
Schmidt leg. (ZKDC). - 1 6,1 9 , Annapurna Mts., S of Ulleri, Ghorepani, 2000 m, 16.vi.1993, 
Schmidt leg. (ZKDC). - 1 9, Kathmandu Prov., Godwari, 1600 m, 31.iii.1984, Löbl leg. 
(MHNG). - 1 9, E-Nepal, Arun valley, Chichila, 1950 m, 31.V.1983, M. Brancucci leg. 
(NHMB). - 1 9, E-Nepal, Biratnagar, 140 m, 21.V.1980, W. Wittmer leg. (NHMB). INDIA: 1 
6, Meghalaya, Shillong (NHMW). - 3 9 9, West Bengal, Nagarkanda, 3.-14.U980, G. Topâl 
leg. (GUPC). - 1 9, West Bengal, Calcutta, 6.VÜ.1972, Basel Exped. 1972 (coll. Bonadona, 
MNHN). - 3 6 6, 1 9, Orissa, Ganjam Prov., N of Berhampur (=Brahmapur), Kalasandrapur 
env., 20.-2 l.ii. 1994, Z. Kejval leg. (ZKDC). - 1 o\ 1 9, Uttaranchal, W of Almora, H. G. 
Champion leg. (BMNH). - 4 6 6. 5 9 9 , Uttaranchal, 22 km N of Rishikesh, 450 m, 30.x. 1979, 
I. Lobi leg. (MNHG, GUPC). - 1 6,3 9 9, Uttaranchal, 16 km of Srinagar, 550 m, 29.x. 1979, 
I. Lobi leg. (MNHG. ZKDC). - 26 6 6, 29 9 9 , Uttaranchal, ca 13 km NW of Nainital, Khairna 
Bridge env., 900-1000 m, 13.-17.vn.2003, Z. Kejval & M. Tryzna leg. (ZKDC). - 8 6 6, 16 9 9 , 
Uttaranchal, 10 km NE of Rishikesh, Henval river valley, Shivpuri env., ca 450 m, 26. vii. 2003, 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 633 



Z. Kejval & M. Tryzna leg. (ZKDC). - 1 6 , 4 5 2 , Uttaranchal, 20 km NE of Rishikesh, Ganga 
river valley. Kaudiyala env., ca 500 m, 25. -27. vii. 2003, Z. Kejval & M. Tryzna leg. (ZKDC). - 

1 6, Uttaranchal Haldwani-Kathgodam, ca 800 m, 21.-22.vii.2003, Z. Kejval & M. Tryzna leg. 
(ZKDC). - 2 6 6, 5 2 2 , Uttaranchal, 30 km N of Rishikesh, NW of Chamba, Arakot env., 1500 
m, 29.-3 l.vii.2003, Z. Kejval & M. Tryzna leg. (ZKDC). -5 6 6\ J 2 5, Uttaranchal, 25 km W 
of Mussoorie, Yamuna river valley, Juido env., ca 750 m, 5. -7. vii. 2003, Z. Kejval & M. Tryzna 
leg. (ZKDC). - 1 1 6 6,1 9 9 , Uttaranchal, Dehra Dun, 12.iii. 1952, R. N. Kothari leg. (DCDC). 
-19, Uttaranchal, Baijnath, 26.vii.-28.vii.2003, E. Kucera leg. (ADBC). - 8 66,5 99, 
Uttaranchal, Gangani, 1250 m, 13.-20.vi.1981, M. Brancucci leg. (NHMB). - 1 6, 1 9, 
Uttaranchal, Barkot, 1000-1200 m, 5.-12.vi.l981, M. Brancucci leg. (NHMB). - 1 9, Rajasthan. 
SE of Bharatpur, Keoladeo Ghana Nat. Park, 27°09'N 77°31'E, ca 170 m, 6.ix.l985, C. W. & L. 
B. O'Brien leg. (DCDC). - 1 9, 10.-12.viii.1989, Hiermeier leg. (ZKDC). - 14 6 6, 16 9 9, sa- 
me locality, 13. -14. vii. 2006, Z. Kejval leg. (ZKDC). - 1 9, Rajasthan, Udaipur, Sajjan Niwas 
Gardens, 24°34'N 73°41'E, 600 m, 3.-8.VÜ.2006, Z. Kejval leg. (ZKDC). -2 66, Maharashtra, 
ca 15 km E of Savantvadi, 15°55'N 75°53'E, riverside, ca 40 m, 22.V.2006, Z. Kejval leg. 
(ZKDC). - 1 6, Goa, Salcete, 13.-16.iv. 1990, E. Heiss leg. (ZKDC). - 1 6, 1 2, Karnataka, 
Ablathi, 12°17' N 76°06' E, x.1984, W. Lorenz leg. (GUPC). - 2 6 6, 3 9 9, Kerala, Palghat 
hills, Malampuzha Dam, 150 m, 27.xi.1972, Besuchet, Lobi & Mussard leg. (MHNG, coll. 
Bonadona MNHN). - 1 9, Kerala, Palghat-Coimbatore, Walayar Forest, 400 m, 23.xi.1972, 
Besuchet, Lobi & Mussard leg. (MHNG). - 8 6 6, 5 2 2, Tamil Nadu, Nilgiri hills, 15 km SE 
of Kotagiri, Kunchappanai env., 76°56' E 11°22' N, ca 900 m, 13.-20.V.1994, Z. Kejval leg. 
(ZKDC). SRI LANKA: 16, 1 2, Kandy, Mahaweli Ganga riv., 450-500 m, 30.i.-l.ii.l970, 
Besuchet, Lobi & Mussard leg. (MHNG).- 1 6, Nuwara Eliya, 18.ix.1963 (BMNH). -2 6 6,4 

2 9. 12.iv.1882, G. Lewis leg. (BMNH). - 2 66, 1 9, Kandy, near Mahaweli Ganga river, 
23.iii.1994, Z. Kejval leg. (ZKDC). - 1 9, Galle Prov., Habaraduwa, 20.8.-4.9.1982, H. J. 
Bremer leg. (GUPC). PAKISTAN: 1 6,5 9 9, Swat, Col de Karakar, 1300 m, 19.V.1983, 
Besuchet & Lobi leg. (MHNG). -2 66,1 9 9, Swat, Madyan, 1400 m, 16.V.1983, Besuchet & 
Lobi leg. (MHNG). - 1 6,3 9 9, Swat, Jowar, 1100 m, 19.V.1983, Besuchet & Lobi leg. 
(MHNG). - 1 6, Swat, Manglaur, 1150 m, 9.V.1983, Besuchet & Lobi leg. (MHNG). -16,2 
9 9 , Hazara, Balakot, 900 m, 4.vi.l983, Besuchet & Lobi leg. (MHNG). - 4 6 6, 2 9 9 , Dir, 
Dir, 1500 m, 20.V.1983, Besuchet & Lobi leg. (MHNG). -2 6 6, 1 9, Punjab, Rawalpindi, Ayub 
Nat. Park, 28.viii.1985, C. W. & L. B. O'Brien leg. (DCDC, ZKDC). -16, Punjab, Rawal Lake 
Dam, 1. ix. 1965, C. W. & L. B. O'Brien leg. (DCDC). 

Redescription: Body length 2.7-3.7 mm. 

Male (Indonesia, Lombok, ZKDC): Head and pronotum rufous; elytra largely 
rufous brown, with rufous base; legs, basal antennomeres and palpi rufous. 

Head: 1.1 times as long as wide, globose, evenly to somewhat widely rounded 
posteriorly; posterior temporal angles indistinct, rounded. Eyes small, moderately con- 
vex. Surface smooth, glossy, conspicuously punctured; dorsal punctures large and rat- 
her shallow, unevenly spaced, separated mostly by less than their diameter, at places 
sparser, especially postero-medially. Setation evenly short, mostly subdecumbent to 
appressed, with sparsely scattered, short erect setae. Antennae short, slightly exceeding 
base of pronotum, moderately enlarged in apical third; antennomere X slightly, 1.1 ti- 
mes, and antennomere XI 1 .7 times as long as wide. 

Thorax: Pronotum 1.3 times as long as wide, much narrower than head 
including eyes, moderately widely rounded anteriorly, strongly narrowing posteriad 
and distinctly impressed postero-laterally (constricted) in dorsal view (Fig. 2); dorsal 
outline of pronotum convex in anterior two thirds, then impressed and distinctly 
bulging before base in lateral view (Fig. 6); pronotai disc anteriorly with conspicuous 
median longitudinal impression/groove, apex of antebasal bulge vaguely divided 
medially by shallow median impression. Surface smooth, glossy, posterior constriction 



634 Z. KEJVAL 

of pronotum distinctly wrinkled laterally to dorso-laterally; dorsal punctation uneven, 
generally much finer and sparser than on head, more distinct at dorsal, convex places 
alonside longitudinal impression, while bottom of impression appears to be nearly 
impunctate. Setation as on head. Both mesosternum and metasternum simple. 

Elytra: 1.7 times as long as wide, convex, clearly truncate apically; humeri 
entirely obsolete; postscutellar impression absent. Surface glossy; punctation uneven, 
ordinary punctures generally much finer and sparser than on head; in addition with fine, 
densely spaced, nearly contiguous punctures forming two paired oblique bands 
(Fig. 2); anterior bands narrow, situated in basal third, directing postero-mediad from 
lateral sides, posterior bands situated shortly behind mid-length, wider, more conspi- 
cuous, directing antero-mediad from lateral sides. Setation mostly as short as on head, 
subdecumbent to appressed, with scattered longer, erect setae, mostly pale, with setae 
of dense punctures contrastingly whitish, thicker, quite appressed and forming distinct 
setose bands. Metathoracic wings almost entirely reduced. 

Legs: Penultimate tarsomeres rather narrow, terminal tarsomere rather subapical 
in metatarsi; all legs simple. Metatibiae rather distinctly and densely punctured, 
especially on inner side; setation normally developed, slightly denser and more raised 
on metatibiae. 

Abdomen: Sternum VII (Fig. 44) truncate posteriorly, its posterior margin near- 
ly straightly sloping towards small median notch/impression in ventral view. Tergum 
VII with posterior margin nearly evenly rounded, surface subapically somewhat more 
convex and with indication of rounded, median longitudinal edge. Sternite VIII (Figs 
45, 46); paired prongs narrow in dorsal view, with lobe-like dilatation in basal half 
ventrally in lateral view, arcuately curved ventrad in apical half, truncate and axe-like 
shaped apically in lateral view, their apical widened portion with three points/ protru- 
sions (Fig. 47); surface of prongs with several longer, stiff setae at apex of ventral di- 
latation. Tergite VIII (Fig. 48) simple, widely rounded posteriorly. 

Aedeagus (Fig. 49). Apical portion of tegmen 0.5 times as long as basal-piece, 
parallel-sided, trilobed apically, middle lobe nearly evenly narrowing towards bluntly 
pointed apex, about as long as apically wider, rounded lateral lobes. 

Female (Indonesia, Lombok, ZKDC): Externally identical with male, except as 
follows: sternum VII simple, evenly rounded posteriorly; tergum VII clearly modified 
(Fig. 50), projecting medially, subapically, and forming conspicuous, laterally flattened 
process, exceeding apical margin of tergum, surface of tergum alongside subapical 
process distinctly impressed. 

Variability: Body colouration varies from rufous to brown black. Head globo- 
se to slightly widely rounded posteriorly in dorsal view, its dorsal surface may be smo- 
oth to somewhat uneven, shallowly longitudinally wrinkled anteriorly. Humeri clearly 
protruding and metathoracic wings fully developed in the six specimens from 
Meghalaya (Shillong), Uttaranchal (16 km of Srinagar) and Rajasthan (Keoladeo 
Ghana Nat. Park). 

Distinctly variable in male/female abdominal characters. Male tergum VII often 
with more or less distinct median subapical protrusion. Process of female tergum VII 
of different size and form; inconspicuous, forming rather rounded edge in the 



STUDIES OF THE GENUS ANTHELEPH1LA HOPE 635 

specimens from Japan, Taiwan and China; more or less protruding in the specimens 
from Nepal (differences even within the same locality sample); female tergum VII 
strongly tapering, evenly shaped without any median, subapical process/edge in the 
specimens from Rajasthan (Keoladeo Ghana Nat. Park). Morphology of male sternum 
VII and sternite VIII varies in details extremely as follows; the specimens from Japan 
and Taiwan: sternum VII deeply emarginate/notched postero-medially, lateral sides of 
emargination moderately lobed (Fig. 51), paired prongs of sternite VIII less dilated in 
basal half, their apical portion simple, moderately enlarged and then narrowing towards 
pointed apex (Figs 52, 53); the specimens from Nepal: postero-median emargi- 
nation/notch of sternum VII less conspicuous and its postero-lateral sides rather evenly 
rounded, paired prongs of sternite VIII distinctly dilated in basal half, dorsal margin of 
their apical portion with minute pointed angle to distinctly projecting process; the 
specimens from Rajasthan (Keoladeo Ghana Nat. Park): sternum VII evenly emar- 
ginate and rounded posteriorly, paired prongs of sternite VIII less dilated in basal half, 
their apical portion simple, nearly evenly narrowing towards pointed apex. 

Differential diagnosis: Anthelephila imperatrix is very conspicuous in having 
the deeply longitudinally impressed and posteriorly clearly constricted pronotum, 
large, globose and rather coarsely punctured head, and whitish setose bands on the 
elytra. It shares this combination of characters only with A. aratrix sp. n., and differs 
by the characters given in the description of the latter species and in the key. 

Distribution: Recorded from Indonesia (Uhmann, 1988), Philippines (Pic, 
1903, 1916a), India (Krekich-Strassoldo, 1931; Uhmann, 1983, 1987; Telnov, 2003), 
Bangladesh (Bonadona, 1978), Laos, Vietnam, Sri Lanka, Nepal and Pakistan 
(Uhmann, 1983, 1987, 1988, 1989; Telnov, 2003), Japan: Honshu, Shikoku, Kyushu, 
Tsushima and Ryukyu Islands (Nomura, 1962; Werner, 1965; Sakai, 1989; Lafer, 1996; 
mostly as F cribriceps), China: Zhejiang, Fukien and Guangxi Prov. (Uhmann, 1988, 
as F cribriceps), Taiwan (Krekich-Strassoldo, 1913; Miwa, 1931; Uhmann, 1983; Hua, 
2002; mostly as F. cribriceps) and from the USA and Japanese territories in the Pacific 
Ocean: Bonin, Volcano, Mariana, Caroline and Palau Islands (Blair, 1942; Werner, 
1965). 

Comments: Nietner (1856) described Anthicus formicarius from Sri Lanka and 
later (Nietner, 1857) proposed replacement name, A. quisquiliarius, because of 
homonymy with A. formicarius LaFerté-Sénectère, 1849. Gemminger (1870) 
overlooked this fact and proposed for the same reasons another replacement name, 
A. myrmecodes. Both former names were synonymized with Anthelephilus imperator 
by Krekich-Strassoldo (1913). 

Having studied the type specimens of Formicomus imperator var. ruficolor and 
F. punctaticeps, I failed to find any significant differences from A. imperatrix and I 
believe them to be identical. The former taxon was first placed in synonymy with A. 
imperatrix, without any comments, by Telnov (2003). All the examined specimens of 
Anthelephilus (or Anthelephila) punctaticeps, listed for India, Nepal and Bhutan by 
Uhmann (1986, 1987, 1990a) and Telnov (2003), were found to be misidentified; they 
form a part of the type series of Anthelephila kresli sp. n., A. lobulicula sp. n., and A. 
sculpta sp. n. 



636 z - KEJVAL 

Formicomus cribriceps was described by Marseul (1876) based on specimens 
from Japan ("Nagasaki et Hiogo") collected by M. G. Lewis. It was synonymized with 
Anthelephilus imperator by Krekich-Strassoldo (1913), who found the specimens from 
Indonesia, Sri Lanka, Taiwan and Japan identical, showing only differences in 
colouration. This synonymy was followed by Winkler (1927) and Telnov (2003), and 
ignored by Miwa (1931), Nomura (1962), Sakai (1989), Lafer (1996), and Hua (2002). 
In his check-list of Japanese insects, Sakai (1989) listed both Anthelephila imperatrix 
and A. cribriceps, based on the differences between specimens from "Japan proper" 
and the Japanese island territories in the Pacific Ocean, treated as Anthelephilus 
imperator by Werner (1965) (M. Sakai, pers. coram.). In my opinion, the examined 
specimens of A. imperatrix from Japan, Taiwan, China (= F cribriceps), and especially 
those from Rajasthan (Keoladeo Ghana Nat. Park) show detailed but distinct 
differences in male/female abdominal characters from the typical form. Despite this, I 
have refrained from treating them as separate taxa (most likely as geographical 
subspecies), mainly because of the variation of these characters observed in the 
specimens from Nepal, and lack of material from China and the riverbasins of the 
Ganges and Brahmaputra in India. 

Based on label data of the examined specimens, A. imperatrix has been 
collected by sweeping grasses, sifting garbage-heap, forest leaf litter, in decaying 
coconut logs and under washed up plant material. In India, I found it locally quite 
common on/near various rooting, vegetable matter, especially near riverbanks. 
Considering its aptery, comparatively less conspicuous variation of the male characters 
and distribution of the related species, A. imperatrix is very probably a native of the 
Asian mainland, and most of its island occurences, especially in the case of Micronesia 
and Hawaii, are to be regarded as rather recent introductions. 

Anthelephila irula sp. n. Figs 22-25 

Type material: Holotype S, S-INDIA, Tamil Nadu state, Nilgiri hills, 15 km SE of 
Kotagiri, Kunjappanai env., 76°56'E 11°22'N, ca 900m, 22-30.V.1999, Z. Kejval & M. Tryzna 
Igt. (NMPC). Paratypes: 7 S â, 8 9 9 , same data as holotype (ZKDC, 1 specim. each in BMNH, 
MHNG, MNHN, NHMB, SMNS, DCDC, GUPC). -IS, Tamil Nadu, 15 km SE of Kotagiri, 
Kunchappanai env., 76°56'E 11°22'N \ Nilgiri hills, alt. 900 m, 17.-28.xi.1993, D. Boukal & Z. 
Kejval Igt. (ZKDC). 

Etymology: Named after Irulas, one of the original tribes inhabiting Nilgiri 

hills. 

Description: Body length 2.6-3.6 mm (holotype 3.4 mm). 

Male (holotype): Head and pronotum rufous brown; elytra largely brown black, 
with rufous brown base, suture, lateral margins and with very vaque indication of paler 
transverse spot in basal third; legs rufous brown, tarsi slightly paler, antennae and palpi 
rufous, apical 2-3 antennomeres slightly paler. 

Head: 1.2 times as long as wide, evenly rounded posteriorly in dorsal view; 
posterior temporal angles obsolete. Eyes small to medium-sized, moderately convex. 
Surface smooth, distinctly but rather sparsely punctured, very glossy; dorsal punctures 
unevenly spaced, separated mostly by about twice their diameter or sparser. Setation 
comparatively long and raised, subdecumbent to decumbent, evenly long, with 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 637 

sparsely scattered erect setae. Antennae rather short, at most moderately exceeding 
base of pronotum, distinctly enlarged in apical third; antennomere X slightly, 1.1 times, 
antennomere XI 1.6 times as long as wide. 

Thorax: Pronotum as long as wide, as wide as head including eyes, rather 
widely rounded anteriorly, strongly narrowing posteriad and shallowly impressed 
postero-laterally before base in dorsal view (not constricted); entire dorsal outline 
convex in lateral view; pronotai disc evenly shaped, without impressions. Surface 
conspicuously sculptured dorsally, smooth and glossy only antero-laterally (near 
procoxal cavities) and dorso-medially before base; dorsal side largely rather coarsely 
and densely corrugated; postero-lateral impressions coarsely, longitudinally wrinkled, 
with rather dorso-laterally situated wrinkles passing into dorsal corrugation; dorsal 
punctation similar to that on head, but mostly obscured by corrugation. Setation as on 
head, less raised setae rather subdecumbent; about four erect setae on each side antero- 
laterally more conspicuous and they appear to be articulated on minute protuberances, 
slightly protruding from lateral outline in dorsal view. Both mesosternum and meta- 
sternum simple. 

Elytra: 1.7 times as long as wide, strongly convex, narrowed and conjountly 
rounded apically in dorsal view; humeri entirely obsolete; postscutellar impression 
absent. Surface smooth, sparsely punctured, very glossy; punctation as coarse as on 
head but much sparser, interspaces seem to be extremely finely and sparsely punctured. 
Setation rather evenly long, distinctly longer than on head, mostly suberect, with 
sparsely scattered, erect setae. Metathoracic wings almost entirely reduced. 

Legs: Penultimate tarsomeres rather narrow, with terminal tarsomere articulated 
before midlength; all legs simple. Setation normally developed, moderately longer and 
denser on inner side of metatibiae. 

Abdomen: Sternum VII (Fig. 22) deeply emarginate posteriorly, with short and 
wide median process, bearing dense, stiff setae apically. Tergum VII simply rounded 
posteriorly, its apical margin slightly emarginate medially. Sternite VIII (Fig. 23); 
paired prongs simple, dorso-ventrally flattened, rather straightly projecting, rounded 
apically; each prong, excepting fine setation of median margin, with numerous long 
and thick setae in apical third (about 17 ventrally and 5 dorsally). Tergite VIII (Fig. 24) 
somewhat truncate posteriorly, its posterior margin moderately sinuous and with 
several long setae laterally. 

Aedeagus (Fig. 25): Apical portion of tegmen 0.6 times as long as basal-piece, 
nearly parallel-sided, trilobed apically, middle lobe wider, evenly narrowing towards 
rounded apex, slightly shorter than evenly narrow lateral lobes. 

Female: Externally identical with male, except as follows: sternum VII simple, 
rather evenly rounded posteriorly; tergum VII evenly rounded posteriorly. 

Variability: Slightly variable in colouration and characters of corrugation; 
paler spots in basal third of elytra mostly quite indistinct; dorsal surface of head in 
several specimens slightly corrugated mesally and near median margins of eyes; dorsal 
corrugation of pronotum somewhat variable in its extent, always prominent mesally. 

Differential diagnosis: Anthelephila irula sp. n. differs from the closely 
related A. besuchen by largely the smooth and more distinctly punctured surface of the 



638 z - KEJVAL 

head, moderately convex lateral subapical margins and rounded apices of the elytra, 
male abdominal sternum VII deeply emarginate posteriorly and with a distinct median 
process (cf. Figs 16, 22), prongs of male sternite VIII longer and narrower, rather 
straightly projecting (cf. Figs 17, 23), and male tergite VIII somewhat truncate 
posteriorly, with posterior margin moderately sinuous and longer setose. 

Distribution: India (Tamil Nadu). 

Comments: The specimens were collected near a stream in 1999, in plant debris 
on a sandy bank and in gaps between large stones. 

Anthelephila kresli sp. n. Figs 54-57, 62 

Type material: Holotype 8, NEPAL, 27. v. 1999 Kathmandu, Swayambunath stupa, 
P. Kresl Igt. (NMPC). - Paratypes: 1 o\ same data as holotype (ZKDC). - 1 8, W-NEPAL, 
Dhawalagiri Myagdi Distr., Kali-Gandaki Khola. 1100-1400m, Tatopani Probst, 27/28.6.1986 
[yellow frame] \ Anthelephilus punctaticeps (Pic) det.G.Uhmann 1988 (ZKDC). - 1 8, W- 
NEPAL, Buri Gandaki Macha Khola-Kholabenesi 1650m, 29.5.-4.6. leg. Probst 1990 [yellow 
frame] \ Anthelephilus punctaticeps (Pic) det.G.Uhmann 1990 (ZKDC). - 1 9, E-NEPAL, 
Dhankuta Arun Valley, Lamobagar Gola 27.5.-3.6.1980, 1000-14000 m leg. C. Holzschuh 
(ZKDC). - 1 8, NEPAL-Expeditionen Jochen Martens \ 246 Gorkha Dist., Darondi Khola un- 
terhalb Barpak bis Doreni 1100-900m Waldreste 12Aug83 Martens & Schawaller \ 
Senckenberg-Museum Frankfurt/Main \ Anthelephilus punctaticeps (Pic) det.G.Uhmann 1984 
(SMF). - 3 8 8, 4 9 9, Nepal420 Kathmandu Distr. Kathmandu-Baneshwar 1350 m, 
18.IV. 1995, Martens & Schawaller \ Anthelephilus punctaticeps (Pic) det.G.Uhmann 1996 
(SMNS, 1 specim. in GUPC). - 1 6,2 9 9, 583 NEPAL: Kathmandu Baneshwar 1300 m, 21.- 
25:JV.1998 leg. W. Schawaller \ Anthelephilus punctaticeps (Pic) det.G.Uhmann 2000 (SMNS). 
-18, Nepal444 Mustang Distr. Kali Gandaki, Dana 1500-1300 m, 14.V.1995 Martens & 
Schawaller \ Anthelephilus punctaticeps (Pic) det.G.Uhmann 1996 (SMNS). - 1 9, Nepal447 
Myagdi Distr. Mahabhir to Beg Khola 1100-1050 m, 15.V.1995 Martens & Schawaller \ 
Anthelephilus punctaticeps (Pic) det.G.Uhmann 1996 (SMNS). - 1 9, Nepal449 Myagdi Distr. 
Beg Khola village to Bega 1050-1650 m, 16.V.1995 Martens & Schawaller \ Anthelephilus 
punctaticeps (Pic) det.G.Uhmann 1996 (SMNS). - 1 9, 210 Dhading Dist., unter Samari 
Banjyang, 1000 - 1300 m, 23 Juli 83, kulturland Martens & Schawaller leg. \ Anthelephilus 
punctaticeps (Pic) det.G.Uhmann 1984 (GUPC). - 2 8 8, 1 9, NEPAL, Prov. Bagmati 
Kathmandu, Bagmati nr. Gorkhana Park 27°43'N; 85°20'E 1340 m NN, 17.VI.1999 leg. A. 
Weigel \ Sammlung Naturkundemuseum Erfurt \ Anthelephila punctaticeps (Pic) det. D. Telnov, 
2000 (NKME, 1 specim. in ZKDC). -Id, NEPAL Kathmandu Swayambhunath Templehiigel 
24VI.1997 leg. A. Weigel \ Anthelephilus punctaticeps (Pic) det. G. Uhmann 1998 (ZKDC). - 
1 8, NEPAL, Kathmandu, N Bagmati River, 1300 m NN, 06VI.1995 leg. M. Hartmann \ 
Anthelephilus punctaticeps (Pic) det. G. Uhmann 1996 (NKME). -18, NEPAL Kathmandu, N 
Safaripark, Mauer 06VI.1995 1300 m Ü.NN leg. A. Weigel \ Anthelephilus punctaticeps (Pic) 
det. G. Uhmann 1996 (NKME). -19, NEPAL oc. 1300 m Kathmandu NW, Balaju Vishnumati 
River, 17VI.1999 leg. A. Weigel \ Sammlung Naturkundemuseum Erfurt \ Anthelephila punctat- 
iceps (Pic) det. D. Telnov, 2000 (NKME). -18, Indien Darjeeling D. Bhakta B. \ Pedong 23- 
28.III.87 \ Anthelephila punctaticeps Pic det. D. Telnov, 1999 (NHMB). 

Etymology: Dedicated to Petr Kresl (Spule, Czech Republic), collector of the 
holotype. 

Description: Body length 3.7-4.6 mm (holotype 4.0 mm). 

Male (holotype): Head brown black; pronotum largely dark rufous brown, 
distinctly darkened, nearly brown black with slight rufous tinge dorsally; elytra dark 
brown to brown black, with rufous brown base, lateral margins, and with indication of 
two pairs of vague, narrow, paler bands (their location identical with that of the setose 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 



639 




Figs 54-61 



Anthelephila kresli sp. n., male: (54) sternum VII. (55) sterilite VIII (half). (56) prong of sternite 
VIII, lateral view. (57) tegmen. A. lobulicula sp. n., male: (58) sternum VII. (59) sternite VIII. 
(60) prong of sternite VIII, lateral view. (61) tegmen. Scale (0.2 mm): A - Figs 55, 59, 61, 
B - Figs 56, 57, 60; (0.5 mm): C - Figs 54, 58. 



640 Z. KEJVAL 

bands, see below); antennae rufous brown basally, becoming darker, brown black in 
apical third, palpi brown black; legs largely brown black, basal narrowed portion of 
femora rather contrastingly yellowish to pale rufous, tarsi rufous brown. 

Head: 1 .2 times as long as wide, somewhat unevenly rounded posteriorly, with 
base slightly produced medially in dorsal view; posterior temporal angles entirely 
obsolete. Surface smooth, conspicuously punctured, glossy; dorsal punctures rather 
coarse, unevenly spaced, separated mostly by about their diameter, at places denser, 
sparser posteriorly. Setation fine, mostly subdecumbent, with sparsely scattered, some- 
what longer, erect setae. Eyes small, rather convex. Antennae moderately exceeding 
base of pronotum, weakly enlarged in apical third; antennomere X 1.4 times, 
antennomere XI 2.2 times as long as wide. 

Thorax: Pronotum 1.3 times as long as wide, moderately narrower than head 
including eyes, evenly rounded anteriorly, distinctly narrowing posteriad and rather 
shallowly impressed postero-laterally in dorsal view; entire dorsal outline convex in 
lateral view; pronotai disc with very slight indication of shallow, median longitudinal 
impression in anterior third. Surface smooth, conspicuously punctured, glossy, postero- 
lateral impression at most very shortly, inconspicuously wrinkled; punctation as on 
head, rather evenly coarse, covering the whole surface (including lateral sides), some- 
what sparser laterally, near procoxal cavities, and postero-dorsally. Setation as on head, 
erect longer setae more numerous. Mesosternum simple. Metasternum with a pair of 
distinct, apically setose protuberances postero-medially, near median margin of 
metacoxae. 

Elytra: 1.7 times as long as wide, strongly convex, subtruncate apically, with 
elytral apices separately rounded; humeri entirely obsolete; postscutellar impression 
absent. Surface smooth, distinctly punctured, glossy; punctation uneven, ordinary 
punctures generally much finer and mostly sparser than on head, especially in apical 
third; additionally with densely spaced, nearly contiguous punctures, forming two 
paired bands, touching neither lateral margins nor suture; anterior bands situated in 
basal third, narrow to interrupted, more oblique, directing postero-mediad from lateral 
sides, posterior bands situated shortly behind mid-length, more conspicuous, wider, 
narrowing towards suture, transverse to slightly oblique antero-mediad from lateral 
sides. Setation mostly as short as on head, subdecumbent, with scattered, long, erect 
setae (somewhat longer than erect setae of head); setation mostly pale, setae of dense 
punctures contrastingly whitish, thicker, nearly appressed, forming distinct setose 
bands, some whitish, more raised setae scattered near base. Metathoracic wings almost 
entirely reduced. 

Legs: Rather robust; penultimate tarsomeres narrow, terminal tarsomere rather 
subapical in metatarsi; metatibiae somewhat uneven on inner side just beyond mid- 
length. Setation normally developed. 

Abdomen: Sternum VII (Fig. 54) with posterior margin very deeply emarginate 
and with conspicuous median process, about as long as lateral lobes of emargination; 
median process wide, laterally flattened, with rounded dorsal and sharper ventral 
longitudinal edge, its apex curved ventrad and bluntly pointed. Tergum VII subtruncate 
posteriorly, with rounded postero-lateral angles, and shallowly impressed subapically. 
Sternite VIII (Figs 55, 56); paired prongs robust, strongly widened in distal half in 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 641 

lateral view, with conspicuous, long and wide, flattened process ventro-medially at 
about mid-length, surface of prongs uneven, variously buckled, their lateral sides with 
several gibbosities in dorsal view. Tergite VIII (Fig. 62) nearly parallel-sided in dorsal 
view, distinctly, widely emarginate posteriorly, its postero-ventral side somewhat 
produced and exceeding median part of emargination in dorsal view; rounded lateral 
lobes of emargination long and rather densely setose. 

Aedeagus (Fig. 57): Apical portion of tegmen 0.8 times as long as basal-piece, 
parallel-sided, trilobed apically, middle lobe wide basally, abruptly narrowed, 
elongated and rather sharply pointed apically, much exceeding narrower and bluntly 
pointed lateral lobes. 

Female: Externally identical with male, except as follows: metasternum simple, 
lacking paired protrusions; metatibiae narrower and rather straight; sternum VU 
simple, evenly rounded posteriorly; tergum VII evenly rounded posteriorly, shallowly 
impressed and with short, median longitudinal edge subapically. 

Variability: Rather variable in colouration; some specimens darker coloured 
with both pale bands of elytra indistinct, in contrast other specimens paler coloured as 
follows: both head and pronotum dark rufous brown, elytra with basal third (as far as 
anterior band), both lateral margins and suture, and posterior transverse bands rufous 
brown, legs brown, nearly basal half of antennae rufous brown. The paratype from 
Dhankuta district (ZKDC) with rounded, but protruding elytral humeri and well 
developed metathoracic wings. 

Differential diagnosis: Anthelephila kresli sp. n. is related to A. lobulicula 
and A. sculpta spp. n., as suggested by the similar form of male sternite VIII (paired 
prongs with large process ventro-medially). It differs from A. lobulicula sp. n. by its 
robust appearance, wider and coarsely punctured pronotum, simple (not trilobed) apex 
of the median process of male sternum VII, and by numerous details in morphology of 
the male sternite and tergite VIII (cf. Figs 55, 56 and 59, 60). See the differential 
diagnosis of the latter species and the key for its separation from A. sculpta sp. n. 

Distribution: Nepal, India (West Bengal). 

Anthelephila lobulicula sp. n. Figs 58-61, 63 

Type material: Holotype S, NEPAL-Expeditionen Jochen Martens \ 344 Taplejung 
Distr., confluence of Kabeli Khola and Tada Khola, 1000-1000 m, mixed broad-leaved forest, 
23.-25.Apr 88 Martens & Schawaller \ Anthelephilus punctaticeps (Pic) det.G.Uhmannl989 

(SMNS). 

Etymology: Composed from Latin lobulus (small lobe) and cuius (posterior, 
abdomen); named in reference to the form of male abdominal sternum VII. 

Description: Body length 3.6 mm. 

Male (holotype): Head and pronotum black; elytra black, with brownish base 
and brownish tinge in apical third; antennae and palpi brown black, basal anten- 
nomeres rufous brown; legs largely brown black to black, basal narrowed portion of 
meso- and metafemora pale yellowish, tarsi rufous brown. 

Head: 1 . 1 times as long as wide, somewhat unevenly rounded posteriorly, with 
base slightly produced medially; posterior temporal angles entirely obsolete. Surface 



642 z - KEJVAL 

smooth, glossy, distinctly punctured, with some shallow wrinkles anteriorly on frons; 
dorsal punctation uneven, punctures mostly coarse, separated by about their diameter, 
finer and more widely spaced near base. Setation mostly rather short, subdecumbent, 
with sparsely scattered, slightly longer, erect setae. Eyes small, moderately convex. 
Antennae moderately exceeding base of pronotum, only slightly enlarged in apical 
third; antennomere X 1.3 times, antennomere XI 2.1 times as long as wide. 

Thorax: Pronotum 1.7 times as long as wide, distinctly narrower than head 
including eyes, evenly rounded anteriorly, distinctly narrowing posteriad and rather 
sharply impressed postero-laterally in dorsal view; entire dorsal outline convex in 
lateral view, strongly convex before midlength; pronotai disc with weak median longi- 
tudinal impression at about midlength. Surface smooth, glossy, somewhat less 
distinctly punctured, postero-lateral impressions finely wrinkled; dorsal punctation 
much finer and sparser than on head. Setation as on head. Mesosternum simple. 
Metasternum with a pair of distinct, apically setose protrusions postero-medially, near 
median margin of metacoxae. 

Elytra: 1.7 times as long as wide, convex, truncate apically; humeri entirely 
obsolete; postscutellar impression absent. Surface smooth, glossy, less distinctly punc- 
tured; punctures generally nearly as fine as on pronotum, unevenly spaced, mostly 
sparser than on head and pronotum; additionally with very densely spaced, nearly 
contiguous punctures, forming paired bands/patches, touching neither lateral margins 
nor suture; anterior band composed more likely of two small, narrowly connected 
patches of dense punctures, situated in basal third, directed obliquely postero-mediad 
from lateral sides, posterior band situated shortly behind mid-length, more conspi- 
cuous, with small patch of dense punctures, narrowly separated medially. Setation 
rather evenly long, moderately longer than on head, mostly decumbent to suberect, 
with sparsely scattered, erect setae; setation mostly fine and pale, setae of dense 
punctures contrastingly whitish, thicker, subdecumbent to appressed, forming distinct 
setose bands. Metathoracic wings almost entirely reduced. 

Legs: Penultimate tarsomeres rather narrow, terminal tarsomere rather subapical 
in metatarsi; all legs simple. Setation normally developed. 

Abdomen: Sternum VII (Fig. 58) with posterior margin very deeply emarginate 
and with conspicuous median process, about as long as lateral lobes of emargination; 
median process wide, distinctly trilobed apically, its ventral side over the whole length 
with keel-like median edge. Tergum VII rounded posteriorly, its apical margin slightly 
emarginate medially. Sternite VIII (Figs 59, 60); paired prongs robust, moderately ar- 
cuate in dorsal view, with conspicuous, flattened, fin-like process ventro-medially at 
about mid-length, apical portion of prongs simple, rounded apically, rather evenly se- 
tose. Tergite VIII (Fig. 63) nearly parallel-sided, subtruncate and moderately emargi- 
nate posteriorly, with postero-ventral margin somewhat produced. 

Aedeagus (Fig. 61): Apical portion of tegmen 0.6 times as long as basal piece, 
nearly parallel-sided, trilobed apically, middle lobe conspicuously wide basally, strong- 
ly narrowed, elongated and sharply pointed apically, moderately exceeding narrow, 
apically rounded and moderately divergent lateral lobes. 

Female: Unknown. 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 643 

Differential diagnosis: Anthelephila lobulicula sp. n. shares the similar 
general form of male sterilite VIII with A. kresli and A. sculpta spp. n., but differs by 
the slender appearance, less distinct punctation of the pronotum (dorsal punctures 
much finer than those on head), and in the male characters. 

Distribution: Nepal. 

Anthelephila ovipennis (Bonadona, 1984) Figs 30-33, 35-37 

Formicomus (Anthelephilus) ovipennis Bonadona, 1984: 486, figs 23, 28-30. 

Type material (not examined): Holotype, 6 , CONGO: Belin \ Anthelephilus ovipennis 
(coll. Pic, MNHN). - Paratypes: 4 specim., Kivu, territoire Uvira, Mulenge, 2010 m, vestige de 
forêt ombrophile, récolté dans lhumus, V.1951 (N. Leleup leg.) \ A. ovipennis Pic (coll. Pic, 
MNHN). - 2 specim., Kivu: Tshibinda, XI. 1932 (L. Burgeon leg.) \ Formicomus près 
subfasciatus \ Anthelephilus ovipennis (coll. Pic, MNHN). 

Other material examined: 1 6, ZAIRE, Kivu, Irangi, 800 m, 22.ii.1985, H. Mühle leg. 
(ZKDC). -4 SS, 1 9, RUANDA, Pref. Cyangugu, Nyakabuye env., 17.ii.1985, H. Mühle leg. 
(GUPC, ZKDC). -3 SS, 5 9 9, same locality, but different dates: 12.-19.vii.1984, 28.xi.1984, 
22.iii.1985, 3.X.1985 or 19.xii.1985 (GUPC, ZKDC). 

Redescription: Body length 3.2-4.1 mm. 

Male (Ruanda, ZKDC): Mesosternum simple; metasternum with a pair of 
robust, apically setose protrusions postero-medially, near median margin of metacoxae. 
Metatibiae rather stout distally, with small patch/tuft of short, thicker, yellowish rufous, 
laterad pointing setae and numerous long, finer, pale setae on inner side subapically 
(Fig. 36); basal protarsomere enlarged, rather long and wide. Sternum VII (Fig. 30) 
moderately emarginate posteriorly and with small, bluntly pointed median process, 
bearing several longer setae apically; dorsal side of median process with short 
longitudinal edge. Tergum VII evenly rounded posteriorly. Sternite VIII (Figs 31, 32); 
prongs wide, strongly narrowed in apical third in dorsal view, their apex somewhat 
obliquely truncate and pointed; ventral side of prongs with longitudinal, distally 
abruptly shortened slat/lobe, its outer edge finely denticulated and terminating distally 
in robust, pointed process; setation less conspicuous, rather fine and sparse, with 
scattered longer setae laterally and apically. Tergite VIII simple, arcuate, evenly 
rounded posteriorly. Aedeagus (Fig. 33); apical portion of tegmen 0.6 times as long as 
basal-piece, parallel-sided, trilobed apically; middle lobe wide basally, strongly 
narrowed at about midlength, elongated and pointed apically, exceeding narrow, 
apically rounded, lateral lobes. 

Female (Ruanda, ZKDC): Externally identical with male, except as follows: 
metasternum simple, lacking paired protrusions; basal protarsomere smaller, narrow; 
metafemora narrower distally, uniformly simply setose; sternum VII simple, somewhat 
unevenly rounded posteriorly, and with a tuft of long, stiff setae apically; tergum VII 
subtriangular, rounded and with longer, dense stiff setae apically (Fig. 37). 

Differential diagnosis: Anthelephila ovipennis is closely related to 
A. congoana, as suggested especially by the very similar form of the prongs of male 
sternite VIII, but differs by the head being rather evenly rounded posteriorly, the male 
metatibiae subapically with a small patch of short, dense, yellowish rufous setae on 



644 z - KEJVAL 

inner side, male sternum VET shallowly emarginate posteriorly and with a small, bluntly 
pointed median process, male tergite VIII evenly rounded apically, female tergum VII 
rather rounded apically, as well as by some details in morphology of male sternite VIH 
and the tegmen. 

Distribution: Zaire, Ruanda, Uganda. 

Comments. As stated by Bonadona (1984), he found the types of Formicomus 
ovipennis in the collection of Maurice Pic (MNHN) and described this species using a 
Pic's manuscript name. 

Anthelephila sculpta sp. n. Figs 4, 64-68 

Type material: Holotype S, INDIA Meghalaya Khasi Hills 5. XI. Nongpoh 700m 
Besuchet-Löbl 78 [1978; p] \ Anthelephilus punctaticeps (Pic) det.G.Uhmann 1986 (MHNG). - 
Paratypes: 5 SS, 5 99, same data as holotype (6 specim. in MHNG, 4 specim. in ZKDC). - 
1 S,\ 9, INDIA Assam Manas 200m 23.X.78 Besuchet-Löbl [p] \ Anthelephilus punctaticeps 
(Pic) det.G.Uhmann 1986 (MHNG, GUPC). -IS, same data, except: 22.X.78 (GUPC). -IS, 
INDIA W. Bengal Darjeeling dist. Teesta 250m 10.X.1978 Besuchet Lobi \ Anthelephilus 
punctaticeps (Pic) det.G.Uhmann 1986 (MHNG). -IS, BHUTAN 1981 Bhakta B. \ 
Phuntsholing 2/400 m 3. IX. \ Anthelephila punctaticeps Pic det. D. Telnov, 1999 (NHMB). 

Etymology: From Latin sculptilis (carved, sculptured); named in reference to 
the coarse sculpture of the head and pronotum. 

Description: Body length 3.9-4.3 mm. 

Male (holotype): Head and pronotum rufous brown; elytra dark rufous brown, 
with vague indication of two narrow, paler bands (their location identical with that of 
setose bands, see below); antennae and palpi rufous; legs brown with rufous tinge, 
basal narrowed portion of femora pale rufous, tarsi rufous brown. 

Head: 1.2 times as long as wide, somewhat unevenly rounded posteriorly, with 
base slightly produced medially in dorsal view; posterior temporal angles entirely 
obsolete. Surface less glossy, very coarsely and rather evenly sculptured, including 
ventral side; punctures situated in deep, pit-like impressions, dorsal impressions 
narrowly separated, at places nearly contiguous. Setation moderately long, sub- 
decumbent to decumbent, with few distinctly longer, erect setae. Eyes small, rather 
convex. Antennae rather short, slightly exceeding base of pronotum, moderately 
enlarged in apical third; antennomere X 1.2 times, antennomere XI 1.9 times as long 
as wide. 

Thorax: Pronotum 1.3 times as long as wide, only slightly narrower than head 
including eyes, regularly rounded anteriorly, distinctly narrowed and shallowly 
impressed postero-laterally in dorsal view; entire dorsal outline convex in lateral view; 
pronotai disc with very slight indication of median longitudinal impression close 
before midlength (obscured by coarse sculpture). Surface characters as on head, pit- 
like impressions rather evenly covering the whole surface, including unwrinkled 
postero-lateral impressions. Setation as on head, long erect setae somewhat more 
numerous. Mesosternum simple. Metasternum with a pair of distinctly protruding, 
apically setose protrusions postero-medially, near median margin of metacoxae. 

Elytra: 1.7 times as long as wide, strongly convex, narrowed, subtruncate to 
nearly conjointly rounded apically in dorsal view; humeri entirely obsolete; post- 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 



645 




A«- 
D- 



Figs 62-70 
Male tergite VIII: (62) Anthelephila kresli sp. n. (63) A. lobulicula sp. n. A. sculpta sp. n., male: 
(64) sternum VII. (65) sterilite Vili. (66) prong of sternite VIII, lateral view. (67) tergite VIII. A. 
subtruncata (Pic), male: (69) metatibia. (70) sternum VII. Scale (0.2 mm): A - Fig. 65, B - Figs 
66, 68, C - Figs 62-64, 67; (0.5 mm): D - Fig. 70, E - Fig. 69. 



scutellar impression absent. Surface smooth, glossy, distinctly punctured; punctation 
uneven, ordinary punctures rather coarse, sparse in basal half, becoming finer and even 
sparser posteriad; additionally with densely spaced, nearly contiguous punctures, 



646 z - KEJVAL 

forming paired, transverse to moderately oblique bands/patches; anterior band 
composed more likely of two separated patches of dense punctures, situated in basal 
third, posterior band situated shortly behind mid-length, more conspicuous, with small, 
narrowly separated patch of dense punctures antero-medially. Setation mostly 
distinctly longer than on head, decumbent to suberect, with sparsely scattered, erect 
setae; ordinary setae fine and pale, setae of dense punctures contrastingly whitish, 
thicker and blunt, nearly appressed, forming distinct setose bands (Fig. 4), some 
whitish setae scattered also along suture in basal and apical third and between setose 
bands laterally. Metathoracic wings almost entirely reduced. 

Legs: Robust; penultimate tarsomere rather narrow, terminal tarsomere sub- 
apical in metatarsi; metatibiae somewhat swollen, with longitudinal, strongly 
protruding ridge on inner side in distal half. Setation rather more distinct, metatibiae 
conspicuously long setose. 

Abdomen: Sternum VII (Fig. 64) with posterior margin distinctly emarginate 
and with small tuft of longer setae medially. Tergum VII evenly rounded posteriorly. 
Sternite VIII (Figs 65, 66); paired prongs rather robust, laterally flattened in distal half, 
with conspicuous, long and wide, flattened process ventrally close behind midlength; 
surface of prongs largely bare, densely setose and with numerous long setae in apical 
portion, median side of ventral process densely short setose. Tergite VIII (Fig. 67) 
simple, its posterior margin somewhat unevenly rounded, with slight lateral angle, with 
some longer setae scattered postero-medially. 

Aedeagus (Fig. 68): Apical portion of tegmen 0.7 times as long as basal-piece, 
moderately narrowed and trilobed apically, middle lobe wide basally, nearly evenly 
narrowing towards pointed apex, much exceeding short lateral lobes. 

Female: Externally identical with male, except as follows: metasternum simple, 
lacking paired protrusions; metatibiae lacking longitudinal inner ridge; sternum VII 
simple, evenly rounded posteriorly; tergum VII flattened to shallowly impressed and 
with short, median longitudinal edge subapically. 

Differential diagnosis: Anthelephila sculpta sp. n. appears to be related to 
A. kresli sp. n. and A. lobulicula sp. n., as suggested by the similar general form of male 
sternite VIII (prongs with large process ventro-medially). It can be easily recognized 
by the coarse sculpture of the head and pronotum, male metatibia with longitudinal 
inner ridge, male sternum VII less deeply emarginate posteriorly and lacking median 
process, and by numerous details in other male characters. 

Distribution: India (Meghalaya, West Bengal, Assam), Bhutan. 

Anthelephila subtruncata (Pic, 1899) Figs 69-74 

Formicomus (Anthelephilus) subtruncatus Pic, 1899: 105. 

Type material: Syntypes: 1 9 , type [h; yellowish label] \ Type [p; red label] \ Museum 
Paris Coll. M.Pic [p; blue label] \ F. subtruncatus Pic [new label, not Pic's handwritting] 
(MNHN). - 1 o\ [h; illegible] \ Af aus' e [h; = Afrique australe, see Comments] \ Museum Paris 
Coll. M. Pic [p; blue label] (MNHN). 

Other material examined: 1 o\ 3 9 $, South Africa, N of Port Elizabeth, Dunbrody, 
8.i., 6.ii. or 10.ii.1903 (coll. Pic, MNHN; BMNH). - 1 9 [nearly completely damaged], South 
Africa, S of Grahamstown, Boknes, lO.i.1948, J. C. vanHille leg. (coll. Pic, MNHN). - 1 $, 



STUDIES OF THE GENUS ANTHELEPH1LA HOPE 647 

2 9 9, South Africa, Cape Prov., Somerset East, 23.-31.xii.1930, R. E. Turner leg. (BMNH, 1 
specim. in ZKDC). -29$, same data, except: x. 1930 (BMNH). - 1 $ , South Africa, East Cape 
Prov., Katberg, xii.1932, R. E. Turner leg. (BMNH). -2 6 6, South Africa, East Cape Prov., 
Kasouga, 30.x. 1977, J. C. vanHille leg. (DCDC). -19, South Africa, Eastern Cape, Alexandria- 
Woody Cape, 10.-13.xii.1997, I. Jenis leg. (ZKDC). - 1 9, South Africa, Eastern Cape Prov., 
Woodi Cape Nat. Reserve near Alexandria, 28.xi.-l.xii.2000, S. Becvâr leg. (ADBC). -3 6 6, 
10 9 9 , South Africa, E Cape Prov., Grahamstown, 19.V.1946, W. E. Collett leg. (AMGS). -19, 
same locality, 12.iii.1953, H. D. Brown leg. (AMGS). - 1 6, 2 9 2, South Africa, E Cape Prov., 
Hogsback, xii.1959, J. C. van Hille leg. (AMGS, ZKDC). - 1 6, 2 9 9, same locality, 
30.ix.1947, E. Rogers leg. (AMGS). - 1 9, South Africa, E Cape Prov., Kasouga, i.1940 
(AMGS). -16, South Africa, E. Cape Prov., Dunbrody, x.1902, J. O'Neil leg. (AMGS). -2 6, 
South Africa, E Cape Prov., Keurboons river, iv.1944, Bester & Collett leg. (AMGS). 

Redescription: Body length 2.9-3.9 mm. 

Male (syntype): Dark brown to brown black, nearly unicoloured. 

Head. 1.2 times as long as wide, somewhat unevenly, slightly widely rounded 
posteriorly in dorsal view; temporal angles indistinct, entirely rounded. Surface with 
fine, dense microsculpture and thus less glossy, somewhat uneven in anterior half on 
frons, distinctly punctured; dorsal punctures unevenly spaced, separated mostly by 
about twice their diameter. Setation short, mostly subdecumbent to appressed, with few 
short, erect setae. Eyes small, at most moderately convex. Antennae distinctly 
exceeding base of pronotum, moderately enlarged in apical third; antennomere X 1 .2 
times and antennomere XI 1.9 times as long as wide. 

Thorax: Pronotum 1.2 times as long as wide, moderately narrower than head 
including eyes, nearly evenly rounded anteriorly, strongly narrowing posteriad and 
distinctly, rather sharply impressed postero-laterally (constricted) in dorsal view; entire 
dorsal outline of pronotum convex in lateral view; pronotai disc with indication of 
median longitudinal impression/furrow in anterior half and posteriorly before base. 
Surface with dense microsculpture and thus less glossy, similarly as in head, in addition 
uneven, longitudinally corrugated postero-dorsally; bottom of postero-lateral 
impressions microsculptured (without coarse wrinkles) and adjacent basal area 
distinctly punctured; dorsal punctation similar to that on head. Setation as on head. 
Mesosternum simple. Metasternum with a pair of conspicuously projecting, apically 
pointed protrusions postero-medially, near median margin of metacoxae. 

Elytra: 1.6 times as long as wide, strongly convex, narrowed and subtruncate 
apically; humeri entirely obsolete; postscutellar impression absent. Surface smooth, 
distinctly punctured, glossy; punctation mostly sparser and finer than on head, espe- 
cially in apical third, at places moderately denser (posterior setose band, see below). 
Setation similar to that on head, at most slightly longer, appressed, with scattered short 
erect setae, mostly pale, some appressed setae whitish and thicker, forming two vague, 
transverse bands, situated in basal third and shortly behind midlength of elytra, some 
whitish setae scattered also laterally and near base. Metathoracic wings strongly 
reduced. 

Legs: Penultimate tarsomeres rather narrow, terminal tarsomere subapical in 
metatarsi; mesofemora slightly dilated on inner side subapically (Fig. 71); mesotibiae 
slightly bent and rather stout; metatibiae enlarged and curved distally, with inner apical 
margin projecting into blunt process, with single apical spine (Fig. 69). Setation mostly 
normally developed; mesofemora and mesotibiae on inner side with numerous, 
conspicuously long, raised setae (Fig. 71). 



648 z - KEJVAL 

Abdomen: Sternum VII (Fig. 70) nearly simple, triangular and apically pointed 
posteriorly. Tergum VII evenly rounded posteriorly. Sternite VIII (Figs 72, 73); paired 
prongs simple, wide, strongly narrowed and moderately converging apically in dorai 
view, their apex curved ventrad and with robust dent-like process on ventral side 
subapically; setation of prongs rather short and scattered, inconspicuous. Tergite VIII 
simple, with posterior margin rounded and slightly emarginate medially. 

Aedeagus (Fig. 74): Apical portion of tegmen 0.6 times as long as basal-piece, 
trilobed apically, apical lobes nearly identical, wide basally and nearly evenly 
narrowing towards blunt apex. 

Female: Externally identical with male, except as follows: metasternum simple, 
lacking paired protrusions; middle legs simple, uniformly short setose; metatibiae 
narrow, simple, lacking subapical process, and with two apical spines; sternum VII 
with posterior margin evenly rounded apically. 

Variability: Some specimens from Somerset East with head more distinctly 
wider than pronotum, surface of pronotum more glossy, unwrinkled postero-dorsally, 
with fine corrugation indistinct, and with dorsal, median longitudinal impression of 
pronotum either rather distinctly or only slightly indicated anteriorly behind collar. 

Differential diagnosis: Of the related Afrotropical species, Anthelephila 
subtruncata resembles especially A. vanhillei sp. n. by its dark colouration and by the 
head somewhat widely rounded posteriorly. It can be distinguished from this species 
by form of the pronotum and by a number of clear differences in male characters (see 
the differential diagnosis of the latter species and key). 

Distribution: South Africa (Eastern Cape Prov.). 

Comments: Pic (1899) described Formicomus subtrunctatus based on 
specimens collected in southern Africa ("Afrique australe") and provided to him by M. 
H. Donckier. The record from Cape Province by Hille (1961) is based on the type 
specimens of A. vanhillei sp. n. The record from Uganda by Uhmann (1990b) is 
dubious and very probably is based on a misidentification. 

Anthelephila vanhillei sp. n. Figs 5, 75-79 

Type material: Holotype 3, S.Afr., S.W. Cape Arniston, dunes 34.39 S - 20.13 E \ 
29.8.1983; E- Y: 1996 coastal dunes, day leg. Endrödy, Penrith (TMP). - Paratypes: Il a 3, 
12 ? 9 , same data as holotype (TMP. 4 specim. in ZKDC). -12$, same data as holotype ex- 
cept: "Arniston, inland" and "E-Y:1994 groundtraps, 59 days" (TMP). - 6 3 3,9 9 9, S.Afr., 
S.W. Cape Struisbaai 34.46 S - 20.03 E \ 28.8.1983; E- Y: 1989 groundtraps, 60 days leg. 
Endrödy, Penrith \ groundtrap with banana [or meat or faeces] bait (TMP, 2 specim. in ZKDC). 
- 1 9, S.Afr., S.W.Cape Gansbaai, 10 km NE 34.31 S - 19.25 E \ 27.8.1983; E- Y: 1983 ground- 
traps, 63 days leg. Endrödy, Penrith \ groundtrap with faeces bait (TMP). - 1 9, S.Afr., 
S.W.Cape Stanford, 13 km S. 34.40 S - 19.26 E \ 27.8.1983; E- Y: 1982 under stones, care. leg. 
Endrödy, Penrith (TMP). - 2 3 3,3 9 9, SOUTH AFRICA Eastern Cape Alexandria - Woody 
Cape 10 - 13.12.1997 Ivo Jenis leg. (ZKDC). -.7 3 3, 6 9 9, S. Afr. Cape Prov. De Hoop Vlei. 
20 miles E Bredasdorp 2.1.51. No. 107 \ Swedish South Africa Expedition 1950-1951 Brinck- 
Rudebeck \ Formicomus subtruncatus Pic det. J C.vanHille Oct. 1960 [three 9 9 in addition with 
sex-mark label] (ZILS). - 1 3, S. Afr. Cape Prov. Mosselbaai 8-9.1.51 No. 127 \ Swedish South 
Africa Expedition 1950-1951 Brinck-Rudebeck \ Formicomus subtruncatus Pic det. J C.vanHille 
Oct. 1960 (ZILS). -13, Cape Province: Mossel Bay. Dec. 1934 [p] \ S.AFRICA: R.E. Turner. 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 



649 




A- 



Figs 71-79 
Anthelephila subtruncata (Pic), male: (71) mesofemur with tibia. (72) sternite VIII (half). (73) 
prong of sternite VIII (outline), lateral view. (74) tegmen. A. vanhillei sp. n., male: (75, 76) 
mesotibia, different views. (77) sternum VII. (78) prong of sternite VIII, lateral view. (79) 
tegmen. Scale (0.2 mm): A - Figs 77, 78, B - Figs 72, 73, C - Fig. 74, D - Fig. 79, E - Figs 71, 
75, 76. 



650 z - KEJVAL 

B.M. 1935-73. [p; with blue line] \ Formicomus subtmncatus Pic det. J.C. van Hille [h] 
(BMNH). -19, Cape Province. Mossel bay. August 1932. [p] \ S. Africa R.E. Turner. Brit. Mus. 
1932-421. [p; with blue line] \ on ... B. M [h; partly illegible] \ Formicomus subtmncatus Pic 
det. J.C. van Hille [h] (BMNH). - 3 3 3, 1 9, Sedgefield 14 Dec. 1977 J. C. van Hille \ 
Formicomus subtmncatus Pic det. J.C. v. Hille (GUPC, 1 3 ZKDC). - 3 3 3, 2 9 9, Boknes 
8-1-1947 J. C. van Hille \ Entomology Dept. Albany Museum Somerset Street Grahamstown 
6139 \ Formicomus subtmncatus Pic det. J. C. v. Hille (AMGS). - 1 3,3 9 9, same data, except: 
10-1-1948 (AMGS). 

Etymology: Named in honour of the late J.C. van Hille, well known specialist 
in the Afrotropical Anthicidae. 

Description: Body length 3.2-3.9 mm (holotype 3.8 mm). 

Male (holotype): Body brown black, nearly unicoloured; legs dark brown to 
brown black; antennae brown black, basal 2-3 antennomeres partly at most slightly 
paler; palpi brown black. 

Head: 1.2 times as long as wide, rather widely rounded posteriorly in dorsal 
view (Fig. 5); posterior temporal angles rounded. Surface with fine, dense micro- 
sculpture and thus less glossy, distinctly punctured; dorsal punctures separated mostly 
by about twice their diameter. Setation short, appressed to subdecumbent, with few 
short, inconspicuous erect setae. Eyes small, at most moderately convex. Antennae 
distinctly exceeding base of pronotum, moderately but distinctly enlarged distally; 
antennomere X 1.2 times and antennomere XI 1.8 times as long as wide. 

Thorax: Pronotum 1.3 times as long as wide, distinctly narrower than head 
including eyes, nearly evenly rounded anteriorly, only moderately narrowing posteriad 
and rather shallowly impressed postero-laterally in dorsal view (Fig. 5); entire dorsal 
outline of pronotum moderately convex in lateral view; pronotal disc with slight indi- 
cation of median longitudinal impression/furrow in anterior half. Surface with dense 
microsculpture and thus less glossy, similarly as in head; bottom of postero-lateral 
impressions microsculpured (without coarse wrinkles) and adjacent basal area 
distinctly punctured; dorsal punctation similar to that on head, finer and sparser 
posteriorly near base. Setation as on head. Mesosternum simple. Metasternum with a 
pair of conspicuously projecting, apically setose protrusions postero-medially, near 
median margin of metacoxae. 

Elytra: 1.7 times as long as wide, strongly convex, narrowed and subtruncate 
apically; humeri entirely obsolete; postscutellar impression absent. Surface smooth, 
glossy, distinctly punctured; punctation generally somewhat finer and sparser than on 
head, especially in apical third, at places somewhat denser (posterior setose band, see 
below). Setation similar to that on head, appressed, with scattered short and incon- 
spicuous erect setae, mostly pale, some appressed setae whitish and thicker, forming 
two vague, transverse bands, situated in basal third and shortly behind midlength of 
elytra, some whitish setae scattered also laterally and near base. Metathoracic wings 
almost entirely reduced. 

Legs: Penultimate tarsomeres rather narrow, terminal tarsomere subapical in 
metatarsi; mesotibiae modified, moderately curved inwards and strongly excavated on 
inner side subapically (Figs 75, 76); metatibiae moderately curved inwards apically 
and thus with inner apical margin slightly produced, with single apical spur. Setation 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 65 1 

largely normally developed; mesotibiae with some longer setae on inner side, near/on 
margins of subapical excavation. 

Abdomen: Sternum VII simple, slightly unevenly rounded posteriorly, its apical 
margin shortly, densely setose. Tergum VII evenly rounded posteriorly. Sternite VIII 
(Figs 77, 78); paired prongs rather simple, wide at base, strongly narrowed before 
midlength, curved ventrad, dorso-ventrally flattened and convergent in apical half, 
their apex rounded; each prong with a tuft of conspicuously long, stiff setae on ventral 
side, and with some short, stiff setae on/along apical margin. Tergite VIII simple, 
rounded posteriorly, with apical margin slightly emarginate medially. 

Aedeagus (Fig. 79): Apical portion of tegmen 0.7 times as long as basal-piece, 
parallel-sided, trilobed apically, middle apical lobe wide basally, nearly evenly 
narrowing towards pointed apex, moderately longer than somewhat divergent lateral 
lobes. 

Female: Externally identical with male, except as follows: metatibiae simple, 
with two apical spurs; metasternum simple, lacking paired protrusions; middle legs 
simple, rather shortly setose; sternum VII with posterior margin quite evenly rounded. 

Variability: Distinctly variable in prominence of surface microsculpture; some 
specimens from Struisbaai and all from Alexandria (Woody Cape) and Sedgefield with 
surface of head and pronotum at least partly rather smooth and glossy. Similarly, 
surface of elytra in some specimens uneven and somewhat less glossy. 

Differential diagnosis: Anthelephila vanhillei sp. n. differs from the most 
closely related and possibly sympatric species, A. subtruncata, by the pronotum rather 
shallowly impressed (less constricted) postero-laterally in dorsal view, by the 
conspicuously modified male mesotibiae, male sternum VII rounded posteriorly, and 
by the form and setation of the prongs of male sternite VIII (cf. Figs 72, 73 and 77, 78). 

Distribution: South Africa. 

ACKNOWLEDGEMENTS 

I am very grateful to all curators and colleagues who enabled study the 
specimens in their care, namely: M. Barclay (BMNH), N. Berti and A. Taghavian 
(MNHN), B. Brancucci (NHMB), D. S. Chandler (University of New Hampshire, 
Durham, USA), R. Danielsson (ZILS), E. Grobbelaar (SANC), J. Jelinek (NMPC), 
A. Kirk-Spriggs (AMGS), D. Kovac (SMF), I. Lobi and G. Cuccodoro (MHNG), 
O. Merkl (HNHM), R. Müller (TMP), H. Schönmann (NHMW), W. Schawaller 
(SMNS), M. Uhlig and B. Jäger (MNHB), G. Uhmann (Pressath, Germany), and 
A. Degiovanni (Bubano, Italy). My sincere thanks are due also to D. S. Chandler, 
G. Nardi (Cisterna, Italy) and M. Sakai (Ehime University, Matsuyama, Japan) for 
providing literature copies and comments on various problems. 

REFERENCES 

Blair, K. G. 1942. Coleoptera Heteromera from Guam. Bernice P. Bishop Museum, Bulletin 

172: 56-60. 
Bonadona, P. 1978. Coléoptères Anthicidae du P.I. M. E. Entomological Museum de Detroit. 

Bollettino della Società Entomologica Italiana 110: 12-11. 



652 z - KEJVAL 



Bonadona, P. 1984. Anthicides nouveaux ou peu connus d'Afrique Noire (Coleoptera, 

Anthicidae). Revue de Zoologie africaine (Tervuren) 98: 469-504. 
Bonadona, P. 1989. Anthicidae (Coleoptera) nouveaux des collections du Muséum d'histoire 

naturelle de Genève. Revue suisse de Zoologie 96: 253-276. 
Chandler, D. S. 1982. A Revision of North American Notoxus with a Cladistic Analysis of the 

New World Species (Coleoptera: Anthicidae). Entomography 1: 333-438. 
Chandler, D. S. 2000. Publication Dates of Papers on the Anthicidae (Coleoptera) by the 

Marquis F. T. de LaFerté-Sénectère. Transactions of the American Entomological Society 

125(4)[1999]: 433-439. 
Gemminger, M. 1870. [replacement names for species and genera]. Coleopterologische Hefte 6: 

119-123. 
Hexe, J.C. van 1961: Chapter III. Coleoptera: Anthicidae. (pp. 217-258). In: Hanström, B., 

Brinck, P. & Rudebeck, G. (eds). South African Animal Life 8. Almqvist & Wiksells, 

Uppsala. 
Hua, L. 2002. List of Chinese Insects, Voll. II. Zhongshan (Sun Yat-sen) University Press, 

Guangzhou, 612 pp. 
Kejval, Z. 2003. The genus Anthelephila (Coleoptera: Anthicidae). European Journal of 

Entomology 100: 381-392. 
Krekich-Strassoldo, H. von 1913. Neue Anthiciden und Mitteilungen über die Verbreitung 

bekanntner Anthiciden. Wiener Entomologische Zeitung 32: 223-232. 
Krekich-Strassoldo, H. von 1931. Beiträge zur Kenntnis indischer Anthiciden II. Folia 

Zoologica et Hydrobiologica 3: 1-41, 3 pis. 
Lafer, G. Sh. 1996. 101. Farn. Anthicidae. (pp. 26-45). In: Ler, P.A. (ed.): A key to the insects 

of the Far East of the USSSR in six volumes. Vol. 3. Coleoptera or beetles. Part 1. 

Dalnauka, Vladivostok (in Russian). 
LaFerté-Sénectère, F. T. de 1849a. Anthelephilus. In: Guérin-Méneville, F.-E. (ed.): Species 

et iconographie générique des Animaux Articulés ou représentation des genres, avec leur 

description et celle de toutes les espèces de cette grande division du règne animal. 

Première Partie: Insectes Coléoptères. Livraison 6, No. 24. Fain et Thunot, Paris, 5 pp., 

lpl. 
LaFerté-Sénectère, F. T. de 1849b. Monographie des Anthicus et genres voisins. Coléoptères 

Hétéromères de la tribu des Trachélides. Sapia, Paris, 340 pp. 
Miwa, Y. 1931. A systematic catalogue of Formosan Coleoptera. Department of Agriculture, 

Government Research Institute of Formosa, Taipei 55: xi + 359 pp. 
Nietner, J. 1856. Entomological papers - being descriptions of new Ceylon Coleoptera, with 

such observations on their habits as appear any way interesting. Journal of the Asiatic 

Society of Bengal [New Series] 25:523-539. 
Nietner, J. 1857. Descriptions of new Ceylon Coleoptera. The Annals and Magazine of Natural 

History [Second series] 19: 374-388. 
Nomura, S. 1962. The Anthicid-Beetles from the Loochoo Islands. The Entomological Review 

of Japan 14: 1-7. 
Pic, M. 1899. Descriptions d' Anthicidae exotiques. Revue française d'Entomologie 18: 105-106. 
Pic, M. 1903. Anthicides de Manille. Annales de la Société entomologique de France 71 (1902): 

643-648. 
Pic, M. 1916a. Notes et descriptions abrégées diverses. Mélanges exotico - entomologiques 17: 

2-8. 
Pic, M. 1916b. Diagnoses spécifiques. Mélanges exotico - entomologiques 17: 8-20. 
Sakai, M. 1989. Anthicidae. In: Hirashtma, Y. (ed.): A check list of Japanese insects, Vol. 1. 

Kyushu University, Fukuoka, xi+ 1-540 pp. 
Telnov, D. 2003. Trictenotomidae und Anthicidae (Insecta: Coleoptera) des Himalayas und an- 
grenzender Regionen, Teil 1. Systematik, Faunistik, Zoogeographie (pp. 279-303). In: 

Hartmann, M. & Baumbach, H. (eds.): Biodiversität und Naturausstattung im 

Himalaya. Verein der Freunde und Förderer des Naturkundemuseums Erfurt e. V., 389 

pp., 16 pis. 



STUDIES OF THE GENUS ANTHELEPHILA HOPE 653 



Uhmann, G. 1981. Anthiciden (Coleoptera) aus Africa, Teil 2. Annales Historico-Naturales 

Musei Nationalis Hungarici 73: 187-196. 
Uhmann, G. 1983. Anthiciden der orientalischen Region (Coleoptera, Anthicidae). Annales 

Historico-Naturales Musei Nationalis Hungarici 75: 185-206. 
Uhmann, G. 1986. Anthicidae aus Nepal, II. (Insecta: Coleoptera). Senckenbergiana Biologica 

66: 305-310. 
Uhmann, G. 1987. Anthiciden der orientalischen Region aus dem Museum in Genf (Coleoptera, 

Anthicidae). Revue suisse de Zoologie 94: 687-701. 
Uhmann, G. 1988. Die Anthicidae der Sammlung Johann Klapperich (Coleoptera). Bonner 

Zoologische Beiträge 39: 395-407. 
Uhmann, G. 1989. Anthicidae (Coleoptera) der orientalischen Region aus dem Natur- 
historischen Museum in Genf, II. Revue suisse de Zoologie 96: 243-252. 
Uhmann, G. 1990a. Anthicidae aus Nepal III. Entomofauna, Zeitschrift für Entomologie 11: 

333-347. 
Uhmann, G. 1990b. Anthicidae aus dem Oberösterreichischen Landesmuseum in Linz 

(Coleoptera, Anthicidae). Linzer biologische Beiträge 22: 577-595. 
Werner, F. G. 1965. Insects of Micronesia, Coleoptera: Anthicidae. Insects of Micronesia 16: 

255-269. 
Werner, F. G. & Chandler, D. S. 1995. Anthicidae (Insecta: Coleoptera). Fauna of New 

Zealand 34. Manaaki Whenua Press, Lincoln, 59 pp. 



Revue suisse de Zoologie 1 14 (3): 655-662; septembre 2007 



Re-description of the holotype of Vipera eriwanensis (Reuss, 1933) 
(Serpentes: Viperidae) 

Laszló KRECSÀK 

Eötvös Lorând University, Department of Systematic Zoology and Ecology, Pâzmâny 

P. s. 1/C, H-1117 Budapest, Hungary. E-mail: lkrecsak@gmail.com 



Re-description of the holotype of Vipera eriwanensis (Reuss, 1933) 
(Serpentes: Viperidae). - The missing holotype of Vipera eriwanensis 
described by Reuss in 1933 is re-described, and the questions relating to the 
source, number of individuals and species in the type series are answered. 
A review of the papers by Reuss relating to this taxon is made, and a 
complete chresonym list is given. The decisions leading to the designation 
of a neotype are discussed, remarks are made on the source of the indi- 
vidual, the revalidation of the holotype and accordingly the change of the 
type locality is proposed. 

Keywords: Taxonomy - neotype - holotype - type locality - Armenia. 

INTRODUCTION 

The systematics of the steppe vipers of the V. ursinii and V. renardi groups is 
one of the most debated subjects in viperid taxonomy. Recently, Nilson & Andren 
(2001) evaluated the taxonomy of these groups and answered, to some extent, many of 
the questions. 

Vipera eriwanensis is a small sized viper from the Vipera (Acridophaga) ursinii 
complex with a distribution restricted to the Armenian plateau, including the political 
regions of Armenia, western Azerbaijan and eastern Turkey (Nilson & Andren, 2001). 

The taxon was described by Reuss (1933) as Acridophaga (renardi) eriwanen- 
sis based on a male specimen in the Zoological Museum, Berlin (ZMB) collected 
according to Reuss by Prof. Ramme during a trip in 1929. The author set the type 
locality as: near Eriwan (Yerevan), at around 2000 m altitude. The one sentence long 
description contained just three characters, 21 dorsal scales, 140+1 ventral scales and 
9 supralabial scales, thus this diagnosis may fit other Viperid taxa in the area as well. 
A more detailed description, containing beneath the previously mentioned three 
characters the number of subcaudal scales (37 1 / 2 on the left, 37 on the right plus 
1 scale), and accompanied by a drawing (Fig. 5a on page 215) and a photograph of the 
head of this individual (Fig. 5 on page 217), was published later (Reuss, 1935b) (Fig. 
2). Interestingly, the details on the source of the individual do not agree in the two 
works. Reuss (1935b) noted that the specimen was collected by Prof. Dr Ramme and 
head of preparatory Richter from the Zoological Museum, Berlin during a research trip 



Manuscript accepted 19.02.2007 



656 L. KRECSAK 

in 1929/30. He also mentioned that he had the snake alive for some time, but now, it is 
in the teaching collection ("Lehrsammlung") of the Museum. 

In general nobody wanted to deal with the taxa described by Reuss. He was a 
splitter, describing in fifteen years over 50 new taxa of Viperidae from Europe and 
Asia. Reuss was severely criticized about his publications throughout his life (e.g. 
Lankes, 1925; Müller, 1929; Werner, 1930; Stucken, 1935) as these new taxa did not 
resolve any questions, but led to complete chaos in the European and Asian Viperidae 
taxonomy. Many of the Reuss types could not be found, and some of his taxa were 
never mentioned in lists of synonymy. The only one who tried, and gave a list of some 
taxa and types described by Reuss, was Schwarz (1936). Most probably his identi- 
fication of the type status of many individuals was based on the personal communi- 
cations with Reuss and not published data. 

Schwarz (1936) had not listed the specimen from Yerevan as being one of the 
known museum specimens at that time. This might have been the reason why Kramer 
(1961) stated that the type had disappeared. His statement was taken as a fact, and later 
several authors, with or without reference to Kramer's work, stated that the type is lost 
(Saint Girons, 1978; Joger, 1984; Bruno, 1985; Golay et al, 1993; McDiarmid et al, 
1999; Nilson et al, 1999; Kutrup et al, 2005). It seems that Kramer (1961) was wrong 
in this case and with some other Reuss types as well. He also stated that the type of 
Acridophaga uralensis (presently a synonym of Vipera renardi renardi) stored in the 
ZMB was destroyed, which is not true; it is still preserved in the collection (Holotype 
ZMB 2856, Paratype ZMB 65910). 

Confusion was caused by a photograph published in 1929 (Reuss, 1929). The 
short paper on snake venoms (Reuss, 1929) included four photographs of different 
Vipera species, one of them depicting a living small sized viper, which was not the 
same as that depicted by Reuss in 1935 (Reuss, 1935b). Nilson et al (1999) and Nilson 
& Andren (2001) supposed that the type series consisted of more than a single 
specimen. This was based on personal communication with Erich Sochurek and a 
reproduction of the 1929 photograph which according to Sochurek showed the type. 
Nilson et al (1999) argued that this photograph of a live specimen shows a viper of the 
Vipera kaznakovi complex, which might be V darevskii, V. pontica or V dinniki, thus 
in the absence of type material and for reasons of stability considered it appropriate to 
designate a neotype. Actually Reuss had only a single male individual, and all of his 
remarks about this taxon were based on this specimen (Reuss, 1933, 1935b, 1937). The 
misleading picture (p. 37 in Reuss, 1929) shows a viper of the Vipera ursinii complex, 
probably V.u. macrops, which is also stated in the photograph's caption ("Orsinsche 
Kreuzotter"-Ursinii Adder). The name erhvanensis has not been used in this paper, and 
no other paper published before the description (i.e. Reuss, 1933). The slide archive of 
the Natural History Museum in Vienna houses a reproduction of this photograph, 
which was probably sent to Nilson and co-workers by Erich Socurek. This work of 
Reuss has been erroneously cited as "Fünf Fotos von Giftschlangen" instead of 
"Schlangengift-ein Verjüngungsmittel?" (Kramer, 1961; Nilson et al, 1999; Nilson & 
Andren, 2001). 

A neotype was designated by Nilson et al (1999), an adult male (GNM Re.ex. 
5158) collected on the mountain Ara-Iler, Armenia, north of Yerevan in May 1972 and 



THE HOLOTYPE OF VIPERA ERIWANENSIS 



657 





Fig. 1 
The holotype: dorsal view (A), ventral view (B), dorsal view of the head (C), and lateral view of 
the left side of the head (D). Photographs by Michael Franzen. 



donated by the Zoological Institute, St. Petersburg to Goteborg Natural History 
Museum. 

The V. eriwanensis type individual was rediscovered in September 2005 during 
a revision of the Vipera material, including the Reuss collection, housed in the ZMB. 

The specimen catalogued as ZMB 55160 remained unnoticed in the teaching 
collection for several decades; it was probably moved into the main collection in the 
1990s and received the actual catalogue number in 1996. The snake was donated by the 
collectors, Ramme and Richter, to the ZMB on 9 May 1928 (together with 21 other 
amphibians and reptiles from Persia, Armenia and Caucasus under number C 869 of 
the access-catalogue). The actual jar contains an original label written by the collectors 
containing the following information: Armenia Goktscha-See (Lake Sevan) above 
Schordsha (Shordzha), at 2000 m altitude, 2.IX.1927 f21.XI.1927, Ramme-Richter 
S. G., Acridophaga renardi ("Bergform"-Mountain form), weight 29 in alcohol, before 
28 gr. 

The area of Shordzha lays within the known range of the species and is a well 
known locality for Vipera eriwanensis (Chernov, 1939; Orlov & Tuniyev, 1990; Nilson 
et al, 1995; Aghasyan, 1996; Nilson & Andren, 2001). 



658 L - KRECSAK 

We can just hypothesize why Reuss has written the locality Yerevan. One 
possibility might have been, that he chose this well known locality because only a few 
people were aware of the location of the Lake Sevan. 

CHRESONYMY 

A.[cridopahaga\ {renardi) eriwanensis Reuss, 1933: 373. 

Acridophaga renardi eriwanensis-Reuss (1935a: 152). 

Acridophaga renardi eriwanensia-Reuss (1935b: 216, 215-Fig. 5a, 217-Fig. 5). [Incorrect 
spelling, lapsus for eriwanensis, hence not available (ICZN 1999: Art. 33.3.).] 

Vipera ursinii renardi (part.)-Schwarz (1936: 186). 

A.[cridophaga] renardi eriwarensis-Reuss (1937: 1788). [Incorrect spelling, lapsus for eriwa- 
nensis, hence not available (ICZN 1999: Art. 33.3.).] 

Vipera ursinii renardi (part.)-Mertens & Wermuth (1960: 203). 

Vipera ursinii renardi (part.)-Kramer (1961 : 698, 699, 700, 701, 715). [Assigned to the western 
group of his southeastern and southwestern steppe form-"südöstliche und südwestliche 
Steppenform".] 

Acridophaga eriwanensis-Kramer (1961: 715). [As cited in the synonymy of Vipera ursinii re- 
nardi.] 

Vipera ursinii ebneri (part.)-Saint Girons (1978: 582, 583). 

Vipera ursinii eriwanensis (part.)-Joger (1984: 62, 63). 

Vipera ursinii eriwanensis (part.)-Golay et al. (1993: 290). 

Vipera ursinii eriwanensis (part. )-Bruno (1985: 74). 

Vipera ursini eriwanensis-Or\o\ & Tuniyev (1990: 2, 6, 8, 10, Piate 3, 19-Fig. 9, 22, 23, 24, 30- 
Fig. 16, 31). [Incorrect subsequent spelling for ursinii.] 

Vipera eriwanensis-Höggren et al. (1993: 12, 17). 

Vipera wrs/m'z'-McDiarmid et al. (1999: 409). 

Vipera eriwanensis-Nüson & Andren (2001: 218, 219, 220, 221, 222, 223, 224). 

Vipera renardi eriwanensis-iogtr & Dely (2005: 343, 345, 348, 349). 

DESCRIPTION OF THE HOLOTYPE 

ZMB 55160; adult male; Lake Sevan above Shordzha at 2000 m a.s.l. [Yerevan], 
Armenia; Ramme and Richter leg. 2. IX. 1927; Ramme and Richter don. 9.V.1928. 

Body proportions (all in mm; measures right/left; *un-measurable): body 
length 370; tail length 55; head length (from the corner of the mouth) 16.32; head 
width (at its widest part) 10.55; head depth (behind the eyes) 5.13; distance between 
the eyes (measured on the dorsal edge of the eyes, across the head) 6.87; distance 
between the eyes and the rostrum (between the anterior edge of the eye and the 
rostrum) 5.49/5.61; eye vertical diameter (longest diameter) */2.02; eye horizontal 
diameter (longest diameter) 2.65/2.75; distance between the lower edge of the eyes and 
the edge of the mouth */1.96; fontal length 4.27; frontal width 2.98; rostral length 2.65; 
rostral width (distance between the contact points with the apical scale) 0.70; rostral 
width (distance between contact points with the nasorostral scales) 2.59; nasal diameter 
(longest) 2.67/2.69; nostril diameter (longest) 1.20/1.24. 

Scalation (counts right/left; *un-countable): 4 preventrals; 135 ventrals; 23 
dorsals on the neck (counted at the 10 th ventral); 21 on midbody (in the middle of the 
rostrum-cloaca length); 17 on the tail (5 ventrals from the anal scale); last row with 23 
dorsal scales, at 14 th ventral; first row with 21 dorsals, at the 15 th ventral; reduction 
from 21 to 19 dorsals, at the 116 th ventral; reduction from 19 to 17 dorsals, 131 st 
ventral; subcaudals 36/36 1 /! + 1; 9/9 supralabial scales; forth and fifth supralabials 



THE HOLOTYPE OF VIPERA ERIWANENSIS 



659 




Fig. 2 
Figure 5 and drawing 5a from the 1935 work by Reuss (1935b). 



bellow the eyes; 10/10 sublabial scales; undivided rostral; rostral separated from the 
nasal scales by 1/1 nasorostral scale; nasal scales divided into 2 in the posteriour 
middline; nostrils situated on the bottom of the nasal scales; 5/5 loreal scales; 11/9 
scales in the first circumocular row; 2/3 scales situated between the circumocular row 
and the supralabials, but do not form a complete second circumocular row; two big, 
undivided supraocular scales, separated from the frontal by 3/3 smaller scales; parietals 
big, not fragmented; one apical scale; 2/2 canthal scales; 5 intercanthal scales; 8 scales 
between the eyes (total number of scales between the supraoculars); 4 second 
chinshields; 6/5 scales on the sides of the mental row (the scales in first longitudinal 
rows are counted); */30 gular scales (total number of gular scales counted, delimited 
by the sublabials and the mental row). 

Colour pattern (pale-caused by the alcohol; counts right/left): Dorsal pattern 
consists of 67/66 brown rounded zigzag windings on the body (the tips of the windings 
counted to the one above the cloaca), and 24/27 on the tail. The windings are bordered 
by a lighter (probably light brown) coloured area, while laterally the ground colour is 
darker. Lateral body pattern consists of narrow brown spots. Head pattern formed by 
two separated dark oblique bands, which run from the frontal scale to the lateral side 
of the head a bit backwards from the corner of the mouth. Two darker spots can also 
be observed: one covering the frontal, supraocular and intersupraocular scales the other 
the apical, canthal and intercanthal scales. Laterally two dark, 2-3 scales wide, bands 
are running from the corner of the eye to the corner of the mouth. Ventral side, both on 
body and tail, light but dark spotted, throat light (Fig. 1). 



660 L. KRECSÂK 



REMARKS 

My scale counts greatly agree with Reuss'. The same number of dorsal and 
supralabial scales have been counted. The difference in the number of ventral scales is 
one, 140 according Reuss (counted probably without using a dissection microscope) 
and 139 (4 preventrals and 135 ventrals) according my counts. Reuss (1935b) counted 
37 l / 2 subcaudal scales on the on the left side, 37 on the right side, and probably 
included into here all paired bigger scales that are found on the ventral surface of the 
tail bellow the anal scale. I did not include the first pair of scales which are situated 
bellow the anal plate but are not connected with each other. As it can be assessed with 
certainty that the individual rediscovered is the holotype described by Reuss in 1933 
and depicted on the figures of the 1935 work (Fig. 2), this should regain its type status, 
according to the Art. 75.8. of the International Code of Zoological Nomenclature 
(ICZN, 1999) and the neotype should be set aside. The erroneous type locality, 
Yerevan, should be corrected to with Lake Sevan above Shordzha at 2000 m a.s.l., the 
correct collection locality of the specimen according to the Recomendation 76A.2. of 
the Code (ICZN, 1999). 

ACKNOWLEDGEMENTS 

I wish to thank the help, hospitality and kindness of Dr Rainer Günther during 
my stay in ZMB, for providing all the background information about the holotype and 
for his comments on the manuscript. Michael Franzen kindly made the photographs of 
the individual. Dr Wolfgang Böhme (ZFMK, Bonn), Dr Ulrich Joger (SNHM, 
Braunschweig), Norbert Otte (Berlin) and Dr Tigran Tadevosyan (Centre for 
Ecological-Noosphere Studies, Yerevan) provided relevant literature. Dr Antonia 
Cabela and Dr Heinz Grillitsch made available a copy of the photograph by Erich 
Sochurek preserved in the slide archive of the Herpetological Collection of the Natural 
History Museum Vienna. David Bird (Dorset, UK) and Dr Wolfgang Böhme (ZFMK, 
Bonn) are acknowledged for their comments on the manuscript. The specimen was 
discovered during a research visit to the Museum für Naturkunde der Humboldt- 
Universität zu Berlin supported through SYNTHESYS funding, which was made avail- 
able by the European Community-Research Infrastructure Action under the FP6 
"Structuring the European Research Area" Programme, project DE-TAF-1014. 

REFERENCES 

Aghasyan, A. L. 1996. The Fauna of Snakes of Armenia and Nakhichevan. Unpublished 
Candidate Thesis, Natinal Academy of Sciences of Republic of Armenia, Institute of 
Zoology, Yerevan, 34 pp (in Russian). 

ICZN, 1999. International Code of Zoological Nomenclature. Fourth Edition adopted by the 
International Union of Biological Sciences. The International Trust for Zoological 
Nomenclature c/o The Natural History Museum, London, xxix+306 pp. 

Bruno, S. 1985. Le vipere d'Italia e d'Europa. Edagricole, Bologna, xiii+269 pp. 

Chernov, S. A. 1939. Herpetological Fauna of Armenian SSR and Nakhichevan ASSR. USSR 
Academy of Science-Armenian Branch, Zoological Papers, I (Transactions of Biological 
Institute, issue III) 7: 77-194 (in Russian). 

Golay, P. H., Smith, H. M., Broadley, D. G., Dixon, J. R., McCarthy, C, Rage, J.-C, Schätti, 
B. & Toriba, M. 1993. Endoglyphs and other major venomous snakes of the World. A 
checklist. Azemiops S.A. Herpetological Data Center, Aire-Genève, xv+478 pp. 



THE HOLOTYPE OF VIPERA ERWANENSIS 661 



Höggren, M., Nilson, G., Andren, C, Orlov, N. L. & TuNiYEV, B. S. 1993. Vipers of the 

Caucasus: Natural history and systematic review. Herpetological Natural History 1(2): 

11-19. 
Joger, U. 1984. The Venomous Snakes of the Near and Middle East. Tübinger Atlas des 

Vorderen Orients, Dr. Ludwig Reichter Verlag, Wiesbaden, Beiheft 12 A, 115 pp. 
Joger, U. & Dely, O. G. 2005. Vipera (Pelias) renardi (Christoph, 1861)— Steppenotter. In: 

Joger, U. & Stümpel, N. (eds). Handbuch der Reptilien und Amphibien Europas, 3/IIB 

Schlangen (Serpentes) III Viperidae. Aula Verlag, Wiesbaden, pp. 343-354. 
Kramer, E. 1961. Variation, Sexualdimorphismus, Wachstum und Taxionomie von Vipera 

ursinii (Bonaparte, 1835) und Vipera kaznakovi Nikolskij, 1909. Revue suisse de 

Zoologie 68: 627-725. 
Kutrup, B., BÜLBÜL, U. & Yilmaz, N. 2005. On the Distribution and Morphology of the Steppe 

Viper, Vipera eriwanensis (Reuss, 1933), from Gavur Mountain (Gümü hane). Turkish 

Journal of Zoology 29: 321-325. 
Lankes, R. 1925. No title (under Rundschau des Vereinslebens-„Isis", München). Blätter für 

Aquarien- und Terrarienkunde 2: 60-62. 
McDiarmid, R. W., Campbell, J. A. & Touré, T. A. 1999. Snake Species of the World. A 

Taxonomic and Geographic Reference. Herpetologists League, Washington DC, Vol. 1, 

xi+5 1 1 pp. 
Mertens, R. & Wermuth, H. 1960. Die Amphibien und Reptilien Europas. (Dritte Liste, nach 

dem Stand vom 1. Januar 1960). Verlag Waldemar Kramer, Frankfurt am Main, 264 pp. 
Müller, L. 1929. No title (under Vereinsnachrichten-München. „Isis"). Wochenschrift für 

Aquarien- und Terrarienkunde 23: 353-355. 
Nilson, G., Tuniyev, B., Orlov, N., Höggren, M. & Andren, C. 1995. Systematics of the 

Vipers of the Caucasus: Polymorphism or Sibling Species? Asiatic Herpetological 

Research 6: 1-26. 
Nelson, G., Tuniyev, B., Andren, C. & Orlov, N. 1999. Vipers of Caucasus: Taxonomic 

Considerations. Kaupia 8: 103-106. 
Nilson, G. & Andren, C. 2001. The meadow and steppe vipers of Europe and Asia— The Vipera 

(Acridophaga) ursinii complex. Acta Zoologica Academiae Scientiarum Hungaricae 

47(2-3): 87-267. 
Orlov, N. L. & Tuniyev, B. S. 1990. Three Species in the Vipera kaznakowi Complex 

(Eurosiberian Group) in the Caucasus: Their Present Distribution, Possible Genesis, and 

Phylogeny. Asiatic Herpetological Research 3: 1-36. 

Reuss, T. 1929. Schlangengift-ein Verjüngungsmittel? Kölnische Illustrierte Zeitung 12. 1. 1929 
2: p. 37 & p. 43. 

Reuss, T. 1933. Fortsetzung und Schluss der Originalberichte (under Vereinsnberichte- 
„Lacerta", Gesellschaft für Terrarienkunde Berlin). Nachrichtenblatt für Aquarien- und 
Terrarien-Vereine 26: 372-373. 

Reuss, F. A. T 1935a. Europäische Giftschlangen. Uebersichtskarte der Giftschlangenzu- 
wanderung in Europa seit der Eiszeit (Quartärzeit). Mit einer Einführung in die 
entsprechende Aufteilung der europäischen Ottern unter Berücksichtigung von Gift- und 
Heilserumfragen. Deutscher Almanach für Aquarien- und Terrarienfreunde, 1935, 145- 
166. 

Reuss, F. A. T. 1935b. Erläuterungen zur Typenfrage in der Zoologie an hand von Bildern 
europäischer Giftschlangen. Nachrichtenblatt für Aquarien- und Terrarien-Vereine 16: 
215-218. 

Reuss, F. A. T 1937. Observations on four Species of European Toxicophidia. Comptes Rendus 
du XII e Congrès International de Zoologie— Lisabonne 1935: 1787-1804. 

Saint Girons, H. 1978. Morphologie externe comparée et systématique des Vipères d'Europe 
(Reptilia, Viperidae). Revue suisse de Zoologie 85: 565-595. 

Schwarz, E. 1936. Untersuchungen über Systematik und Verbreitung der europäischen und 
mediterranen Ottern. In: Bieling, R., Demnitz, A., Schaumann, O., Schlossberger, H.. 



662 L - KRECSAK 



v. Schuckman, W. & Schwarz, E. (eds). Die europäischen und mediterranen Ottern und 

ihre Gifte. Grundlagen zur Darstellung eines wirksamen Schlangenserums. 

Behringwerk-Mitteilungen, Heft 7, Selbstverlag der Behringwerke Marburg-Lahn, 

159-355+1 map. 
Stucken 1935. No title (under Rundschau des Vereinslebens-München. „Isis", Gesellschaft für 

biologische Aquarien- und Terrarienkunde). Blätter für Aquarien- und Terrarienkunde 6: 

141-142. 
Werner, F. 1930. Über Vipera (Mesocoronis) bosnensis Wem. u. T.Reuss. Zoologischer 

Anzeiger 88(1/4): 89-92. 



REVUE SUISSE DE ZOOLOGIE 
Tome 114 — Fascicule 3 

Pages 

Lienhard, Charles. Description of a new African genus and a new tribe of 

Speleketorinae (Psocodea: 'Psocoptera': Prionoglarididae) 441-469 

Ng, Heok Hee & Kottelat, Maurice. A review of the catfish genus Hara, 

with the description of four new species (Siluriformes: Erethistidae) 471-505 

Roy, Roger. Deroplarys indica, nouvelle espèce de l'Inde (Dicty optera, 

Mantodea) 507-51 1 

Lourenço, Wilson R. Further additions to the scorpion fauna of the 

Guayana region of South America 513-519 

Scheller, Ulf. Records of Pauropoda (Pauropodidae; BrachypauropoT 
didae; Eurypauropodidae) from Singapore, Indonesia and Malaysia 
with the description of 18 new species (Pauropoda and Symphyla of 
the Geneva Museum XV) 521-572 

Stebnicka, Zdzislawa T. New species of Iarupea Martinez and morpho- 
logical specializations among related taxa associated with ants and 
termites (Coleoptera: Scarabaeidae: Eupariini) 573-590 

Azpelicueta, M. de las Mercedes, Almirón, Adriana E., C asciutta, 
Jorge R. & Koerber, Stefan. Hisonotus hungy sp. n. (Siluriformes, 
Loricariidae) a new species from arroyo Tirica, Misiones, Argentina 591-598 

Ziegler, Thomas, Heidrich, Astrid, Schmitz, Andreas & Böhme, 
Wolfgang. Phylogenetic position, morphology and natural history of 
the Vietamese water skink Tropidophorus noggei Ziegler, Vu & Bui, 
2005 (Sauria: Scincidae) 599-614 

Kejval, Zbynëk. Studies of the genus Anthelephila Hope (Coleoptera: 

Anthicidae) - 10. Species related to A. imperatrix 615-653 

Krecsâk, Lâszlo. Re-description of the holotype of Vipera erixvanensis 

(Reuss, 1933) (Serpentes: Viperidae) 655-662 



REVUE SUISSE DE ZOOLOGIE 

Volume 114 — Number 3 

Pages 

Lienhard, Charles. Description of a new African genus and a new tribe of 

Speleketorinae (Psocodea: 'Psocoptera': Prionoglarididae) 441-469 

Ng, Heok Hee & Kottelat, Maurice. A review of the catfish genus Hara, 

with the description of four new species (Siluriformes: Erethistidae) 471-505 

Roy, Roger. Deroplatys indica, new species from India (Dictyoptera, 

Mantodea) 507-511 

Lourenco, Wilson R. Further additions to the scorpion fauna of the 

Guayana region of South America 513-519 

Scheller, Ulf. Records of Pauropoda (Pauropodidae; Brachypauropo- 
didae; Eurypauropodidae) from Singapore, Indonesia and Malaysia 
with the description of 18 new species (Pauropoda and Symphyla of 
the Geneva Museum XV) 521-572 

Stebnicka, Zdzislawa T. New species of Iarupea Martinez and morpho- 
logical specializations among related taxa associated with ants and 
termites (Coleoptera: Scarabaeidae: Eupariini) 573-590 

Azpelicueta, M. de las Mercedes, Almirón, Adriana E., Casciotta, 
Jorge R. & Koerber, Stefan. Hisonotus hungy sp. n. (Siluriformes, 
Loricariidae) a new species from arroyo lìrica, Misiones, Argentina 591-598 

Ziegler, Thomas, Heidrich, Astrid, Schmitz, Andreas & Böhme, 
Wolfgang. Phylogenetic position, morphology and natural history of 
the Vietamese water skink Tropidophorus noggei Ziegler, Vu & Bui, 
2005 (Sauria: Scincidae) 599-614 

Kejval, Zbynëk. Studies of the genus Anthelephila Hope (Coleoptera: 

Anthicidae) - 10. Species related to A. imperatrix 615-653 

Krecsäk, Laszló. Re-description of the holotype of Vipera eriwanensis 

(Reuss, 1933) (Serpentes: Viperidae) 655-662 

Indexed in Current Contents, Science Citation Index 



PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE 

CATALOGUE DES INVERTÉBRÉS DE LA SUISSE, Nos 1-17 (1908-1926) série Fr. 285.— 

(prix des fascicules sur demande) 

REVUE DE PALÉOBIOLOGIE Echange ou par fascicule Fr. 35.— 

LE RHINOLOPHE (Bulletin du centre d'étude des chauves-souris) par fascicule Fr. 35. — 

THE EUROPEAN PROTURA: THEIR TAXONOMY, ECOLOGY AND 
DISTRIBUTION, WITH KEYS FOR DETERMINATION 
J. Nosek, 345 p., 1973 Fr. 30.— 

CLASSIFICATION OF THE DIPLOPODA 

R. L. Hoffman, 237 p., 1979 Fr. 30.— 

LES OISEAUX NICHEURS DU CANTON DE GENÈVE 
P. Géroudet, C. Guex & M. Maire 
351 p., nombreuses cartes et figures, 1983 Fr. 45. — 

CATALOGUE COMMENTÉ DES TYPES D'ECHINODERMES ACTUELS 
CONSERVÉS DANS LES COLLECTIONS NATIONALES SUISSES, 
SUIVI DUNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ 
À LA CONNAISSANCE DES ECHINODERMES ACTUELS 
M. Jangoux, 67 p., 1985 Fr. 15.— 

RADULAS DE GASTÉROPODES LITTORAUX DE LA MANCHE 
(COTENTLN-BAIE DE SEINE, FRANCE) 
Y. Finet, J. Wüest & K. Mareda, 62 p., 1991 Fr. 10.— 

GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN: 
SHELLS AND RADULAS 
Y. Finet, J. Wüest & K. Mareda, 1992 Fr. 30.— 

O. SCHMIDT SPONGE CATALOGUE 

R. Desqueyroux-Faundez & S.M. Stone, 190 p., 1992 Fr. 40.— 

ATLAS DE RÉPARTITION DES AMPHIBIENS 
ET REPTILES DU CANTON DE GENÈVE 

A. Keller, V. Aellen & V. Mahnert, 48 p., 1993 Fr. 15. — 

THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: 
A DOCUMENTED FAUNAL LIST 
Y. Finet, 180 p., 1995 Fr. 30.— 

NOTICE SUR LES COLLECTIONS MALACOLOGIQUES 
DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE 
J.-C. Cailliez, 49 p., 1995 Fr. 22.— 

PROCEEDINGS OF THE XHIth INTERNATIONAL CONGRESS 

OF ARACHNOLOGY, Geneva 1995 (ed. V. Mahnert), 720 p. (2 vol.), 1996 Fr. 160.— 

CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE) 

{Instrumenta Biodiversitatis I), I. LÖBL, xii + 190 p., 1997 Fr. 50. — 

CATALOGUE SYNONYMIQUE ET GÉOGRAPHIQUE DES SYRPHIDAE (DIPTERA) 
DE LA RÉGION AFROTROPICALE 
(Instrumenta Biodiversitatis II), H. G. DlRlCKX, x +187 p., 1998 Fr. 50. — 

A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE 
WEST PALAEARCTIC REGION 
(Instrumenta Biodiversitatis III), S. Bowestead, 203 p., 1999 Fr. 60. — 

THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE 
SOKOTRA ARCHIPELAGO 

(Instrumenta Biodiversitatis IV), B. SchAtti & A. Desvoignes, 
178 p., 1999 Fr. 70.— 

PSOCOPTERA (INSECTA): WORLD CATALOGUE AND BIBLIOGRAPHY 

(Instrumenta Biodiversitatis V), C. Lienhard & C. N. Smithers, 

xli + 745 p., 2002 Fr. 180.— 

REVISION DER PALÄARKTISCHEN ARTEN DER GATTUNG BRACHYGLUTA 
THOMSON, 1859 (COLEOPTERA, STAPHYLINIDAE) (1. Teil) 
(Instrumenta Biodiversitatis VI), G. Sabella, Ch. Buckle, V. Brachat 
& C. Besuchet, vi + 283 p., 2004 Fr. 100.— 

PHYLOGENY, TAXONOMY, AND BIOLOGY OF TEPHRITOID FLIES 
(DIPTERA, TEPHRITOIDEA) 

Proceedings of the "3rd Tephritoid Taxonomist's Meeting, Geneva, 19.-24. July 2004" 
(Instrumenta Biodiversitatis VII). B. Merz, vi + 274 p., 2006 Fr. 100. — 



Volume 1 14 - Number 3 - 2007 

Revue suisse de Zoologie: Instructions to Authors 

The Revue suisse de Zoologie publishes papers by members of the Swiss Zoological Society and scientific 
results based on the collections of the Muséum d'histoire naturelle, Geneva. Submission of a manuscript implies 
that it has been approved by all named authors, that it reports their unpublished work and that it is not being 
considered for publication elsewhere. A financial contribution may be asked from the authors for the impression of 
colour plates and large manuscripts. All papers are refereed by experts. 

In order to facilitate publication and avoid delays authors should follow the Instructions to Authors and refer to a 
current number of R.S.Z. for acceptable style and format. Papers may be written in French, German, Italian and 
English. Authors not writing in their native language should pay particular attention to the linguistic quality of the text. 

Manuscripts must be typed or printed, on one side only and double-spaced, on A4 (210 x 297 mm) or equi- 
valent paper and all pages should be numbered. All margins must be at least 25 mm wide. Authors must submit 
three paper copies (print-outs), including tables and figures, in final fully corrected form, and are expected to retain 
another copy. Original artwork should only be submitted with the revised version of the accepted manuscript. 

We encourage authors to submit the revised final text on a CD-R, using MS-WORD or a similar software. The 
text should be in roman (standard) type face throughout, except for genus and species names which should be for- 
matted in italics (bold italics in taxa headings) and authors' names in the list of references (not in other parts of the 
text!), which should be formatted in Small Capitals. LARGE CAPITALS may be used for main chapter headings 
and Small Capitals for subordinate headings. Footnotes and cross-references to specific pages should be avoided. 
Papers should conform to the following general layout: 

Title page. A concise but informative full title plus a running title of not more than 40 letters and spaces, full 
name(s) and surname(s) of author(s), and full address(es) including e-mail address(es) if possible. 

Abstract. The abstract is in English, composed of the title and a short text of up to 200 words. It should sum- 
marise the contents and conclusions of the paper and name all newly described taxa. The abstract is followed by up 
to 10 keywords, separated by hyphens, which are suitable for indexing. Some of the terms used in the title may be 
omitted from the list of keywords in favour of significant terms not mentioned in the title. 

Introduction. A short introduction to the background and the reasons for the work. 

Material and methods. Sufficient experimental details must be given to enable other workers to repeat the work. 
The full binominal name should be given for all organisms. The International Code of Zoological Nomenclature 
must be strictly followed. Cite the authors of species on their first mention. 

Results. These should be concise and should not include methods or discussion. Text, tables and figures should 
not duplicate the same information. New taxa must be distinguished from related taxa. The abbreviations gen. n., sp. 
n., syn. n. and comb. n. should be used to distinguish all new taxa, synonymies or combinations. Primary types must 
be deposited in a museum or similar institution. In taxonomic papers the species heading should be followed by 
synonyms, material examined, description, distribution, and comments. All material examined should be listed in 
similar, compact and easily intelligible format; the information should be in the same language as the text. Sex sym- 
bols should be used rather than "male" and "female" (text file: $ = â, £ = 9 ). 

Discussion. This should not be excessive and should not repeat results nor contain new information, but should 
emphasize the significance and relevance of the results reported. 

References. The author-date system (name-year system) must be used for the citation of references in the text, 
e.g. White & Green (1995) or (White & Green, 1995). For references with three and more authors the form Brown 
et al. (1995) or (Brown et al., 1995; White et al., 1996) should be used. In the text authors' names have to be 
written in standard type face. However, in the list of references they should be formatted in Small Capitals (see 
below). The list of references must include all publications cited in the text and only these. References must be 
listed in alphabetical order of authors, in the case of several papers by the same author, the name has to be repeated 
for each reference. The title of the paper and the name of the journal must be given in full in the following style: 
Penard, E. 1888. Recherches sur le Ceratium macroceros. Thèse, Genève, 43 pp. 
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. 

Tables. These should be self-explanatory, not integrated in the text-file, with the title at the top, organised to fit 
122 x 180 mm, each table on a separate sheet and numbered consecutively. 

Figures. These may be line drawings or half tones, not integrated in the text-file, and all should be numbered 
consecutively. Figures should be arranged in plates which can be reduced to 122 x 160 mm. Drawings and lettering 
should be prepared to withstand reduction. Magnification should be indicated with scale lines. Authors should refrain 
from mixing drawings and half tones. Originals of figures (ink drawings, photographs, slides) should be submitted 
together with the revised version of the accepted manuscript. Original drawings will not be returned automatically. 
The Revue suisse de Zoologie declines responsibility for lost or damaged slides or other documents. If scanned figures 
are submitted on CD, this should be clearly indicated on the print-out. Scanned line drawings must be saved as TIF files 
in bitmap mode with a resolution of at least 600 dpi. Half tone illustrations and photos must have at least 300 dpi reso- 
lution. 

Legends to figures. These should be typed in numerical order on a separate sheet. 

Proofs. Only page proofs are supplied, and authors may be charged for alterations (other than printer's errors) if 
they are numerous. 

Offprints. The authors receive a total of 25 offprints free of charge; more copies may be ordered at current prices 
when proofs are returned. 

Correspondence. All correspondence should be addressed to 

Revue suisse de Zoologie, Muséum d'histoire naturelle, CP 6434, CH-121 1 Genève 6, Switzerland. 
Phone: +41 22 418 63 33 - Fax: +41 22 418 63 01. E-mail: danielle.decrouez@ville-ge.ch 
Home page RSZ: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 




3 9088 01 495 8144