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

Full text of "Revue suisse de zoologie"

\I_ES 



de la 


SOCIÉTÉ SUISSE DE ZOOLOGIE 


et du 


MUSÉUM D'HISTOIRE NATURELLE 


de la Ville de Genève 


tome 1 1 1 


fascicule 1 


2004 



o 
o 

N 

LU 

o 



LU 
O 

O 






o 
o 

N 

LU 

o 

> I I I 

o ■■■ 



CD 



LU 

> 

LU 

JeL GENEVE MARS 2004 ISSN 0035 - 418 X C^ HÉÌ 



O 



</5 
C/5 



REVUE SUISSE DE ZOOLOGIE 



TOME 111— FASCICULE 1 

Publication subventionnée par: 

Académie suisse des Sciences naturelles ASSN 

Ville de Genève 

Société suisse de Zoologie 



VOLKER MAHNERT 

Directeur du Muséum d'histoire naturelle de Genève 

CHARLES LIENHARD 

Chargé de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des instituts de zoologie des universités 
suisses. 

Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le 
sujet étudié. 

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

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 

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



Prix de l'abonnement: 

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

(en francs suisses) 

Les demandes d'abonnement doivent être adressées 

à la rédaction de la Revue suisse de Zoologie, 

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



ANNALES 

de la 

SOCIÉTÉ SUISSE DE ZOOLOGIE 

et du 

MUSÉUM D'HISTOIRE NATURELLE 

de la Ville de Genève 



tome 1 1 1 
fascicule 1 
2004 




LU 
O 

o 



o 
o 

N 



> 



û 

LU 



O 




> 

uu 

J L GEf JEVE MARS 2004 ISSN 003 5 - 418 X C/) HÉK 



C/5 
CD 



REVUE SUISSE DE ZOOLOGIE 



TOME 111— FASCICULE 1 

Publication subventionnée par: 

Académie suisse des Sciences naturelles ASSN 

Ville de Genève 

Société suisse de Zoologie 



VOLKER MAHNERT 

Directeur du Muséum d'histoire naturelle de Genève 

CHARLES LIENHARD 

Chargé de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des instituts de zoologie des universités suis- 
ses. 

Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le 
sujet étudié. 

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

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 

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



Prix de l'abonnement: 

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

(en francs suisses) 

Les demandes d'abonnement doivent être adressées 

à la rédaction de la Revue suisse de Zoologie, 

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



Revue suisse de Zoologie 111 (1): 3-10; mars 2004 



New earthworms of Pheretima and Pithemera (Oligochaeta: 
Megascolecidae) from Mt. Arayat, Luzon Island, Philippines 

Samuel W. JAMES \ Yong HONG 2 * & Tae Heung KIM 2 

1 Department of Life Sciences, Maharishi University of Management, Fairfield, 
IA 52557, U.S.A. 

2 Faculty of Biological Resources Science, College of Agriculture, Jeonbuk National 
University, Jeonju 561-756, Republic of Korea. 



New earthworms of Pheretima and Pithemera (Oligochaeta: Mega- 
scolecidae) from Mt. Arayat, Luzon Island, Philippines. - Three new 
species of Pheretima and two new species of Pithemera are described from 
Mt. Arayat: Pheretima arayatensis sp. n., Pheretima simsi sp. n., and Phere- 
tima castilloi sp. n., Pithemera rotunda sp. n., and Pithemera philippinensis 
sp. n. Pheretima arayatensis sp. n., Pheretima simsi sp. n., and Pheretima 
castilloi sp. n. have spermathecae in segment ix, vi, and vi-viii, respectively. 
Pithemera rotunda sp. n., and Pithemera philippinensis sp. n. have sper- 
mathecae in segment v-ix. The former species has numerous circular genital 
markings, but the latter Pithemera species has no genital markings. 
Descriptions of the new species are provided, including illustrations of the 
ventral view, male pore region, and spermathecae. 

Key- words: Earthworms - Pheretima - Pithemera - Megascolecidae - 
Oligochaeta - Mt. Arayat - Philippines - taxonomy. 

INTRODUCTION 

Mt. Arayat is an isolated volcanic mountain in the large plains of central Luzon, 
north of Manila. Its forested upper elevations have always been isolated from other 
mountain ecosystems, such as the central Cordillera to the north and west, the 
Caraballo Range to the north, and the southern Sierra Madre to the east. The remain- 
ing forests of Mt. Arayat are an important refuge for biota whose habitats would other- 
wise be destroyed in the region. Its biogeographic importance as an isolated peak is 
now accentuated by deforestation and conversion of land to intensive agriculture. As 
part of a biotic survey of the earthworms of the Philippines, specimens were collected 
from Mt. Arayat in March, 2001. In this paper, we report three new species of 
Pheretima Kinberg, 1867 and two new species of Pithemera Sims & Easton, 1972. 
James (2004) described 21 new species of Megascolecidae of which 18- were new 
species of the genus Pheretima from the Mt. Kitanglad Range, Mindanao Island. He 
followed the systematics of Sims & Easton (1972). These authors assigned species 



* To whom correspondence and reprint requests should be addressed 
E-mail: geoworm@hanmail.net 
Manuscript accepted 27.10.2003 



4 S. W. JAMES ETAL. 

with the intestinal caeca(um) originating in or near xxii to this genus, Pithemera. They 
also defined four species groups within Pithemera, one of which is the bicincta group, 
containing the generotype Pithemera bicincta (Perrier, 1875). We report 2 new species 
of the bicincta group herein. 

Holotypes and paratypes are deposited in the National Museum of Annelids of 
the Philippines (NMA). Paratypes are deposited in the Field Museum of Natural 
History (FMNH), and the Museum of Natural History of Geneva (MHNG). 

DESCRIPTIONS 

Pheretima arayatensis James & Hong sp. n. Figs 1A-B 

Material: Holotype: One clitellate (NMA 0003741): Philippines, Luzon Island, 
Pampanga Province, Mt. Arayat (15°12.29'N, 120°43.25'E), 410 m, litter layers in forest, 14 
March 2001, Y. Hong & A. Castillo colls. 3 paratypes: 1 clitellate (NMA 0003746), 1 clitellate 
(FMNH 10012), 1 clitellate (MHNG 34807): Same data as for holotype. Other material: Same 
data as for holotype, 4 clitellate, 18 aclitellate specimens; Mt. Arayat (15°12.34'N, 120° 
43.90'E), 330 m, litter layers in forest, 1 clitellate, 1 semiclitellate, 13 March 2001. 

Etymology: The species is named for its type locality. 

Diagnosis: Spermathecal pore in 8/9, brown dorsal pigment, genital marking 
absent. 

Description: Brown dorsal pigment. Dimensions 88-141 mm by 5.8-6.3 mm at 
segment x, 7.0-7.2 mm at xxx, 5.1-5.5 mm at clitellum, segments 96-115; body cylin- 
drical in cross-section. Setae regularly distributed around segmental equators, number- 
ing 54 at vii, 87 at xx; 5-6 between male pores, size, distance regular; setal formula 
AA:AB:YZ:ZZ = 2:1:2:4 at xiii. Clitellum annular xiv-xvi; setae invisible externally. 

First dorsal pore 12/13, spermathecal pore midventral deep in intersegmental 
furrow 8/9, with thickened surrounding tissue, monothecate. Female pore single in xiv, 
0.3 mm oval shape, 0.5 mm openings of copulatory bursae paired in xviii, distance 
between male pores 2.1 mm. Genital marking lacking. 

Septa 5/6-7/8 thick, 8/9 absent, 9/10 thin, only a small membrane connected to 
10/11, not to body wall, 10/11-13/14 thick, muscular. Gizzard large in viii-x, intestine 
begins in xv, small paired lymph glands from xxviii along dorsal vessel; intestinal 
caeca simple, originating in xxvii, and extending anteriorly to xxii, each consisting of 
a finger-shaped sac; typhlosole low simple fold from xxvii. Hearts x-xiii esophageal; 
ix lateral, right side large, left very reduced. 

Ovaries and funnels in xiii, single spermatheca in ix with nephridia on sper- 
mathecal ducts; spermatheca with small ampulla, ducts shorter than ampulla, 2 diver- 
ticula each composed of 5 separate chambers on single stalks or double stalks termi- 
nating in two chambers, club-shaped. Male sexual system holandric, testes and funnels 
in ventral paired sacs in x, xi. Seminal vesicles two pairs in xi, xii with dorsal lobes, 
pseudovesicles xiii, prostates in xviii, with stout muscular duct entering center of the 
copulatory bursae without stalked glands. Vas deferens muscular xiv-xviii. Pads or 
other structures like penis on inside of copulatory bursae. 

Remarks: The species shares the brown dorsal pigment, dimensions, setae, and 
monothecate condition with Pheretima ambonensis Cognetti, 1913 from Ambon, 
Indonesia. Pheretima arayatensis has 2 spermathecal diverticula each composed of 5 



PHERETIMA AND PITHEMERA FROM MT. ARAYAT, PHILIPPINES 






Fig. 1 
A-B. Pheretima arayatensis sp. n. A: ventral view; B: spermathecae. C-D. Pheretima simsi sp. 
n. C: ventral view; D: spermathecae. E-F. Pheretima castilloi sp. n. E: ventral view; F: sper- 
mathecae. Scale bars = 5 mm (A, C, E), 2 mm (B), 1 mm (D, F). 

separate chambers on single stalks or double stalks terminating in two chambers, but 
P. ambonensis has a thick duct and only 2 diverticula on single stalks. Also Pheretima 
arayatensis has its spermathecal pore in 8/9 and spermatheca in ix, but P. ambonensis 
has the spermathecal pore in 7/8 and spermatheca in vii. 



Pheretima simsi James & Hong sp. n. 



Figs 1C-D 



Material: Holotype: One clitellate (NMA 0003742): Philippines, Luzon Island, 
Pampanga Province, Mt. Arayat (15°12.41'N, 120°44.45'E), 1050 m, litter layers in forest, 13 
March 2001, Y. Hong & O. Castillo colls. 3 paratypes: 1 clitellate (NMA 0003747), 1 clitellate 
(FMNH 10013), 1 clitellate (MHNG 34808): Same data as for holotype. Other material: Same 
data as for holotype, 4 clitellate, 6 aclitellate specimens; Mt. Arayat (15°12.34'N, 120°43.90'E), 
330 m, litter layers in forest, 9 clitellate, 2 aclitellate, 13 March 2001. 

Etymology: Named for Dr R. W. Sims, who made good contributions to earth- 
worm systematics. 

Diagnosis: Spermathecal pores in 5/6. Diverticulum, as long as ampulla, cham- 
ber and stalk showing spermatozoal iridescence, nephridia covering duct-ampulla 
junction, part of ampulla. 

Description: Brown dorsal pigment. Dimensions 68-91 mm by 2.6-2.8 mm at 
segment x, 2.8-3.1 mm at xxx, 2.7-3.0 mm at clitellum, segments 82-97; body cylin- 



6 S. W. JAMES ETAL. 

drical in cross-section. Setae regularly distributed around segmental equators, number- 
ing 31-33 at vii, 39-47 at xx; 6-8 between male pores, size, distance regular; setal for- 
mula AA:AB:YZ:ZZ = 1.5:1:2:3 at xiii. Clitellum annular xiv-xvi; setae invisible 
externally. 

First dorsal pore 11/12, 12/13, spermathecal pores in 5/6, at lateral margin of 
ventrum, white color, with slightly protuberant lips. Female pore single in xiv, 0.4 mm 
oval shape, 0.5 mm openings of copulatory bursae paired in xviii, distance between 
male pores 2.5 mm. Genital marking lacking. 

Septa 5/6-8/9 thin, 9/10 absent, 10/11-12/13 thick, muscular, 13/14 thin. 
Gizzard usual in viii, intestine begins in xv, small paired lymph glands from xxviii 
along dorsal vessel; intestinal caeca simple, originating in xxvii, and extending 
anteriorly about to xxvi, each consisting of a finger-shaped sac; typhlosole low simple 
fold from xxvii. Hearts x-xiii esophageal; ix lateral, viii to gizzard, vii lateral, with 
branches to anterior gizzard wall. 

Ovaries and funnels in xiii, paired spermathecal in vi with nephridia on sper- 
mathecal ducts; spermatheca with small ampulla, duct very short, diverticulum, as long 
as ampulla, chamber and stalk showing spermatozoal iridescence, nephridia covering 
duct-ampulla junction, part of ampulla. Male sexual system holandric, testes and 
funnels in ventral paired sacs in x, xi. Seminal vesicles two pairs in xi, xii with narrow 
dorsal lobes, prostates in xvii- xix, with stout muscular duct entering center of the 
copulatory bursae without stalked glands. Copulatory bursa opening flanked by 
anterior, posterior pads, male pore within large convoluted opening on roughly conical 
protrusion from copulatory bursa roof. 

Remarks: Pheretima simsi keys to the urceolata group in Sims & Easton (1972), 
which is composed of two species, P. urceolata Horst 1893 and P. baweanensis 
Michaelsen, 1924. Gates (1961) synonymized these two species and P. ditheca 
Michaelsen, 1928. The species shares the lack of setae in the clitellum, 31 setae per 
segment at vii, and spermathecal pores in 5/6 with P. urceolata. Pheretima simsi 
differs from P. urceolata in having as the ampulla long as the diverticulum, and in hav- 
ing short caeca. The locality of Pheretima simsi is close to human disturbance and 
activity, as was the collection site of P. urceolata. The present species is similar to 
Pheretima kitangladensis (James, 2004), but P. kitangladensis has no dorsal or ventral 
setal gaps, and has vertical lamellae in the esophagous. Also, P. kitangladensis is quite 
remote from human disturbance at a high elevation site (2250 m), but Pheretima simsi 
and P. urceolata were collected from low and mid- elevation sites (330-1050 m, 1100 
m). P. urceolata was collected from Sumatura, Kupang, Timor, Indonesia and P. 
kitangladensis from Mt. Kitanglad Range, Mindanao Island, Philippines. 

Pheretima castilloi James & Hong sp. n. Figs 1E-F 

Material: Holotype: One clitellate (NMA 0003743): Philippines, Luzon Island, 
Pampanga Province, Mt. Arayat (15°12.43'N, 120°44.38'E), 900 m, litter layers in forest, 13 
March 2001, Y. Hong & A. Castillo colls. 3 paratypes: 1 clitellate (NMA 0003748), 1 clitellate 
(FMNH 10014), 1 clitellate (MHNG 34809): Same data as for holotype. Other material: Mt. 
Arayat (15°12.41'N, 120°44.45'E), 1050 m, litter layers in forest, 4 clitellate, 1 aclitellate, 13 
March 2001. 



PHERETIMA AND PITHEMERA FROM MT. ARAYAT, PHILIPPINES 7 

Etymology. The species is named after the type's collector, Augusto Castillo. 

Diagnosis: Spermathecal pores in 5/6-7/8, genital marking absent. 

Description: Brownish dorsal pigment. Dimensions 59-81 mm by 3.1-3.3 mm 
at segment x, 3.2-3.3 mm at xxx, 3.0-3.3 mm at clitellum, segments 64-81 body cylin- 
drical in cross-section. Setae regularly distributed around segmental equators, number- 
ing 26-28 at vii, 46-50 at xx; 7-12 between male pores, size, distance regular; setal for- 
mula AA:AB:YZ:ZZ = 1:1.5:2:6 at xiii. Clitellum annular xiv-xvi; setae invisible 
externally. 

First dorsal pore 12/13, paired spermathecal pores in 5/6-7/8, at lateral margin 
of ventrum, distance between spermathecal pores 3.0 mm. Female pore single in xiv, 
0.4 mm oval shape, 0.4 mm openings of copulatory bursae paired in xviii, distance 
between male pores 2.5 mm. Genital marking lacking. 

Septa 5/6-7/8 thin, 8/9 absent, 9/10 very thin, attached to 10/11, 10/11 thick, 
11/12-13/14 thin. Gizzard usual in viii-x, intestine begins in xv, small paired lymph 
glands from xxviii along dorsal vessel; intestinal caeca simple, originating in xxvii, and 
extending anteriorly about to xxiv, each consisting of a small finger-shaped sac; 
typhlosole very low simple fold from xxvii. Hearts x-xiii esophageal; ix lateral, very 
small viii to gizzard, vii lateral, extra esophageal vessel pharynx to ventral esophageal, 
then to clitellum area xiii to xiv, xv. 

Ovaries and funnels in xiii, paired spermatheca in vi-viii, viii with nephridia on 
spermathecal ducts; spermatheca with small ampulla pouch, duct shorter than ampulla, 
diverticulum golf club-shaped, shorter than ampulla. Male sexual system holandric, 
testes and funnels in ventral paired sacs in x, xi. Seminal vesicles two pairs in xi, xii 
with dorsal lobes, prostates in xvi- xviii, with stout muscular duct entering center of 
copulatory bursae without stalked glands. Copulatory bursa opening flanked by 
circular pads anterior, posterior, pads backed by glandular chambers, male pore within 
long slit on side of conical protrusion from copulatory bursa roof. 

Remarks: In Sims & Easton (1972) Pheretima castilloi keys to a couplet offer- 
ing a choice between one and four thecal segments, corresponding to the urceolata and 
lejokana (genital marking present), darnleiensis (genital marking absent) species 
groups, respectively. However, the present species has three pairs of spermathecal 
pores in 5/6-7/8. Therefore Pheretima castilloi is unlike all known Pheretima. 

Pithemera rotunda James & Hong sp. n. Figs 2A-B 

Material: Holotype: One duellate (NMA 0003744): Philippines, Luzon Island, 
Pampanga Province, Mt. Arayat (15°12.41'N, 120°44.45'E), 1050 m, litter layers in forest, 13 
March 2001, Y. Hong & A. Castillo colls. 3 paratypes: 1 clitellate (NMA 0003749), 1 clitellate 
(FMNH 10015), 1 clitellate (MHNG 34810): Same data as for holotype. Other material: Same 
data as for holotype, 3 clitellate, 15 aclitellate specimens; Mt. Arayat (15°12.43'N, 
120°44.38'E), 900 m, litter layers in forest, 7 clitellate, 1 aclitellate, 13 March 2001. 

Etymology: The name rotunda is Latin for circular, referring to the shape of the 
genital marking. 

Diagnosis: Spermathecal pores in 4/5-8/9, male pores in xviii, on 0.3 mm small 
hardened slightly protuberant circular porophore. Genital markings numerous circular 
within thickened epidermal area. 



S. W. JAMES ETAL. 




FlG. 2 
A-B. Pithemera rotunda sp. n. A: ventral view; B: spermathecae. C-D. Pithemera philippinensis 
sp. n. C: ventral view; D: spermathecae. Scale bars = 5 mm (C), 2 mm (A), 1 mm (B, D). 



Description: Worm unpigmented. Dimensions 31-44 mm (aclitellates 25-32) by 
2.0-2 .4 mm at segment x, 2.3-2.6 mm at xxx, 1.9-2.0 mm at clitellum, segments 79-91; 
body cylindrical in cross-section. Setae regularly distributed around segmental equa- 
tors, numbering 49-54 at vii, 48-61 at xx; 9 between male pores, size, distance regular; 
setal formula AA: AB: YZ:ZZ = 2:1:1:3 at xiii. Clitellum annular xiv-l/2xvi; xvi setae 
visible, dorsal pore invisible externally. 

First dorsal pore 12/13, five pairs of spermathecal pores in 4/5-8/9, very small 
white spot in lateral margin of ventrum. Female pores paired in xiv, 0.3 mm oval shape, 
male pores in xviii, on 0.3 mm small hardened slightly protuberant circular porophore, 
pores 1.9 mm apart. Circular raised genital markings numerous; 2 on right side xviii, 
3 between male pores xviii, 3 presetal in xix, xx, xxi, xxii; genital markings within 
white thickened epidermal area. 

Septa 5/6 thin, 6/7 thick, 7/8 thin, 8/9 absent, 9/10, 10/11 thin, 11/12, 12/13 
some muscular thickening, 13/14 thin. Gizzard usual in viii-ix, intestine begins in xv; 
intestinal caeca simple, originating in xxii, and extending anteriorly about to xxi, each 
consisting of a small triangle-shaped sac; typhlosole about 1/3 lumen diameter simple 
fold from xxii. Hearts x-xii esophageal; ix lateral, viii absent, vi, vii lateral. 

Ovaries and funnels in xiii, spermathecae in v-ix; no nephridia on spermathecal 
ducts; spermathecae with small strawberry-shaped ampulla, ducts shorter than 
ampulla, diverticulum club-shaped, shorter than ampulla. Male sexual system holan- 



PHERETIMA AND PITHEMERA FROM MT. ARAYAT, PHILIPPINES 9 

dric, testes and funnels in single large sacs in x, xi; sacs attached to front side of septum 
10/11, 11/12, sacs do not enclose hearts, sacs of xi enclose seminal vesicles. Seminal 
vesicles two pairs in xi, xii without dorsal lobes, prostates multilobed in xvi- xx, ectal 
half of duct expanded, muscular. 

Remarks: Pithemera rotunda keys to the bicincta group in Sims & Easton 
(1972), which is composed of two species, Pithemera bicincta (Perrier, 1875) and 
Pithemera violacea (Beddard, 1895). Like P. bicincta and P. violacea, P. rotunda has 
spermathecal pores in 4/5-8/9, and the spermathecal ampulla is longer than the diver- 
ticulum. Pithemera rotunda has three circular papillae in segments xviii to xxii, but P. 
bicincta has paired genital papillae extending from segments xviii to xix, and 1 pair of 
genital papillae just behind the male pores on 18/19. Only one pair of genital papillae 
is present in P. violacea just lateral to and behind the male pores (Beddard, 1895). 

Pithemera philippinensis James & Hong sp. n. Figs 2C-D 

Material: Holotype: One clitellate (NMA 0003745): Philippines, Luzon Island, 
Pampanga Province, Mt. Arayat (15°12.41'N, 120°44.45'E), 1050 m, litter layers in forest, 13 
March 2001, Y. Hong & A. Castillo colls. 2 paratypes: 1 clitellate (FMNH 10016), 1 clitellate 
(MHNG 34811): Same data as for holotype. Other material: Same data as for holotype, 11 
aclitellate specimens; Mt. Arayat (15°12.43'N, 120°44.38'E), 900 m, litter layers in forest, 3 
clitellate, 1 aclitellate, 13 March 2001. 

Etymology: The species is named for its type locality. 

Diagnosis: Spermathecal pores in 4/5-8/9, male pores xviii on foot print- shaped 
pads extending to 17/18-18/19. Genital marking absent. 

Description: Light brown dorsal pigment. Dimensions 51-54 mm (semiclitellate 
41-48) by 2.3-2.5 mm at segment x, 2.8-3.3 mm at xxx, 2.5-2.6 mm at clitellum, 
segments 78-90; body cylindrical in cross-section. Setae regularly distributed around 
segmental equators, numbering 56-68 at vii, 49-61 at xx; 9 between male pores, size, 
distance regular; setal formula AA:AB:YZ:ZZ = 3:1:1:2 at xiii. Clitellum annular xiv- 
l/2xvi; xvi setae visible, dorsal pore invisible externally. 

First dorsal pore 12/13, five pairs of spermathecal pores in 4/5-8/9, small white 
spots, lateral. Female pores paired in xiv, 0.4 mm oval shape, male pores xviii on foot 
print-shaped pads extending to 17/18-18/19, 2.0 mm between male pores. Genital 
marking lacking. 

Septa 5/6-7/8 thin, 8/9 absent, 9/10-11/12 thick, 12/13, 13/14 thin. Gizzard 
usual in viii-ix, intestine begins in xv; intestinal caeca simple, originating in xxii, and 
extending anteriorly about to xxi, each consisting of a small triangle-shaped sac; 
typhlosole simple fold about 1/4 lumen diameter from xxii. Hearts x-xii esophageal; ix 
lateral, viii lateral. 

Ovaries and funnels in xiii, spermathecae in v-ix; no nephridia on spermathecal 
ducts; spermathecae with small pouch, diverticulum sausage-shaped, as long as 
ampulla. Male sexual system holandric, testes and funnels in ventral paired sacs in x, 
xi. Seminal vesicles two pairs in xi, xii, prostates in xvii- xix, two main lobes, ental 
half of ducts slender, non muscular; ectal half of ducts enlarged, muscular tapered at 
both ends; body wall surface over external male pads covered with densely packed, 
tiny tubules, probably nephridia or modified nephridia serving a glandular function. 



10 S. W. JAMES ETAL. 

Remarks: The species also keys to the bicincta group in Sims & Easton (1972), 
but unlike the other members, Pithemera philippinensis has foot print-shaped pads 
extending over 17/18-18/19. 

ACKNOWLEDGEMENTS 

This study was supported by National Science Foundation grant DEB-0072764. 

REFERENCES 

Beddard, F. E. 1895. A monograph of the order Oligochaeta. Oxford, Clarendon Press, i-xii + 

769 pp. 
Cognetti de Martiis, L. 1913. Oligochetes (de Ceram et de Waigeu). Bijdragen tot de 

Dierkunde 19: 37-41. 
Gates, G. E. 1961. On some species of the oriental earthworm genus Pheretima Kinberg, 1867. 

Zoologische Mededelingen, Leiden 37: 293-312. 
James, S. W. 2004. New species of Amynthas, Pheretima, Pleionogaster (Oligochaeta: Mega- 

scolecidae) of the Mt. Kitanglad Range, Mindanao Island, Philippines. Raffles Bulletin 

of Zoology, Singapore. (In press). 
Michaelsen, W. 1924. Oligochäten von Niederländisch-Indien. Treubia, Journal of the Zoo- 
logical Museum and Laboratory 5: 379-401. 
Michaelsen, W. 1928. Die Oligochäten Borneos. Arkiv for Zoologie 20: 1-60. 
Perrjer, E. 1875. Sur les vers de terre des îles Philippines et de la Cochinchine. Comptes Rendus 

Hebdomadaires des Séances de l'Académie des Sciences, Paris 81: 1043- 1046. 
Sims, R. W. & Easton, E. G. 1972. A numerical revision of the earthworm genus Pheretima auct. 

(Megascolecidae: Oligochaeta) with the recognition of new genera and an appendix on 

the earthworms collected by the Royal Society North Borneo Expedition. The Biological 

Journal of the Linnean Society, London 4: 169-268. 



Revue suisse de Zoologie 111 (1): 11-20; mars 2004 



Redescription of Monticellia magna (Rego, dos Santos & Silva, 
1974) (Eucestoda: Monticelliidae) parasite of Pimelodus spp. 
(Pisces: Silurif ormes) from Argentina, and morphological study of 
microtriches 

Alicia A. GIL de PERTIERRA 

Laboratorio de Helmintologia. Dpto. Biodiversidad y Biologìa Experimental. Facultad 
de Ciencias Exactas y Naturales. Universidad de Buenos Aires. Ciudad Universitaria, 
Pab. n, 4° Piso. CP: C1428EHA-Buenos Aires, Argentina. 
E-mail: helminto@bg.fcen.uba.ar 



Redescription of Monticellia magna (Rego, dos Santos & Silva, 1974) 
(Eucestoda: Monticelliidae) parasite of Pimelodus spp. (Pisces: Siluri- 
formes) from Argentina, and morphological study of microtriches. - 

Monticellia magna (Rego, dos Santos & Silva, 1974) is redescribed and the 
microtriches and their distribution are studied for the first time. This species 
is characterised by the following combination of characters: (1) vagina 
anterior to cirrus pouch; (2) muscular asymmetrical sphincter present; (3) 
testes in one layer and in two fields connected anteriorly and posteriorly; (4) 
vitelline follicles distributed cortical, paramuscular and a few follicles 
medullary; (5) internal longitudinal musculature strongly developed; and 
(6) scolex with filiform microtriches in apical region, filiform and spiniform 
microtriches in central cavity, marginal ring and nonadherent surface of 
suckers, and spiniform microtriches in neck and immature proglottides. The 
species parasitised fishes of Pimelodus spp. Strict specificity of South 
American proteocephalids for their hosts is placed into consideration since 
recently new host records have been reported, especially for hosts commer- 
cially exploited. 

Key- words: micro triches - Proteocephalidea - Monticellia magna - 
Pimelodidae - host specificity - Argentina. 

INTRODUCTION 

Monticellia magna (Rego, dos Santos & Silva, 1974) was originally described 
from Pimelodus clarias (Bloch, 1782) (junior synonym) (= P. blochii Valenciennes, 
1840, original combination) as Nomimoscolex magna. However, de Chambrier & 
Vaucher (1997) studied the type material and found it composed by specimens that 
belong to two different genera, Proteocephalus and Monticellia; and transferred N. 
magna to Monticellia. 



Manuscript accepted 21.07.2003 



12 A. A. GIL DE PERTŒRRA 

Latter, Rego & Pavanelli (1992) described Monticellia loyolai from Pimelodus 
maculatus Lacépède, 1803. De Chambrier & Vaucher (1999) synonymized this species 
with M. magna based on the study of the type material and new material collected in 
Paraguay. Eventhough, these authors exposed remarkable and distinctive characters for 
M. magna, and did not present drawings of this species. 

During a survey of proteocephalidean cestodes from freshwater teleost fishes in 
Argentina, specimens of M. magna were collected from Pimelodus albicans (Valen- 
ciennes, 1840), P. argenteus Perugia, 1891, and P. maculatus. M. magna is described 
in detail for the first time in this paper, based on type and the new material. The surface 
of the tegument of the scolex and portions of the strobila were studied using scanning 
electron microscopy (SEM). 

MATERIALAND METHODS 

Eighty specimens off. albicans, 6 of P. argenteus and 212 of P. maculatus from 
Colastiné, La Plata, and Parana rivers were examined for helminths. Worms found in 
the intestine were isolated and fixed in hot 4% v/v formaldehyde solution and stored in 
75% v/v ethanol. Entire tapeworms were stained with Langeron 's alcoholic chlorhy- 
dric carmine (Langeron, 1949), differentiated in acid ethanol, dehydrated through a 
gradual ethanol series, cleared in beechwood creosote and mounted in Canada balsam. 
Thick transverse hand-cutting serial sections of proglottides were stained following the 
same procedure. Eggs were mounted in distilled water, after fixation for drawing. 
Three specimens were prepared for SEM as follows: post-fixed in 1% osmium 
tetroxide, dried with tetrametylsilane (Analyticals', Carlo Erba), mounted on stubs 
with adhesive tape, sputter coated with gold in a Thermo VG Scientific Polaron SC 
7630 and examined with a Philips XL 30 scanning electron microscope. Microthrix 
density values (D) were obtained by counting microtriches from randomly selected 
areas of 1 urn . Voucher specimens of M. magna from Argentina were deposited at 
Colección Parasitológica del Museo Argentino de Ciencias Naturales "Bernardino 
Rivadavia", Buenos Aires, Argentina (MACN-Pa), and at the Natural History Museum, 
Geneva, Switzerland (MHNG). Syntypes of M. magna (Rego, dos Santos & Silva, 
1974) and M. loyolai (Pavanelli & Machado dos Santos, 1992) from Helminthological 
Collection of the Instituto Oswaldo Cruz (CHIOC) were also studied. The information 
on taxonomic classification of fishes was obtained from FishBase Online (www.fish- 
base.org). All measurements are given in micrometers, unless otherwise stated, with 
the range followed by the mean, the standard deviation and the number of measure- 
ments (n) in parentheses. Measurements of microtriches were determined from photo- 
micrographs. Illustrations were made with the aid of a camera lucida using Nomarski 
interference contrast in a Zeiss Axioscope microscope. 

RESULTS 

Monticellia magna (Rego, dos Santos & Silva, 1974) Figs 1-14 

Nomimoscolex magna Rego, dos Santos & Silva, 1974 
Monticellia loyolai Pavanelli & Machado dos Santos, 1992 

Type host: Pimelodus clarias (Bloch, 1782) (junior synonym) (= P. blochii Valenciennes, 
1840, original combination). 



REDESCRIPTION OF MONTICELL1A MAGNA 



13 




Figs 1-2 
Monticellia magna (Rego, dos Santos & Silva, 1974). 1. Scolex, apical region showing unicel- 
lular gland cells. 2. Gravid proglottis, ventral view. Scale-bars: 1 = 250 um; 2 = 500 um. 



Additional hosts: Pimelodus albicans (Valenciennes, 1840), vernacular name: moncho- 
lo, bagre bianco; Pimelodus argenteus Perugia, 1891, vernacular name: bagre bianco; Pimelodus 
maculatus Lacépède, 1803, vernacular name: bagre amarillo (Siluriformes: Pimelodidae). 

Material studied: Argentina: 1) Buenos Aires Province, Buenos Aires Port, La Plata riv- 
er (34°37'S, 58°22'W), MACN-Pa 419/1-3, collected from P. albicans on 23/10/1989, and 
MACN-Pa 423/1-2 collected from P. maculatus on 10/09/1994. 2) Santa Fé Province, Santo 
Tome City, Colastiné river (tributary of Parana river) (31°40'S, 60°46'W), MACN-Pa 423/3-4 
and MNHG 34660 IN VE, collected from P. maculatus on 15/02/2002 and 31/07/2001 respec- 
tively, and MACN-Pa 424/1-2 and MNHG 34661 INVE collected from P. argenteus on 
13/12/2002. Type specimens from Brazil: 1) Mato Grosso State, Esperanza Port, syntypes M. 
magna (Rego, dos Santos & Silva, 1974) CHIOC 31049 a-c, 2 contracted specimens; CHIOC 
33137 (= 4476), fragment of strobila; CHIOC 33139 (= 4480), fragment of strobila. 2) Parana 
State, Parana river, M. loyolai (Pavanelli & Machado dos Santos, 1992), CHIOC 32715 (holo- 
type); CHIOC 32716 c-d and 32717 a-b transverse sections of proglottides (paratypes). 

Prevalence: 40% (80 P. albicans examined), 33% (6 P. argenteus examined), 70% (212 
P. maculatus examined). 



14 



A. A. GIL DE PERTIERRA 




4 




Figs 3-5 
Monticellia magna (Rego, dos Santos & Silva, 1974). 3. Mature proglottis, dorsal view. 4-5. 
Transverse sections of proglottides showing internal longitudinal musculature, and topography 
of the genitalia. 4. Transverse section anterior to ovary. 5. Transverse section at level of ovary. 
Scale-bars: 3-5 = 500 um. 



REDESCRIPTION OF MONTICELLIA MAGNA 



15 




Figs 6-7 
Monticellia magna (Rego, dos Santos & Silva, 1974). 6. Detail of cirrus pouch and vagina show- 
ing the asymmetrical sphincter. 7. Eggs drawn in distilled water, after fixation. Scale-bars: 6 = 
200 urn; 7 = 100 urn. 



Intensity of infection: 2-20 worms per fish. 

Site of infection: anterior and middle part of intestine. 

Description (based on 15 specimens and measurements on 9 specimens from 
Argentina): Proteocephalidea, Monticelliidae, Monticelliinae. Testes, ovary and uterus 
cortical. Vitelline follicles partly in the cortex and partly in the medulla. Medium size 
worms, 23-125 mm, flattened dorsoventrally. Strobila acraspedote, anapolytic, consist- 
ing of 53-125 (n = 9) proglottides: 20-60 immature, 8-22 mature, and 12-60 gravid. 

Scolex wider than proliferation zone (Figs 1, 8), 480-830 (582 ± 58, n = 9) wide. 
Apical organ absent, numerous spherical-shaped glandular cells with granular inclu- 
sions, distributed in apical region. Apical region of scolex proper covered with densely 
packed filiform microtriches 1.1-1.3 (n = 4) long, 0.1 wide, D = 21-25 (n = 4) (Fig. 9). 
Suckers spherical to oval, unilobate, uniloculate, strongly muscular, 220-300 (246, n = 
18) long, 160-300 (217) wide. Central cavity surface of suckers covered with filiform 
microtriches 0.8-1.2 (n = 5) long, 0.1 wide, interspersed with spiniform microtriches 
0.9-1.1 (n = 6) long, 0.2-0.3 wide, D = 23 filiform : 1-2 spiniform (n = 4) (Fig. 11). 
Marginal ring surface of suckers covered with filiform microtriches 0.9-1.2 (n = 5) 
long, 0.1 wide, interspersed with spiniform microtriches 1.0-1.3 (n = 6) long, 0.2-0.3 
wide, D = 14-20 filiform : 1-2 spiniform (n = 4) (Fig. 10). Nonadherent surface of 
suckers covered with filiform 1.0-1.2 (n = 5) long, 0.1 wide, interspersed with spini- 
form microtriches, 1.1-1.3 (n = 4) long, 0.2 wide, D = 8-14 filiform : 2-3 spiniform 
(n = 4) (Fig. 12). Proliferation zone (neck), 800-3000 (1422, n = 9) long, surface 
covered with spiniform microtriches, 0.6-0.8 (n = 4) long, 0.2 wide, D = 31-35 (n = 4) 
(Fig. 13). 



16 



A. A. GIL DE PERTIERRA 




Figs 8-14. Monticellia magna (Rego, dos Santos & Silva, 1974) SEM micrographs. 8. Scolex in 
dorsoventral view, and positions of high magnification views for Figs 9-13. 9. Apical region 
surface, filiform microtriches. 10-12. Suckers: marginal ring, central cavity, and nonadherent 
surface, respectively, filiform microtriches interspersed with spiniform microtriches. 13. Proli- 
feration zone surface, spiniform microtriches. 14. Immature proglottis surface, spiniform 
microtriches. Scale-bars: 8 = 100 um; 9-12, 14 = 1 um; 13 = 2 um. Full arrows show filiform 
microtriches, empty arrows show spiniform microtriches. 



REDESCRIPTION OF MONTICELLI A MAGNA 17 

Immature proglottides wider than long, 120-590 (338, n = 12) long, 430-800 
(598) wide. Immature proglottides surface covered with spiniform microtriches 0.6-0.7 
(n = 5) long, 0.2 wide, D = 38-43 (n = 4) (Fig. 14). The surface of proliferation zone 
and the immature proglottis is covered with spiniform microtriches, and they are the 
regions with higher densities (D = 31-35 and 38-43 respectively). Mature proglottides 
wider than long or longer than wide, 400-1010 (674 ± 154, n = 37) long, 880-2620 
(1340 ± 547) wide (Fig. 3). Gravid proglottides wider than long or longer than wide, 
500-2200 (1150 ± 373, n = 34) long, 870-2740 (1602 ± 621) wide (Fig. 2). 

Internal longitudinal musculature strongly developed, forming thick fibre bun- 
dles, delimiting a reduced medulla. Osmoregulatory canals situated between testes and 
vittelline follicles. Ventral canal, 12-20 in diameter, with secondary osmoregulatory 
canals ending on ventral surface lateral to ovarian lobes. Dorsal canal, 5-10 in diame- 
ter (Figs 4, 5). 

Testes cortical, total number 85-146 (106, n = 22) in mature proglottides, 40- 
100 (70, n = 18) in diameter; in one layer, in two fields connected anteriorly and pos- 
teriorly (Fig. 3). Occasionally, 1-3 testes overlapping vas deferens and distal part of 
cirrus pouch. Cirrus pouch pyriform with thin muscular wall, 210-320 (260 ± 36, n = 
31) long, 40-85 (66 ± 13) wide; occupying 21-29% (24% ± 2, n = 31) of proglottis 
width in mature proglottides. Cirrus occupying about 30-62% of cirrus pouch length. 
Vas deferens coiled, 20-25 in diameter, usually not surpassing mid-line of body in 
mature proglottides. Genital pores irregularly alternating, situated anteriorly at 20-42% 
(28% ± 4, n = 31) of proglottis width. 

Vagina anterior (99%) to cirrus pouch, 13-20 in diameter, strongly asymmetrical 
and muscular sphincter present (Fig. 6). Ovary cortical, with 2 lobulate lobes; occupy- 
ing 56-74% (67% ± 4, n = 31) of proglottis width in mature proglottides. 

Vitelline follicles cortical and paramuscular, with 1-2 follicles lying in medulla. 
Forming 2 lateral bands concentrated in mid-lateral region of proglottis, interrupted at 
cirrus pouch and vagina level on ventral side, reaching 98-100% of total proglottis 
length (Figs 3, 4, 5). 

Uterine primordium stem and uterine branches cortical. Uterine branches 
occupying up to 70% of gravid proglottis width; 19-29 (22, n = 14) lateral branches op- 
posite to cirrus pouch side, and 18-28 (20) on cirrus pouch side. Cortical uterine 
diverticula nearly completely overlap the ovary. Mature eggs released by a ventral 
longitudinal slit (Fig. 2). Eggs with thick hyaline outer envelope, 160-275 (228, n = 7) 
in diameter; embryophore , 35-48 (41, n = 7) in diameter; oncosphere 18-20 (n = 7) in 
diameter; hooks 10-12 long (Fig. 7). 

DISCUSSION 

The genus Monticellia La Rue, 1911 includes 11 species, all distributed in the 
Neotropical region: M. amazonica de Chambrier & Vaucher, 1997, M. belavi stensi s 
Pavanelli, Machado, Takemoto & dos Santos, 1994, M. coryphicephala (Monticelli, 
1891), M. dlouhyi de Chambrier & Vaucher, 1999, M. lenha Woodland, 1933, M. 
magna, M. mandi (Pavanelli & Takemoto, 1996), M. megacephala Woodland, 1934, 
M. ophisterni Scholz, de Chambrier & Salgado-Maldonado, 2001, M. spinulifera, 
Woodland, 1935, and M. ventrei de Chambrier & Vaucher, 1999. M. diesingii 



18 A. A. GIL DE PERTIERRA 

(Monticelli, 1891) and M. macrocotylea (Monticelli, 1892) are considered species 
inquirendae (Rego et al, 1999). 

Among the morpho-anatomical features studied in this redescription the follow- 
ing combination of characters are important to characterise M. magna: (1) vagina an- 
terior to cirrus pouch; (2) muscular asymmetrical sphincter present; (3) testes in one 
layer and in two fields connected anteriorly and posteriorly; (4) vitelline follicles 
distributed cortical, paramuscular and a few follicles medullary; (5) internal longitu- 
dinal musculature strongly developed; and (6) scolex with filiform microtriches in 
apical region, filiform and spiniform microtriches in central cavity, marginal ring and 
nonadherent surface of suckers, and spiniform microtriches in proliferation zone and 
immature proglottides. 

The syntypes of M. magna were studied and the conspecificity with the speci- 
mens from Argentina was confirmed. Transverse sections were not available from the 
material examined. The holotype and the paratypes of Monticellia loyolai (= M. 
magna) were also studied. The asymmetrical vaginal sphincter and uterine branches 
nearly completely overlapping the ovary were clearly observed in type specimens. The 
vitelline follicles are situated cortical, paramuscular, and medullary (Figs 15, 16). The 
same topography was observed in the specimens collected from P. albicans, P. urgen- 
tem and P. maculatus from Argentina, thus the conspecificity with M. magna was also 
confirmed. 

In the genus Monticellia, only M. spinulifera has been partially examined with 
SEM (Rego, 1999). Even when this author studied the sucker at very low magni- 
fication, and the giant spiniform microtriches on the marginal ring of the sucker could 
be easily observed. However, from the photomicrograph it is not known if the spini- 
form microtriches are the only kind of microtriches on the marginal ring surface or if 
they are interspersed with other types of microtriches. Therefore, it is necessary to 
study in detail all the regions of the scolex proper to confirm the microthrix distribu- 
tion of M. spinulifera. To date, in the Neotropical proteocephalids only Nomimoscolex 
semenasae Gil de Pertierra, 2002 (Monticelliidae, Zygobothriinae), and M. magna in 
this paper were completely analysed for the microthrix distribution with SEM. 

Host-specificity varies widely among different taxa of fish helminths. Highly 
host-specific parasites are restricted to one host species and specificity declines as the 
number of suitable host species increases (Poulin, 1998). The South American proteo- 
cephalideans have been considered to be specific to one fish host species. In fact, a few 
examples of species having more than one final host were registered: (1) Amazotaenia 
yvettae de Chambrier, 2001 from Br achyplaty stoma filamentosum Lichtenstein, 1819, 
and B. vaillantii Valenciennes, 1840; (2) Choanoscolex abscissus (Riggenbach, 1896) 
from Zungaro zungaro (Humboldt, 1821) [(= Paulicea luetkeni (Steindachner, 1876)], 
Pseudoplaty 'stoma corruscans (Agassiz, 1829), P. fasciatimi Linnaeus, 1766, and 
Raphiodon vulpinus Spix & Agassiz, 1829 (see Rego et al, 1999); (3) Harriscolex ka- 
parari (Woodland, 1934) from/ 5 , corruscans, and P. tigrinum (Valenciennes, 1840); (4) 
Monticellia magna from Pimelodus albicans (new record), P. argenteus (new record), 
P. clarias, and P. maculatus; (5) M. ventrei de Chambrier & Vaucher, 1999 from 
Luciopimelodus pati (Valenciennes, 1836) (new record), and Pinirampus pirinampu 
(Spix & Agassiz, 1829); (6) Nomimoscolex microacetabula Gil de Pertierra, 1995 from 



REDESCRIPTION OF MONT1CELLIA MAGNA 



19 




Figs 15-16 
Monticellia magna (Rego, dos Santos & Silva, 1974), transverse sections of proglottides drawn 
from holotype CHIOC 32715a, showing internal longitudinal musculature, and topography of 
the genitalia. 15. Transverse section anterior to ovary. 16. Transverse section at level of ovary. 
Scale-bar: 15, 16 = 500 urn. 

P. albicans, and P. maculatus; (7) N. suspectus Zehnder, de Chambrier, Vaucher & 
Mariaux, 2000 from B. filamentosum, B. vaillantii, and Z. zungaro [(= B. flavicans 
(Castelnau, 1885)]; (8) Nupelia tornasi de Chambrier & Vaucher, 1999 from 
Trachelyopterus galeatus (-Parauchenipterus galeatus) (Linnaeus, 1766), and T. cf. 
striatulus (= P. striatulus) (Steindachner, 1877); (9) Peltidocotyle rugosa Diesing, 
1 850 (see Zehnder & de Chambrier, 2000) from P. corruscans, P. fasciatimi, and P. 
tigrinum; (10) Peltidocotyle lenha (Woodland, 1933) (see Zehnder & de Chambrier, 
2000) from Z. Zungaro, and Sorubimichthys planiceps (Spix & Agassiz, 1829); and 
(11) Proteocephalus microscopicus Woodland, 1935 from Cichla monoculus Spix & 
Agassiz, 1831, and Cichla ocellaris Bloch & Schneider, 1801. 

Poulin (1992, 1997) stated that high host-specificity can be an artefact of 
inadequate sampling, and among species of parasites of freshwater fishes sampling 
effort explains much of the variability in host-specificity. The number of South 
American proteocephalid species known parasitising more than one fish host might be 
correlated with a larger sampling effort. Among the fishes mentioned previously 8 1 % 
are commercially exploited (Cichla, 2 species; Brachyplaty stoma, 2 species; Luciopi- 



20 A. A. GBL DE PERTffiRRA 

melodus pati; Pimelodus, 3 species; Pinirampus pirinampu; Pseudoplaty stoma, 3 
species; Z. zungaro) 

M. magna is widespread within Pimelodus spp., these host species are very 
frequent, and is common food among people living nearby the Argentinian rivers. 

In this study accurate drawings of the scolex, mature and gravid proglottides, 
transverse sections of the proglottides at different levels, detail of the vaginal sphinc- 
ter, eggs are revisited for the first, and the types of microtriches and their distribution 
are presented for the first time, the ranges of measurements and the mean are in agree- 
ment with the values given by de Chambrier & Vaucher (1999), differences were 
registered only for the number of uterine branches (18-36 vs 37-57 in this paper), and 
cirrus length/cirrus pouch length (30% vs 30-62% in this paper). 

ACKNOWLEDGEMENTS 

The author thanks Dr V. Ivanov, at Universidad de Buenos Aires, for fruitful dis- 
cussions during the preparation of this manuscript and its critical revision; the review- 
er for constructive critisism; and Dr Dely Noronha, at Instituto Oswaldo Cruz, for the 
loan of specimens. The research was supported by SEPCYT Grant BID 1201 OC- AR 
PICTN° 1-6604. 

REFERENCES 

de Chambrier, A. & Vaucher, C. 1997. Révision des cestodes (Monticelliidae) décrits par 
Woodland (1934) chez Br achy platy stoma filamentosum avec redéfinition des genres 
Endorchis Woodland, 1934 et Nomimoscolex Woodland, 1934. Systematic Parasitology 
37: 219-233. 

de Chambrier, A. & Vaucher, C. 1999. Proteocephalidae and Monticelliidae (Eucestoda: 
Proteocephalidea) parasites de poissons d'eau douce au Paraguay, avec descriptions d'un 
genre nouveau et de dix espèces nouvelles. Revue suisse de Zoologie 106: 165-240. 

Langeron, M. 1949. Précis de Microscopic 7 th ed. Paris: Masson & Cie, 1429 pp. 

Rego, A. A. 1999. Scolex morphology of proteocephalid cestodes parasites of Neotropical fresh- 
water fishes. Memórias do Instituto Oswaldo Cruz 94: 37-52. 

Rego, A. A., Chubb, J. C. & Pavanelli, G. C. 1999. Cestodes in South American freshwater 
teleost fishes: keys to genera and brief description of species. Revista Brasileira de 
Zoologia 16: 299-367. 

Pavanelli, G. C. & Machado dos Santos, M. H. 1992. Goezeella agostinhoi sp. n. e Monti- 
cellia loyolai sp. n., cestóides proteocefalideos parasitas de peixes pimelodideos do rio 
Parana, Parana, Brasil. Revista Brasileira de Parasitologia Veterinaria 1: 45-50. 

Poulin, R. 1992. Determinants of host-specificity in parasites of freshwater fishes. International 
Journal for Parasitology 22: 753-758. 

Poulin, R. 1997. Parasite faunas of freshwater fish: the relationship between richness and the 
specificity of parasites. International Journal for Parasitology 27: 1091-1098. 

Poulin, R. 1998. Evolutionary ecology of parasites: from individuals to communities. Chapman 
& Hall, 212 pp. 



Revue suisse de Zoologie 111 (1): 21-30; mars 2004 



Austrolebias arachan (Cyprinodontiformes, Rivulidae), a new 
species of annual fish from northeastern Uruguay 

Marcelo LOUREIROi, Maria de las Mercedes AZPELICUETA2 & 
Gracida GARCÎA3 

1 Section Vertebrados, Fac. de Ciencias, Iguâ 4225, Montevideo 11400, Uruguay. 

2 Division Zoologìa Vertebrados, Facultad de Ciencias Naturales y Museo de 
La Plata, Paseo del Bosque, 1900 La Piata, Argentina. 

3 Sección Genètica Evolutiva, Fac. de Ciencias, Iguâ 4225, Montevideo 11400, 
Uruguay. 

Austrolebias arachan (Cyprinodontiformes, Rivulidae), a new species 
of annual fish from northeastern Uruguay. - The annual fish Austrolebias 
arachan is described from temporary ponds of Rio Tacuari (Los Patos- 
Merin system) and Rio Negro (Rio Uruguay drainage) basins. Austrolebias 
arachan is easily distinguished from other Austrolebias species by the com- 
bination of the following characters: body of males dark brown to black, 
with light narrow yellow vertical bands, dorsal fin dark brown with verti- 
cally elongated light yellow dots in the proximal half, pectoral fin light grey 
with margin black; females with anterior anal fin rays elongated, forming a 
triangular- shaped fin. Both sexes with pelvic fins joined to each other in dif- 
ferent degrees; two to four parietal neuromasts; three pectoral radiais in 
most individuals. The new species is closely related to A. adloffi species 
group and to A. viarius. 

Key-words: Cyprinodontiformes - Rivulidae - Austrolebias - Uruguay 
basin - Los Patos System. 

INTRODUCTION 

At least seven species of the annual fish genus Austrolebias (Rivulidae) are en- 
demic of the Los Patos-Merin system, southern Brazil and eastern Uruguay (Costa & 
Cheffe, 2001; Garcia et al., 2000). Most of them are distributed in lowlands and 
wetlands associated to that drainage (Costa & Cheffe, 2001; Azpelicueta & Garcia, 
2001), between and 50 meters above sea level. However, at least two species, A. 
cyaenus (Amato, 1987) and A. vazferreirai (Berkenkamp et al., 1994), inhabit "high- 
land" localities of that system, between 50 and 100 meters above sea level. Further- 
more, they can also be found in temporary ponds of the Rio Negro drainage, Uruguay 
(Berkenkamp et al, 1994; Azpelicueta & Garcia, 2001). 

In the present article, we describe a new species of Austrolebias with a disrupt- 
ed distribution that includes "highland" localities from both drainages, Laguna Merin 
and Rio Negro (Rio Uruguay basin). We also perform chromosomal studies to con- 
tribute with the diagnosis of the populations under study. 



Manuscript accepted 17.10.2003 



22 M. LOUREIRO ETAL. 



MATERIAL AND METHODS 

Measurements were taken with digital calliper following Costa (1988), except 
peduncle length that was measured from the vertical through last anal fin ray to pos- 
terior margin of hypural plate and snout length which was measured from anterior tip 
of upper lip to a vertical through anterior eye margin. 

Measurements were expressed as percentages of SL, or other lengths as indi- 
cated. Vertebral counts included the composed caudal centrum as one element. 
Cephalic neuromasts nomenclature followed Costa (2001). Osteological preparations 
were made following Dingerkus & Uhler (1977). 

In the cytogenetics analysis, metaphases were studied following the protocols 
of Kligerman & Bloom (1977), with modifications by Bertollo et al. (1978). Better 
metaphase preparations of spleen, kidney, and gill epithelium were selected under a 
microscope Nikon Microphot-Fx with inmersion objective. Photographs were taken 
with Kodalith (25 asas). Chromosome nomenclature followed Levan et al. (1964) and 
Denton (1973) for fishes. Based on arm relation (AR), chromosomes were classified as 
metacentric (M), submetacentric (S), subtelocentric (ST) and acrocentric (A). The 
diploid number (2n) and the number of chromosome arms (FN) were also established. 
Nucleolar Organizer Regions (NORs) were identified with silver-staining method by 
Howell & Black (1980) and constitutive heterochromatic C-band regions were 
detected according to Sumner (1972). Meiotic studies were performed in male indi- 
viduals to corroborate haploid number (N) following standard protocols (Kligerman & 
Bloom, 1977). 

The examined specimens were deposited in Muséum d'histoire naturelle de 
Genève (MHNG) and Vertebrate Collection of Facultad de Ciencias, Montevideo, 
Uruguay (ZVC-P). Tissues and voucher of A. arachan used for chromosomic analysis 
are deposited in the Sección Genètica Evolutiva Facultad de Ciencias, Montevideo, 
Uruguay (FCMVD-GP) with the numbers 657-661, 775-781, and 1130-1132 (all from 
Departamento de Cerro Largo, Parque Municipal, Melo city). 

Comparative material (SL in mm). Austwlebias adloffi Ahi: ZVC-P 724, 25 ex., (8 
C&S), 30.75-38.26 mm, Brazil, Rio Grande do Sul, Niteroi, pond into the town; ZVC-P 747, 25 
ex., 26.46-29.55 mm, Brazil. Rio Grande do Sul. pond close to Rio Gravataf; ZVC-P 854, 16 ex., 
26.00-35.10 mm, Brazil, Rio Grande do Sul, Niteroi, pond close to Rio Gravataf; ZVC-P 855, 
13 ex., 27.85-33.70 mm, Brazil, Rio Grande do Sul, Niteroi, Villa Rio Branco; MCP 10288, 12 
ex., 38.95-54.29 mm, Brazil, Rio Grande do Sul, Canoas, Rio dos Sinos; MCP 15040, 2 ex., 
32.00-39.00 mm, Brazil, Rio Grande do Sul, Rio Gravataf; MCP 10933, 20 ex., 29.37-36.92 mm, 
Brazil, Rio Grande do Sul, Alvorada. rio, Rio Gravataf; UMMZ 168844, 4 ex., 37.25-50.75 mm, 
Brazil, Canoas; UMMZ 181681, 13 ex., 37.50-44.55 mm, Brazil, vicinity of Porto Alegre; 
UMMZ 181682, 11 ex., 32.60-40.70 mm, Brazil, vicinity of Porto Alegre. Austwlebias bellot- 
tii Steindachner: ZVC-P 876, 61 ex. (19 C&S), 25.20- 46.93 mm, Uruguay, Colonia, Carmelo, 
pond close to Rio Uruguay. Austwlebias melanoorus Amato: ZVC-P 4322, 10 ex. (2 C&S), 
29.82-38.00 mm, ZVC-P 4323. 10 ex., 26.74-44.64 mm, Uruguay, Tacuarembó, Pueblo Ansina, 
pond close to Rio Tacuarembó 10 ex. (2 C&S). Austwlebias vandenbergi Huber: ANSP 175282, 
16 ex., 28.73-47.71 mm, Paraguay, Boquerón, pond along road Filadelfia-Teniente Montana; 
ANSP 175289, 20 ex., 37.30-55.10 mm, Paraguay, Boquerón, pond along road Filadelfia- 
Teniente Montana; ANSP 175290, 25 ex., 24.88-47.44 mm, Paraguay, Boquerón, pond along 
road Filadelfia-Teniente Montana. Austwlebias viarius Vaz-Ferreira. Sierra & Paulette: ZVC-P 
525, 50 ex. (8 C&S), 28.63-49.63 mm, Uruguay, Rocha, pond in the vicinity of A° Valizas; ZVC- 
P 596, 14 ex. (2 C&S), 27.57-41.67 mm, Uruguay, Rocha, pond next to A° Valizas. 



A NEW AUSTROLEBIAS FROM URUGUAY 23 



RESULTS 

Austrolebias arachan sp. n. Figs 1, 2, Tables I, II 

Cynolebias uruguayensis Nion et ah, 1994, nomen nudum. 

Holotype. ZVC-P 4336, male, 38.6 mm SL, Departamento de Cerro Largo, pond close 
to Route 7 and Arroyo Chuy, Laguna Merin basin, Uruguay, col. M. Loureiro, F. Teixeira, A. 
D'Anatro, L. Bocardi, September 2000. 

Paratypes. ZVC-P 4574, 6 males (1 C&S), 31.9-43.7 mm SL, 8 females (1 C&S), 24.8- 
31.3 mm SL, collected with the holotype. ZVC-P 4314, 2 males, 22.9-24.9 mm SL, 1 female 
21.6 mm SL, Departamento de Cerro Largo, pond close to Melo city, road to the airport, 
Uruguay, coll. M. Loureiro, F. Teixeira, A. D'Anatro, L. Bocardi, September 2000. ZVC-P 43 17, 
4 males (2 C&S), 26.0-29.5 mm SL, 4 females (2 C&S), 23.2-28.3 mm SL, Departamento Cerro 
Largo, pond at Parque Municipal, Melo city, Uruguay, coll. M. Loureiro, F Teixeira, A. 
D'Anatro, L. Bocardi, September 2000. ZVC-P 4329, 4 males (all C&S), Departamento de Cerro 
Largo, pond close to Route 26 and Negro River, Uruguay, coll. M. Loureiro, G. Yemini, C. 
Hernandez, October 1999. ZVC-P 4331, 14 males (3 C&S), 26.9-33.8 mm SL, 7 females, 24.7- 
33.0 mm SL, Departamento de Cerro Largo, pond close to Route 26 and Rio Negro, Uruguay, 
coll. M. Loureiro, F Teixeira, A. D'Anatro, L. Bocardi, September 2000. ZVC-P 4332, 10 males, 
23.6-28.2 mm SL, 5 females, 16.4-22.7 mm SL, Departamento de Cerro Largo pond close to 
Melo city, road to the airport, Uruguay, coll. M. Loureiro, F Teixeira, A. D'Anatro, L. Bocardi, 
September 2000. ZVC-P 4333, 5 males (3 C&S) 28.5-35.6 mm SL, 14 females (5 C&S), 24.5- 
35.6 mm SL, Departamento de Cerro Largo pond close to Melo city, road to the airport, Uruguay, 
coll. M. Loureiro, G. Yemini, C. Hernandez, October 1999. ZVC-P 4335, 2 males, 30.95-32.45 
mm SL, 4 females, 26.3-31.9 mm SL, Departamento de Tacuarembó, Route 26, km 331 pond 
next to the road, Uruguay, coll. M. Loureiro, F Teixeira, A. D'Anatro, L. Bocardi, September 
2000. ZVC-P 4528, 3 males (1 C&S), 25.0-29.4 mm SL, 3 females (1 C&S), 23.6-25.9 mm SL, 
Departamento de Cerro Largo, a pond at Parque Municipal, Melo city, Uruguay, coll. G. Garcia, 
July 2000. MHNG 2641.88, 3 males, 30.58-33.89 mm SL, 3 females, 24.53-29.38 mm SL, 
Departamento de Cerro Largo, pond close to Route 26 and Rio Negro, Uruguay, coll. M. 
Loureiro, F Teixeira, A. D'Anatro, L. Bocardi, September 2000. 

Diagnosis. Austrolebias arachan is distinguished by the following combination 
of characters: medium size body (no more than 45.8 mm SL); dorsal-fin insertion 
anterior to anal-fin origin; long dorsal- and anal-fin bases; male body dark brown to 
black with light yellow narrow vertical bands, dorsal fin dark brown with vertically 
elongated light yellow dots on basal half, yellowish brown without dots on distal half; 
proximal half of anal fin dark brown with vertically elongated light blue dots, distal 
half light blue without dots; caudal fin light blue with light yellow dots proximally, 
hyaline on distal fourth; pectoral fin light grey with the ventral margin black. Males 
with contact organs on scales of flanks, specially distributed over anal fin and around 
pectoral-, pelvic-, and caudal-fin bases; contact organs on distal third of second to fifth 
anal fin rays and on three or four uppermost pectoral fin rays distally. Anal fin of 
females with anterior rays elongated, forming a triangular- shaped fin. Pelvic fins joined 
to each other at their bases or by an epithelial membrane covering completely or 
partially (one third) of inner edge in both fins. Two to four parietal neuromasts, some- 
times continuous with supra-orbital series. Three pectoral radiais in 92 % of indi- 
viduals examined, four radiais in the remaining 8 %. 

Description. Meristic and morphometric values are presented in tables I and II 
respectively. Dorsal profile of head straight or slightly concave; dorsal profile strongly 
convex from rear of head to dorsal-fin origin, less convex in females; straight below 
dorsal fin, and straight or scarcely concave on caudal peduncle. Ventral profile arched 



24 M. LOUREIRO ETAL. 

from tip of lower jaw to end of anal-fin. Maximum body depth at pelvic-fin origin 
(Fig. 1); females with lower body. 

Dorsal fin with 21-26 rays in males; in females, 16-20. Dorsal-fin tip of males 
rounded, acute in females. Second proximal pterygophore of dorsal fin through a ver- 
tical between vertebrae 7 and 8 in males; through vertebrae 9 and 11 in females. Anal 
fin with 22-27 rays in males, 17-22 in females. Anal-fin tip rounded in males; anterior 
rays elongated, forming a triangular fin in females, ratio between longest anal-fin 
ray/anal-fin base 0.76-1.21 (mean 0.94). Anal-fin origin through a vertical between 2 nd 
to 5 th dorsal-fin rays in males and females. Caudal fin with distal margin slightly 
rounded in males, with 20-29 rays; caudal fin rounded with 20-26 rays in females. 
Caudal-fin skeleton supported by last three or four vertebrae. Pectoral fin with 11-13 
rays in males, 10-14 in females; pectoral-fin margin rounded, with middle rays longer; 
tip of pectoral fin reaching bases of 2 nd to 6 th anal-fin rays in males, and half or two 
thirds of pelvic-fin length of females. Pelvic fin with 4-7 rays, covering urogenital 
papilla and surpassing 3 to 5 anal-fin ray bases in males, scarcely surpassing anal-fin 
origin in females; distal fin margin rounded. Pelvic fins joined to each other at their 
bases or joined by an epithelial membrane covering completely or partially (one third) 
of inner edge in both fins. 

Total number of vertebrae 27-30, number of precaudal vertebrae 11-13; second 
vertebrae pre-zygapophysis poorly developed; post-temporal ventral process variably 
developed, process reduced or absent in 47 % of the individuals, in rest process mod- 
erate or well developed. 

Scales cycloid, 27-34 on lateral series; transverse scales 11-18; scales around 
caudal peduncle 14-20; 1-3 series over caudal fin. Suborbital and postorbital regions 
scaled. 

Contact organs on scales placed over anal-fin base, many contact organs dis- 
tributed around pectoral and pelvic fin bases, peduncular area completely covered by 
contact organs. Contact organs on distal half of uppermost pectoral fin rays 1-3 and on 
distal third of anal fin rays 2-6. 

Supraorbital neuromasts 13-22, parietal neuromasts 2-4. Ventral gill rakers 
7-11, dorsal gill rakers 2-4. Basihyal slightly widened, basihyal cartilage 50-60 % of 
total basihyal length. Two to 4 teeth in second pharyngobranchial. Fourth cerato- 
branchial with 1 or 2 series of teeth mono-, bi- or multicuspidate. Dermosphenotic 
absent. 

Coloration in life: Male: Body dark brown with transverse well defined narrow 
light yellow bands, between those bands may appear some poorly defined bands with 
the same coloration. Opercular region bluish. Suborbital and supraorbital bands black. 
Dorsal fin dark brown with vertically elongated light yellow dots in the proximal half, 
yellowish brown without dots in the distal half; proximal half of anal fin dark brown 
with vertically elongated light blue dots, distal half light blue without dots; caudal fin 
light blue with light yellow dots proximally, hyaline in distal fourth; pectoral fin grey 
with lower margin black; pelvic fin bluish. 

Female: Pale brown body, with darker brown irregular dots or stripes on flanks. 
Dorsum darker than rest of body. Opercular region yellowish green iridescent. Sub- 
orbital band faint, supraorbital absent. Dorsal and anal fins proximally covered by 



A NEW AUSTROLEBIAS FROM URUGUAY 



25 




Fig. 1 
Austrolebias arachan, female and male, non-preserved specimens collected in a pond at Parque 
Municipal, Melo city, Uruguay (one of the paratypes localities). 



ST-Â 



r it it 



i< 



15 



1< 



10 



11 



Fig. 2 
Karyogram of Austrolebias arachan, somatic cells stained with Giemsa. Diploid number 2n= 48, 
NF= 60. Scale= 10 mm. 



irregular black to brown dots or diffuse vertical bands or completely hyaline; most 
specimens with dots on basal one third of caudal fin; a few specimens, more or less one 
third of them, without dots on caudal fin. Pectoral fin hyaline; pelvic fin light grey. 

Cytogenetic analysis. The mitotic metaphase studies from 15 individuals (ten 
males and five females) showed 2n=48, NF=60 and gonial meiotic cells from testis 
N=24. All individuals with following chromosome formula: three pairs of biarmed 
chromosomes of submetacentric (SM) type, and 23 pairs of ST-A chromosomes (Fig. 
2); three pairs of large acrocentric chromosomes; two Ag-NORs placed at terminal 
position in small arms of first SM chromosome pair (see arrows, Fig. 2). C-bands lo- 



26 



M. LOUREIRO ETAL. 



Table I. Meristic data of Austrolebias arachan from Laguna Merin and Rio Negro basins. 
Holotype and minimum, maximum, and mean of 90 paratypes. 







Males (n= 


= 45) 


Females (n= 


= 45) 




Holotype 


min-max 


mean 


min-max 


mean 


Dorsal fin rays 


22 


21-26 


23.6 


16-20 


18.0 


Anal fin rays 


23 


22-27 


24.1 


17-22 


19.8 


Caudal fin rays 


23 


20-29 


23.4 


20-26 


23.3 


Pectoral fin rays 


13 


11-13 


12.1 


10-14 


12.1 


Pelvic fin rays 


6 


4-6 


5.4 


5-7 


5.6 


Lateral scales 


31 


27-34 


30.4 


28-34 


30.5 


Transverse scales 


14 


11-18 


14.5 


12-16 


13.3 


Peduncle scales 


18 


14-20 


16.8 


14-18 


16.2 


Predorsal scales 


19 


14-24 


19.9 


19-30 


22.0 


Supraorbital scales 


4 


0-4 


1.8 


0-4 


20.7 


Supraorbital neuromast series 


21 


13-24 


20.7 


15-27 


2.0 



Table II. Morphometries of Austrolebias arachan from Laguna Merin and Rio Negro basins. 
Holotype and maximum, minimum, and mean of 86 paratypes in percentage of SL. 



Males (n= 39) 



Females (n= 47) 



Holotype 



Standard length 
Predorsal length 
Basidorsal length 
Preanal length 
Basianal length 
Prepelvic length 
Body depth 
Peduncle depth 
Peduncle length 
Pectoral fin length 
Pelvic fin length 
Head length 
Head width 
Head depth 
Eye diameter 
Interorbital length 
Snout length 



38.6 


22.9-45.8 


30.2 


16.4-42.4 


26.7 


49.9 


47.5-54.1 


50.3 


51.8-62.0 


58.4 


43.3 


38.3-48.1 


43.2 


25.9-35.7 


29.4 


51.2 


49.3-59.0 


53.2 


55.2-65.9 


60.2 


38.2 


33.8-48.0 


41.4 


21.5-31.0 


24.9 


49.6 


43.5-49.6 


46.9 


46.9-57.8 


51.8 


39.0 


30.6-42.1 


35.2 


25.6-37.1 


31.0 


13.6 


11.2-15.9 


13.5 


9.4-14.2 


12.3 


12.7 


7.7-15.3 


10.4 


14.9-22.6 


18.6 


23.9 


10.2-27.9 


23.6 


20.9-30.1 


25.4 


9.1 


7.9-13.3 


9.8 


9.4-14.0 


11.7 


34.2 


28.9-34.5 


32.2 


28.4-36.9 


32.4 


59.4 


49.7-65.7 


57.3 


50.2-70.5 


59.0 


93.9 


79.5-106.1 


88.7 


71.9-98.7 


81.5 


24.1 


23.8-32.9 


27.8 


23.9-37.9 


28.8 


41.7 


37.7-50.4 


44.3 


35.0-48.7 


41.9 


19.7 


15.9-24.3 


19.9 


11.6-23.3 


19.1 



cated at centromeric and telomeric positions in some chromosome pairs. Large hetero- 
chromatic blocks of C-bands occuring at interstitial regions of extra large ST-A type. 

Etymology. The name arachan, originally refers to native inhabitants from the 
vicinity of Porto Alegre city and Laguna de los Patos system (Brazil). At present, it 
refers to people from Departamento de Cerro Largo (Uruguay), where several popul- 
ations of the new species of Austrolebias occur. 

Distribution. The new species is known from localities of the upper Rio Tacuari 
drainage (Laguna Merin basin), and upper Rio Negro drainage (Rio Uruguay basin); 
all localities are between 50 and 100 meters above sea level (Fig. 3). 



A NEW AUSTROLEBIAS FROM URUGUAY 



27 




Fig. 3 
Geographical distribution of Austrolebias arachan. Black star: type locality; white stars: 
paratypes localities. Dark grey areas represent Cucinila Grande (more than 150 m above sea lev- 
el), white areas represent land between 150 and 100 m above sea level, ligth grey areas repre- 
sent land between 100 and 50 m above sea level. Scale bar= 10 km. 



DISCUSSION 

Nion et al. (1994), in a meeting occurred in Montevideo (Uruguay), presented 
an abstract entitled "Dos nuevas especies del gènero Cynolebias Steindachner, 1876 
(Actinopterygii, Aplocheilidae) del Uruguay". Those authors recognized two new 
species but they never described them, neither in the abstract nor in other subsequent 
paper. Therefore, the name Cynolebias uruguayensis, used in that abstract for a taxon 
with similar distribution of the new species here described, became a nomen nudum. 
Furthermore, the material used for that record is lost and this taxon was never de- 
scribed until the present paper. 

Austrolebias arachan is related to a group of medium size species that present 
the anterior anal-fin rays of females elongated (Costa, 2002), pelvic fins joined to each 
other in different degrees, and two to four parietal neuro masts. The species included in 
this group are: A. bellottii, A. melanoorus, A. vandenbergi, A. viarius, and the A. adlof- 
fi species group, in which A. adloffi, A. charrua, A. minuano, A. nigrofasciatus are in- 
cluded (Costa & Cheffe, 2001; Costa, 2002). Austrolebias arachan also shares with all 
the species mentioned, except A. viarius, the presence of pectoral fin with ventral mar- 
gin black. The pigmentation pattern in the body male of A. arachan easily distinguishes 
it from most of the species aforementioned, except from A. viarius. However, A. 
viarius is distinguished from males of A. arachan by the darker coloration of body and 
fins (body dark brown to black vs. brown orange, Vaz Ferreira et al., 1964), a relatively 
larger dorsal fin base (mean 43.2 vs. 38.8 % of SL), and a relatively narrower head 
(mean 57.3 vs. 62.2 % of HL); both sexes of A. arachan and A. viarius are also dif- 
ferentiated by the absence in the former of black dots on the flanks. Austrolebias 
arachan is further distinguished from A. adloffi species group by the absence of verti- 
cally arranged black dots in the caudal peduncle. Furthermore, the proportion of indi- 
viduals with reduced postemporal process is higher in A. arachan (45 %) than in all 
species abovementioned (less than 30 %). 



28 M. LOUREIRO ETAL. 

Parsimony analysis based on 27 data set of cyt-b sequence including eighteen 
species of Austrolebias and six other Rivulidae genera (Garcia et al., 2002) confirmed 
that A. arachan (C. sp GG-661, in that article) is closely related to the A. adloffi 
species group, and to A. viarius. Also, cytogenetic data revealed that A. arachan 
presents a generalized karyotype composed by 48 chromosomes of predominantly ST- 
A chromosome type as was reported for other Cyprinodontiforms groups (Ebeling & 
Chen, 1970) and other taxa within Austrolebias (Garcia et al, 1993, 1995, 2002). 
However, A. arachan has distinctive karyological characters from those of A. adloffi, 
and A. viarius. Austrolebias arachan has three pairs of SM chromosome type, while A. 
adloffi bears one pair of biarmed chromosomes and A. viarius one or two pairs. 
Furthermore, A. arachan presents three extra-large ST-A chromosomes as markers. 
This type of chromosomes were found in species included in other clades of Austro- 
lebias such as A. alexandri and A. gymnoventris but they were not found in A. viarius 
and A. adloffi (Garcia et al., 1995). 

Austrolebias arachan shows two NORs at terminal positions of the first biarmed 
chromosomes. The variability in number and position of NOR regions observed in this 
taxon is very low in comparison with the high variability detected in previous studies 
within the genus Austrolebias and among sister taxa of the clade (Garcia et al., 1993, 
1995, 2001). Peculiar large heterochromatic blocks of C-bands were detected at inter- 
stitial regions of the extra large ST-A type; this is an additional different cytogenetic 
character in A. arachan, not found in A. viarius and A. adloffi. 

Austrolebias arachan, together with A. vazferreirai, has a unique geographical 
distribution within the genus, because no other species is found in localities of the two 
different drainages of the Rio Negro and the Laguna Merin. In addition, both drainages 
belong to different ichthyogeographical regions such as the Rio de la Plata basin and the 
Coastal Atlantic rivers (Ringuelet, 1975). The distribution pattern of the new species 
could be an indicator of an ancient connection between both systems and regions. The 
Rio Negro basin is separated from the Laguna Merin basin by the Cuchilla Grande (up 
to 300 m above sea level). However, this hills system is interrupted by an area between 
150 and 100 meters above sea level, close to A. arachan distribution (Fig. 3). This dis- 
tribution pattern would suggest a relatively ancient origin of these species and also the 
possibility of dispersion caused by birds (Santamaria & Klassen, 2002). Annual fish 
present drought resistant eggs which could rise the probability of successful dispersion 
by this way. However, this kind of phenomena has not been proved for these fish yet. 

ACKNOWLEDGEMENTS 

Authors thank the late Juan Reichert for the aquarium pictures of the new 
species, D. Nelson and M. Sabaj for the loan of comparative material from UMMZ and 
ANSP, respectively, and R. Reis for the allowance to check comparative material from 
MCP This work was partially funded by PEDECIBA (UdelaR, Uruguay). 

REFERENCES 

Amato, L. H. 1987. Description de Cynolebias cyaneus n. sp., nuevo pez anual del estado de 
Rio Grande do Sul, Brasil (Cyprinodontiformes, Rivulidae). Comunicaciones 
Zoológicas del Museo de Historia Natural de Montevideo 163: 2-11. 



A NEW AUSTROLEBIAS FROM URUGUAY 29 



Azpelicueta, M. de las M. & Garcia, G. 2001. The fish fauna of a reserve of Biosphere, The 
"Humedales del Este", in Uruguay. Biogeographica 11: 1-13. 

Berkenkamp, H. O., Etzel, V., Reichert, J. J. & Salvia, H. 1994. Ein neuer Fächerfisch aus 
Uruguay. Cynolebias vazferreirai sp. n. Das Aquarium 306: 11-19. 

Bertollo, L. A. C., Takahashi, C. S & Moreira-Filho, O. 1978. Karyotipic studies of two al- 
lopatric populations of the genus Hoplias (Pisces, Erythrinidae). Revista Brasileira de 
Genetica 2: 17- 37. 

Costa, W. J. E. M. 1988. Sistematica y distribuiçâo do complexo de espécies Cynolebias min- 
imus (Cyprinodontif ormes, Rivulidae), com a descriçào de duas espécies novas. Revista 
Brasileira de Zoologia 5: 557-570. 

Costa, W. J. E. M. 2001. The Neotropical annual fish genus Cynolebias (Cyprinodontiformes: 
Rivulidae): phylogenetic relationships, taxonomic revision, and biogeography. Ichthyo- 
logical Exploration of Freshwaters 12: 333-383. 

Costa, W. J. E. M. 2002. Monophyly and phylogenetic relationships of the Neotropical annual 
fish genera Austrolebias, and Megalebias (Cyprinodontiformes: Rivulidae). Copeia 2002 
(4): 916-927. 

Costa, W. J. E. M. 2001 & Cheffe, M. M. 2001. Three new annual fishes of the genus 
Austrolebias from the Laguna Dos Patos System, southern Brazil, and a redescription of 
A. adloffi (Ahi) (Cyprinodontiformes: Rivulidae). Comunicaçôes do Museu de Ciências 
e Tecnologia da PUCRS, série Zoologia, Porto Alegre 14: 179-200. 

Denton, T. E. 1973. The Fish Karyotype (pp. 69-86). In: Thomas, C. E. (ed.). Fish chromosome 
methodology. Springfield, Illinois. 

Dingerkus, G. & Uhler, L. D. 1977. Differential staining of bone and cartilage in cleared and 
stained fish using alcian blue to stain cartilage and enzymes for clearing flesh. Stain 
Technology 52: 229-232. 

Ebeling, A. W. & Chen, T. R. 1970. Heterogamety in teleostean fishes. Transactions of the 
American Fish Society 99: 131-138. 

Garcia, G., Scvortzoff, E., Màspoli, M. C. & Vaz-Ferreira, R. 1993. Analysis of karyotypic 
evolution in natural populations of Cynolebias (Pisces, Cyprinodontiformes, Rivulidae) 
using banding techniques. Cytologia 58: 85-94. 

Garcia, G., Scvortzoff, E. & Hernandez, A. 1995. Karyotipic heterogeinity in South 
American Annual Killifishes of the genus Cynolebias (Pisces, Cyprinodontiformes, 
Rivulidae). Cytologia 60: 103-110. 

Garcia, G., Wlasiuk, G. & Lessa, P. E. 2000. High levels of mitochondrial cytochrome b di- 
vergence in the annual killifishes of the genus Cynolebias (Cyrpinodontiformes, Rivu- 
lidae). Zoological Journal of the Linnean Society 129: 93-110. 

Garcia, G., Lalanne, A. I., Aguirre, G. & Cappetta, M. 2001. Chromosome evolution in the 
annual killifish genus Cynolebias and mitochondrial phylogenetic analysis. Chromosome 
Research 9: 437-448. 

Garcia, G., Alvarez- Valin, F & Gómez, N. 2002. Garcia G, Alvarez- Valin, F & Gomez, N. 
2002. Mitochondrial genes: Signals and Noice in phylogenetic reconstruction within kil- 
lifish genus Cynolebias (Cyprinodontiformes, Rivulidae). Biological Journal of the 
Linnean Society 76: 49-59. 

Howell, W. M. & Black, D. A. 1980. Controlled silver staining nucleolus organizer with pro- 
tective colloidal developer: a 1-step method. Experientia 36: 1014-1015. 

Kligerman, A. D. & Bloom, S. E. 1977. Rapid chromosome preparations from solid tissues of 
fishes. Journal of Fisheries, Research Board of Canada 34: 266-269. 

Levan, A., Fredga, K. & Sandberg, A. A. 1964. Nomenclature for centfomeric position on 
chromosomes. Hereditas 52: 201-220. 

Nión, H., Rios, C, Reichert, J., Salvia, H. & Prieto, F 1994. Dos nuevas espécies del gènero 
Cynolebias Steindachner, 1 876 (Actinopterygii, Aplocheilidae) del Uruguay. Resumenes 
del Primer encuentro nacional sobre fauna acuâtica, octubre de 1994, Montevideo, 
Uruguay. 



30 M. LOUREIRO ET AL. 



Ringuelet, R. A. 1975. Zoogeografia y ecologia de los peces de aguas continentales de la 
Argentina y consideraciones sobre las areas ictiológicas de America del Sur. Ecosur, 
Corrientes, 2(3): 1-122. 

Santamaria, L. & Klaassen, M. 2002. Waterbird-mediated dispersal of aquatic organisms: an 
introduction. Acta Oecologica 23: 115-119. 

Sumner, A. T. 1972. A simple technique for demonstrating centromeric heterochromatin. 
Experientia Cellular Research 75: 304-306. 

Vaz Ferreira, R., Sierra de Soriano, B. & Scaglia de Paulete, S. 1964. Très especies nuevas 
del gènero Cynolebias Steindachner, 1876 (Teleostomi, Cyprinodontidae). Comunica- 
ciones Zoológicas del Museo de Historia Natural de Montevideo 8: 1-36. 



Revue suisse de Zoologie 111(1): 31 -3 3; mars 2004 



Edaphus comellinii sp. n. aus Sri Lanka 

(Coleoptera: Staphylinidae) 

88. Beitrag zur Kenntnis der Euaesthetinen 

Volker PUTHZ 

c/o Limnologische Fluß-Station, MPI-Limnologie, Damenweg 1, D-36110 Schlitz, 

Deutschland. E-Mail: Stenus.Puthz@t-online.de 



Edaphus comellinii sp. n. from Sri Lanka (Coleoptera: Staphylinidae). 
88th Contribution to the knowledge of Euaesthetinae. - Description of 
Edaphus comellinii sp. n. from Sri Lanka, a species of the group of E. 
cribricollis Schaufuss, which is now represented by 3 species in Sri Lanka. 

Key-words: Coleoptera - Staphylinidae - Edaphus - new species - taxo- 
nomy. 

EINLEITUNG 

Unter den Paratypen des Edaphus cinghalicus Comellini fand sich im Muséum 
d'histoire naturelle de Genève (MHNG) eine neue Art der Gattung aus Sri Lanka, die 
ich hier beschreibe und zu Ehren von Monsieur André Comellini, der sich um die 
Erforschung der Edaphus von Sri Lanka verdient gemacht hat, benenne. Aus Sri Lanka 
sind bisher 18 Arten der weltweit verbreiteten Gattung bekannt, die meisten von A. 
Comellini beschrieben. Während der Revision des Artenkomplexes um Edaphus 
cinghalicus Comellini (Puthz, 2003) habe ich auch die Typen der genannten Art unter- 
sucht und dabei unter den Paratypen zwei Stücke der neuen Art entdeckt; ein weiteres 
Stück fand sich unter unbestimmtem Material. 

Edaphus comellinii sp. n. 

Material: 3-Holotypus und 1 c?-Paratypus: CEYLON (= SRI LANKA): Central, 
Matale, 400 m, 17.1.1970, Mussard, Besuchet & Löbl (7) (beide Stücke tragen auch die 
Paratypus-Etiketten von Comellini „E. cinghalicus n. sp.")(MHNG); 1 Ç-Paratypus: Polo- 
nuaruwa, 31. VI. 1965, R. Mussard no. 5 (coll. Puthz). 

Beschreibung: Länge: 1,0 mm (Vorderkörperlänge: 0,6 mm). Hell bräunlich, 
glänzend, Pronotum sehr fein und wenig dicht punktiert; Beborstung fein, anliegend. 
Stirn mit Seitenkielen. 

Proportionsmaße des Holotypus: Kopfbreite: 27,4; Augenabstand: 19; Wangen- 
länge: 5; Schläfenlänge: 1,5; Augenlänge: 8; Pronotumbreite: 28,5; Seitenfalten- 
abstand: 18; Pronotumlänge: 26; größte Elytrenbreite: 39,5; größte Elytrenlänge: 33; 
Nahtlänge: 28. 



Manuskript angenommen am 17.10.2003 



32 



V. PUTHZ 






1 



Figs 1-3 
Edaphus comellinii sp. n. (Holotypus): 8. Sternit (1), Aedoeagus, ausgestülpt (2) und Sperma- 
pumpe (3).- Maßstab = 0,1 mm. 



Männchen: 8. Sternit (Fig. 1). Spermapumpe (Fig. 3). Aedoeagus (Fig. 2), aus- 
gestülpt. 

Kopf wenig schmäler als das Pronotum, Augen fein facettiert, vergleichsweise 
klein, Schläfen deutlich ausgeprägt, Stirn mit deutlichen Seitenkielen, hintere 
Querfurche und vordere Längsfurchen deutlich eingeschnitten, vorderer Mittelteil 
breiter als jedes der Seitenstücke, breitrund erhoben, unpunktiert, vom Clypeus durch 
eine schmale Querrandung abgesetzt, vordere Seitenteile mit wenigen winzigen 
Punkten. Fühler kurz, 7.-9. Glied fast so lang wie breit, Keule zweigliedrig, deutlich 
abgesetzt, vorletztes Glied etwa so lang wie breit. Pronotum wenig breiter als lang, 
hinten kräftig eingeschnürt, mit deutlichen Seitenfältchen, die durch eine feine Linie 
über die Pronotummitte hinaus nach vorn verlängert sind (dies wird besonders bei 
schrägem Lichteinfall erkennbar), Mittelfältchen bis zur Basis durchgehend, und mit 6 
basalen Grübchen. Punktierung sehr fein, aber erkennbar und ziemlich weitläufig. 
Elytren trapezoid, erheblich breiter als lang, Schultern einfach, Punktierung un- 
deutlich, sehr fein, flach und weitläufig. Abdomen zerstreut mit winzigen Punkten 
versehen, Mittelkiel des 3. Tergits gut bis zur Tergitmitte reichend. 

Bemerkungen: Diese neue Art ist dem E. sparsus Puthz aus Malaysia zum 
Verwechseln ähnlich; sie unterscheidet sich aber von ihm durch längere Seitenfältchen 
des Pronotums, feinere Punktierung desselben und durch kleineren Aedoeagus. Von E. 
cinghalicus unterscheidet sie sich sofort durch fehlende lange Mittelfurche und kürzere 
Seitenfalten des Pronotums. In meiner Bestimmungstabelle (Puthz, 1979) muss die Art 



EUDAPHUS COMELLINU SP. N. AUS SRI LANKA 33 

bei Leitziffer 13 eingefügt werden. Comellini nennt in seiner Arbeit bei E. cinghalicus 
2 Paratypen von Matale, 1 S und 1 ? : beide Stücke sind jedoch Männchen und 
gehören zu der neuen Art. 

LITERATUR 

Comellini, A. 1977. Nouvelles espèces d' Edaphus Motsch. du Sri Lanka (Coleoptera Staphy- 

linidae). Mitteilungen der schweizerischen entomologischen Gesellschaft 50 : 251-268. 
Puthz, V. 1979. Die vorder- und hinterindischen Arten der Gattung Edaphus Motschulsky 

(Coleoptera, Staphylinidae). Annales historico-naturales Musei Nationalis Hungarici 

71: 107-160. 
Puthz, V; 2003. Beiträge zur Kenntnis der Euaesthetinen LXXXVII. Edaphus cinghalicus 

Comellini, ein Komplex aus mehreren Arten (Staphylinidae, Coleoptera). Philippia 11: 

69-76. 



Revue suisse de Zoologie 111 (1): 35-55; mars 2004 



Two new taxa of Leptodirini (Coleoptera: Leiodidae, Choie vinae) 
from the Cantabrian cornice (Asturias, Spain). 
Biogeographical observations 

José Maria SALGADO COSTAS 1 & Javier FRESNEDA 2 

1 Dpto. de Biologia Animal, Facultad de Biologia, Universidad de Leon. 
E-24071 Leon, Espana. E-mail: dbajsc@unileon.es 

2 Ca de Massa, E-25526 Llesp-El Pont de Suert, Lleida, Espana. 
E-mail: ffresned@pie.xtec.es 



Two new taxa of Leptodirini (Coleoptera: Leiodidae, Cholevinae) from 
the Cantabrian cornice (Asturias, Spain). Biogeographical observ- 
ations. - Quaestus (Asturianella) incognitus subgen. n., sp. n, and Quaestus 
(Speogeus) mermejaensis sp. n. are described from caves located in the 
eastern foothills of the Sierra Mermeja (Asturias, Spain). Both taxa are 
included in the section Quaestus (sensu Salgado, 2000). Keys for a better, 
more precise placement of the new taxa are also given as well as biogeo- 
graphical observations and a study on colonization by the species captured 
in the four caves examined in this paper, in comparison with other morpho- 
logically similar species or those living in neighbouring areas. 

Key-words: Coleoptera - Leiodidae - Leptodirini - taxonomy - Quaestus 
(Asturianella) incognitus sp. n. - Quaestus (Speogeus) mermejaensis sp. n. - 
caves - colonization. 

INTRODUCTION 

One of the karstic areas with the largest troglobitic fauna stretches over a series 
of small mountain ranges situated towards the south of Asturias comprising the 
Crespón, Mermeja, Corteguero, Pintacanales, Ques and Giblaniella sierras. They cover 
an area of approximately 600 km^ between the Ponga, Sella, Infierno, Marea, Nalón, 
S. Isidro and Aller river basins (fig. 27). Eleven species of troglobitic Leptodirini are 
known from this area. They have been described over the last 20 years (Salgado, 1982, 
1984, 1985, 1988, 1989) and are at different stages of evolution, indicating that the 
caves were colonized during different glacial periods in the Pleistocene. 

Undoubtedly, the presence of such a high number of species in such a small area 
is explained by the occurrence of great orogenic convulsions causing the formation of 
small mountain ranges and numerous faults, landslides, overthrusts, etc. Well-isolated 
areas were formed and this interrupted the genetic flow of the populations, the main 
reason for speciation, which in this case is clearly allopatric (Belles & Martinez, 1980; 
Galân, 1993; Salgado, 1995; Peck, 1998). 



Manuscript accepted 19.08.2003 



36 J- M. SALGADO COSTAS & J. FRESNEDA 

In one of the previously-mentioned mountain ranges, the Mermeja, various 
samplings were carried out intermittently in 4 caves. A known species, Quaestus 
(Quaestus) longicornis (Salgado, 1989) was captured together with one proximate to 
Q- (Q.) jeannei (Coiffait, 1965) and Q. (Q.) recordationis (Salgado, 1982), Quaestus 
(Quaestus) sp., this last species is still under study, due to the complexity of its struc- 
tures and distribution. Two new taxa, Quaestus (Asturianella) incognitus subgen. n., 
sp. n. and Q. (Speogeus) mermejaensis sp. n. were also captured. 

TAXONOMIC TREATMENT 

Description of the new subgenus of the genus Quaestus 

Asturianella subgen. n. 

Type species: Quaestus (Asturianella) incognitus sp. n. 

Diagnosis. The basic distinguishing characters are: antennal segments of club 
longer than wide; pronotum strongly transverse, parameres with 4 apical setae; very 
long, robust stylet with subrectangular basal plate; spermatheca with small spherical 
apical lobe, basal lobe highly developed and oblong, spermathecal duct long and fine. 

Description. Ovoide, anophthalmia small, less than 3 mm (fig. 1). Antennae not 
surpassing half body length, all segments longer than wide (fig. 6). Elliptical elytra 
following curve of prothoracic sides. Mesosternal carina developed, with rounded tip 
and expanded ventral margin (fig. 9). Legs short. Male protarsi with 4 clearly dilated 
segments (fig. 7); females with gracile 4-segmented protarsi. 

Aedeagus robust, weakly arcuate but with apex strongly curved towards ventral 
face. Ventral lamina of tegmen slightly wider than long. Parameres somewhat longer 
than median lobe, distally forming dilated club with 4 setae inserted. Internal sac with 
two bands almost the same length as sac; long developed stylet inserted in subrectan- 
gular basal plate, no other sclerotized pieces discernible, only scales in basal region and 
fine spines and pilosity near median region (figs 2, 3 and 5). 

Apical lobe of spermatheca small and spherical, basal lobe expanded and long 
spermathecal duct weakly dilated along anterior region (fig. 4). 

Etymology. The subgeneric name is a reference to Asturias, the region of Spain 
where the specimens belonging to this new taxon were captured. The name is of 
feminine gender. 

Discussion. Some of the characters of the external morphology of the new 
taxon, including, 8th, 9th and 10th segments scarcely long; pronotum transverse with 
uniformly arcuate sides, elytra quite narrow in posterior region, and in particular, the 
genitalia, with a well-developed stylet and the spermatheca exhibiting an easily-distin- 
guishable spherical apical lobe, are also observed in the genus Quaestus. However, 
there is one unique and completely distinguishing character in this new taxon, the 4 
setae in the apical region of the parameres, which is only present in the genus 
Speocharinus Espanol & Escolà, 1997, as all the Quaestus have only 3. In exceptional 
cases some specimens may exhibit 4 setae, as indicated by Belles (1977) in Quaestus 
(Quaesticulus) minos (Jeannei, 1909) and Salgado (1982) in Quaestus (Quaestus) 
jeannei (Coiffait, 1965), both species have a supernummary seta, but it is only present 
in one specimen and on one paramere. 



TWO NEW TAXA OF LEPTODIRINI 



37 




Fig. 1 
Quaestus (Asturianella subgen. n.) incognitus sp. n. (paratype, male), habitus. 



38 J- M. SALGADO COSTAS & J. FRESNEDA 

Considering the importance of this character, 4 setae in the parameres, with 3 
setae never being observed in any specimen, and considering that the aedeagus and 
spermatheca are quite similar to those in specimens belonging to Quaestus, especially 
the group jeannei (Salgado, 1982), the category of genus should not be assigned to this 
new taxon. The category of monospecific subgenus is sufficient to establish its taxo- 
nomic indépendance. 

Taxonomy and key of the subgenera belonging to the genus Quaestus 

The section Quaestus (Salgado, 2000) includes the following valid genera: 
Breuilia Jeannei, 1909; Breuilites Salgado, 1980; Cantabrogeus Salgado, 2000; Espa- 
noliella Guéorguiev, 1976; Leonesiella Salgado, 1996; Quaestus Schaufuss, 1861; 
Oresigenus Jeannei, 1948 and Speocharinus Espanol & Escolà, 1997. 

The genera belonging to this section are characterized by the long elevated or 
low mesosternal carina (short in the subgenus Speogeus only). Internal sac of aedeagus 
with pilosity, scales, spinules, spines, sclerotised plates or generally long stylet, or a 
combination of all these structures. Spermatheca with generally discernible basal lobe, 
more developed than spherical apical lobe. The species belonging to these genera are 
found along the Cantabrian cornice, except for Quaestus (Quaesticulus) cisnerosi 
(Perez-Areas, 1872) which inhabits the Central Mountain Range of the Iberian 
Peninsula. 

Using studies by Salgado (1985, 1993, 2000), Newton (1998) and Perreau 
(2000) as a basis, the genus Quaestus includes 6 subgenera. 

1 Elytra lacking suturai striae or with very rudimentary weakly discernible 
stria, only visible in anterior region of elytrum. Stylet of internal sac of 
aedeagus bearing plate or basal bowl. Spermatheca with small apical 

lobe and generally long, fine spermathecal duct 2 

1' Elytra with suturai stria along all the elytrum, usually easily discernible. 
Sylet of internal sac of aedeagus lacking plate or basal bowl. Sper- 
matheca with big or small apical lobe and short spermathecal duct 4 

2 Parameres bearing 4 setae in apical region Asturianella subgen. n. 

2' Parameres with 3 setae in apical region 3 

3 Mesosternal carina short, if the apophysis is near the end of the metaster- 
num, the carina is always low. Angle of carina strongly obtuse. Ventral 
edge of carina always narrow. Mid posterior region of metasternum 
often bearing small fovea Speogeus Salgado, 1985 

3' Mesosternal carina long and elevated, apophysis always reaching meta- 
sternum. Angle of carina just over 90°. Ventral edge narrow or wide. 
Mid posterior region of metasternum lacking fovea . Quaestus Schaufuss, 1861 

4 Apical region of elytra pointed and diverging. Median lobe of aedeagus 
strongly expanded in mid region, with carina in ventral face. Internal sac 
lacking fanerae, only stylet present Samanolla Salgado, 2000 



TWO NEW TAXA OF LEPTODIRINI 39 

4' Apical region of elytra rounded and not diverging. Median lobe of 
aedeagus not strongly expanded in mid region, lacking carina in ventral 
face. Internal sac with fanerae and stylet 5 

5 Male metafemur with small denticle on posterior margin. Internal sac of 

aedeagus with robust teeth and spines, spinules or both 

Amphogeus Salgado, 2000 

5' Male metafemur lacking denticle in posterior margin. Internal sac of 

aedeagus with more or less developed spiny formations 

Quaesticulus Schaufuss, 1861 

Descriptions of the new species 

Quaestus (Asturianella) incognitus sp. n. 

Type material. Holotype, S, Cave of Julio, Caleao, T. M. de Caso (Asturias), 31 -VII- 
2002, J.M. Salgado leg., in Muséum d'histoire naturelle de Genève. Paratypes: 48 S S and 74 
9 9 , same data as Holotype. Cave of the Prau, Puente de Piedra-Buspriz, T. M. de Caso 
(Asturias), 17-VI-1989, 60 SS and 96 ? 2; 28-X-1989, 68 SS and 73 5 9; 31-VII-2002, 3 
S S and 10 9$, J.M. Salgado and D. Rodriguez leg. Cave of Carretera, Puente de Piedra- 
Buspriz, T. M. de Caso (Asturias), 28-X-1989, 2 S S and 3 9 9 , J.M. Salgado and D. Rodriguez 
leg. Cave of Rearco, Coballes, T. M. de Caso (Asturias), 22-IV-1989, 33 S S and 50 9 9 ; 15- 
rV-2003, 45 S S and 41 9 9, J.M. Salgado and D. Rodriguez leg. Muséum d'histoire naturelle 
de Genève; J.M. Salgado Coll., Dpto. de Biologìa Animal, Universidad de Leon (Spain); J. 
Fresneda Coll., Lleida (Spain); P.M. Giachino, Museo Regionale di Scienze Naturali, Torino 
(Italy); Museo de Zoologia de Barcelona (Spain); Muséum National d'Histoire Naturelle, Paris 
(France); Field Museum of Natural History, Chicago (USA). 

Diagnosis. 11th antennal segment twice the length of 10th; 8th, 9th and 10th 
segments slightly longer than wide; male protarsi as wide as maximum width of tibia; 
aedeagus with tip of median lobe curved towards ventral face; parameres somewhat 
longer than median lobe, with 3 short apical setae and one longer preapical seta; 
internal sac of aedeagus exhibiting pilosity, scales and very long robust stylet with sub- 
rectangular basal plate; spermatheca with small spherical apical lobe, and wide oval 
basal lobe, spermathecal duct very long. 

Description. Male. Holotype 6, 2.56 mm (paratypes: 2.53-2.70 mm). 
Elongate, elliptic body, not very narrow posteriorly (fig. 1). Uniformly reddish-brown. 
Shiny tegumenta. Antennae not surpassing half the body length, segments noticeably 
expanded from 7th (fig. 6). Of particular interest is the 11th segment which is twice the 
length of 10th; 8th, 9th and 10th segments slightly longer than wide; 3rd and 5th seg- 
ments equal and slightly longer than 4th and 6th, which are the same (Table I). 

Pronotum strongly transverse (maximum width/maximum length relationship: 
2.05), sides uniformly arcuate and widest at base. Elytra elongate, less than one and a 

Table I. Maximum lengths (L) and widths (W) of antennal segments (1 mm = 50 units). 



Segments 


I 


II 


III 


IV 


V 


VI 


VII 


vni 


IX 


X 


XI 


L 


6.5 


7.0 


5.0 


4.0 


5.0 


4.0 


5.5 


3.2 


4.1 


3.9 


8 


W 


2.5 


2.7 


2.0 


2.0 


2.0 


2.0 


2.8 


2.4 


3.1 


3.3 


3.6 



40 



J. M. SALGADO COSTAS & J. FRESNEDA 




Figs 2-4 
Quaestus (A.) incognitus sp. n.: (2) aedeagus, dorsal view; (3) apical region of the paramere; (4) 
spermathecal complex. 



half times width of pronotum, margins continuous, gradually narrowing towards tip. 
Suturai stria absent. Mesosternal carina elevated, apophysis reaching metasternum; 
obtuse angle with rounded tip and ventral margin wide (fig. 9). Legs slender, modera- 
tely long, with first segment of protarsi as wide as maximum width of protibiae (fig. 7). 
Aedeagus robust and long (0.77 mm), in dorsal view, median lobe with sides 
almost parallel as far as apical fourth then narrowing progressively, apical area weakly 
truncate (fig. 2). In lateral view distal region in extended pointed tip and curved 
towards ventral part (fig. 5). Parameres just surpassing tip of median lobe, apical region 



TWO NEW TAXA OF LEPTODIRINI 41 

expanded, resembling an axe, with 4 setae inserted, 3 short equal ones in apical mar- 
gin, a more robust one in club base near internal margin (fig. 3). Long robust stylet of 
internal sac with subrectangular basal plate, flanked by two thin very weakly sclero- 
tized plates; scales near basal region set out in shape of an arch, fine spinules and 
pilosity in two narrow rows near median region. Two reinforcement bands almost as 
long as sac (fig. 2). 

Female. As robust as male, with shorter antennae and proportionally thicker 
segments (Table II). Protarsi not expanded and tetramere. Eighth urite with short, 
narrow spiculum ventrale, twice as short as length of sternite (fig. 8). Spermathecal 
complex showing a spermatheca with very small spherical apical lobe, basal lobe twice 
as wide and almost three times longer than apical lobe; the joining between these lobes 
short, narrow and not sclerotized; spermathecal duct long, approximately 9 times as 
long as spermatheca which is slightly expanded in one third of its anterior length 
(fig. 4). 

Table II. Maximum lengths (L) and widths (W) of antennal segments (1 mm = 50 units). 



Segments 


I 


II 


III 


IV 


V 


VI 


VII 


VIII 


IX 


X 


XI 


L 


5.7 


6.2 


4.0 


3.7 


4.0 


3.7 


5.0 


2.7 


4.0 


3.8 


7.5 


W 


2.5 


2.7 


2.0 


2.0 


2.0 


2.0 


2.7 


2.3 


3.0 


3.2 


3.6 



Etymology. "Incognitus=unknown" refers to the long period of time that passed 
from when this new species was first captured (22-IV-1989) until it was described. 

Discussion. This new taxon shows certain similarities to two species in the 
group jeanne i, Quaestus (Quaestus) jeannei Coiffait, 1965 and Q. (Q.) recordationis 
Salgado, 1982, in their external morphology, shape of the median lobe, structures of 
the internal sac of the aedeagus and shape of the spermatheca (figs 10-17). Q. 
(Asturianella) incognitus sp. n. is differentiated from the two previously-mentioned 
species by the more slender antennal segments, all of which are longer than wide, and 
a more transverse pronotum; also, the median lobe of the aedeagus is more robust and 
has a slightly truncate apical area, more parallel sides in the median region and the 
apical curvature more pronounced. However, two characters make this new species 
unmistakeable and clearly distinguish it from any other species in the genus Quaestus, 
the shape of the club of the parameres and the 4 setae inserted. 

Finally, it should be mentioned that the antennal segments of the specimens 
captured in the Rearco cave are slightly longer than those of the specimens captured in 
the other three. 

Quaestus (Speogeus) mermejaensis sp. n. 

Type material. Holotype, 6 . Cave of the Prau, Puente de Piedra-Buspriz, T. M. de Caso 
(Asturies, Spain), 17-VI-1989, J.M. Salgado leg., in Muséum d'histoire naturelle de Genève. 
Paratypes: Cave of the Prau, Puente de Piedra-Buspriz, T. M. de Caso (Asturies, Spain), 17-VI- 
1989, 35 6 6 and 47 9 9 ; 31-VII-2002, 10 6 6 and 10 9 9, J.M. Salgado and D. Rodriguez 
leg. Cave of the Carretera, Puente de Piedra-Buspriz, T. M. de Caso (Asturies, Spain), 28-X- 
1989, 2 6 6 and 4 $ $ , J.M. Salgado and D. Rodriguez leg. Cave of Rearco, Coballes, T. M. de 
Caso (Asturies, Spain), 22-IV-1989, 2 6 6 and 2 $ 9; 15-IV-2003, 4 6 6 and 12 $ 9, J.M. 



42 



J. M. SALGADO COSTAS & J. FRESNEDA 




Figs 5-7 
Quaestus (A.) incognitus sp. n.: (5) aedeagus, lateral view; (6) antenna; (7) protarsum, dorsal 



Salgado and D. Rodriguez leg. Muséum d'histoire naturelle de Genève; J.M. Salgado Coll., 
Dpto. de Biologia Animal, Universidad de Leon (Spain); J. Fresneda Coll., Lleida (Spain); P.M. 
Giachino, Museo Regionale di Scienze Naturali, Torino (Italy); Museo de Zoologia de Barcelona 
(Spain); Muséum National d'Histoire Naturelle, Paris (France); Field Museum of Natural 
History, Chicago (USA). 

Diagnosis. A species of the tribe Leptodirini characterized by: elongate weakly 
convex body; 8th antennal segment transverse, 9th and 10th segments slightly longer 
than wide; elytral margins parallel along mid region; mesosternal carina low and short 
with narrow ventral edge and strongly obtuse angle; parameres clearly surpassing tip 
of median lobe, dilated club with 3 setae and one or two easily discernible piliferous 
pores. 



TWO NEW TAXA OF LEPTODIRINI 43 

Description. Male. Holotype 6, 2.62 mm (paratypes: 2.58-2.82 mm). Body 
elongate, elliptical weakly convex, reddish-brown and covered in short laid back yel- 
lowish pilosity. Antennae quite long, clearly surpassing half body length (fig. 23); 11th 
segment three times longer than wide and three times as long as 10th segment; 8th seg- 
ment transverse; 3rd to 6th segments similar in length (Table III). 

Table III. Maximum lengths (L) and widths (W) of antennal segments (1 mm = 50 units). 



Segments 


I 


II 


III 


IV 


V 


VI 


VII 


VIII 


IX 


X 


XI 


L 


7.2 


8.0 


5.1 


4.9 


5.0 


5.0 


6.4 


3.0 


4.1 


4.0 


11.7 


W 


2.7 


2.4 


2.0 


2.0 


2.0 


2.0 


3.4 


3.0 


3.7 


3.8 


4.0 



Pronotum transverse, almost twice as wide as long (1.88), sides uniformly 
curved, attenuated towards front and weakly narrow towards back; basal area as wide 
as elytra. Elytra elongate, 1.65 times longer than wide, with parallel margins along two 
thirds of their length, gradually narrowing to the tip. Suturai stria not discernible. 
Mesosternal carina low, with very obtuse angle and very rounded tip; ventral margin 
narrow, apophysis slightly surpassing mesosternum (fig. 21). Legs slender and quite 
long, protarsi dilated, the first being wider than the maximum width of the protibiae 
(first protarsi/protibiae ratio = 1.25). 

Aedeagus weakly wide and quite long (0.60 mm) in lateral view, with uniformly 
arcuate median lobe and curved pointed tip (fig. 22). In dorsal view, tip of median lobe 
truncate and slightly arcuate inwards in median region (fig. 18). Parameres clearly sur- 
passing tip of median lobe, with expanded shovel-like club, internal margin pointed, 
resembling a peak; of the three setae inserted in the club, the lower one is long and 
inserted in the base, fairly distant from the two small apical ones, with insertion pores 
close together and one or two pores above the apical setae which have no setae (fig. 
19). Internal sac with numerous weakly-sclerotised scales scattered in mid basal re- 
gion; stylet very long and fine, almost as long as median lobe, with well-developed 
subrectangular insertion plate. Two reinforcement bands as long as internal sac 
(fig. 18). 

Female. Generally a little more robust than male, with elytra slightly more 
convex. Antennae somewhat shorter and segments comparatively thicker than in male 
(Table IV). Spiculum ventrale of 8th urite very short and narrow, 3 times shorter than 
length of sternite. Spermathecal complex with spermatheca quite rectilinear, apical 

Table IV. Maximum lengths (L) and widths (W) of antennal segments (1 mm = 50 units). 



Segments 


I 


II 


III 


IV 


V 


VI 


VII 


VIII 


IX 


X 


XI 


L 


6.9 


7.8 


5.0 


4.7 


5.0 


4.8 


6.0 


2.8 


4.0 


4.0 


9.0 


W 


2.7 


2.5 


2.1 


2.1 


2.1 


2.1 


3.5 


2.8 


3.7 


3.8 


4.0 



lobe small and spherical, basal lobe much longer than wide, both well-sclerotised, the 
joining region more narrow and translucent; spermathecal duct fine, just over 4 times 
as long as spermatheca, and slightly more expanded along anterior fifth (fig. 20). 



44 



J. M. SALGADO COSTAS & J. FRESNEDA 




0.5mm 




0.5mm 



Figs 8-9 
Quaestus (A.) incognitus sp. n.: (8) urite female, ventral view; (9) mesosternal carina, lateral 
view. 



Etymology. The name refers to the caves situated in the foothills of the Sierra 
Mermeja where specimens belonging to this new species were found. 

Discussion. The only species that Quaestus (Speogeus) mennejaensis sp. n. 
could be mistaken for is Q. (S.) nietoi (Salgado, 1988) as they are similar morpholo- 
gically. There are, however, constant and significant differences in some external mor- 
phological characters and in the genitals. These differences are: 8th antennal segment 
in Q. (S.) mennejaensis transverse, 9th and 10th slightly longer than wide; apophysis 
of mestosternal carina only just surpassing mesosternum; parameres widely surpassing 
median lobe, insertion pores of both apical setae more separated, pore of basal seta at 
a considerable distance. Whilst all the antennal segments in Q. (S.) nietoi are clearly 
longer than wide, the apophysis of the mesosternal carina almost reaches the metaster- 
num, the club of the parameres has a different shape and only just surpasses the tip of 
the median lobe, the insertion pores of the apical setae are closer together and the 
insertion pore of the basal seta is slightly separated (figs 24, 25); finally, the area 
joining the spermathecal lobes is shorter and the spermathecal duct is clearly longer 
(fig. 26). Added to all this, the populations of these species are isolated by barriers such 
as overthrusts and sandstone areas separating them, as can be seen on the distribution 
map (fig. 27). 



TWO NEW TAXA OF LEPTODIRINI 



45 




Figs 10-12 

Quaestus (Quaestus) jeannei (Coiffait, 1965): (10) aedeagus, dorsal view; (11) apical region of 
the paramere; (12) spermathecal complex. 

KEY TO THE SPECIES OF THE SUBGENUS SPEOGEUS 

1 Basal plate of stylet of internal sac rounded 2 

1' Basal plate of stylet of internal sac subrectangular 3 

2 Male protarsi more narrow or slightly wider than maximum width of 
protibiae; 11th antennal segment 3 times or just over 3 times as long as 
10th; parameres narrow, with tip in the shape of an oblong club 
Q. (S.) amicalis (Salgado, 1984) 



46 J- M. SALGADO COSTAS & J. FRESNEDA 

2' Male protarsi almost twice as wide as maximum width of protibiae; 11th 
antennal segment at most 2.5 times as long as 10th; parameres wide, tip 
curved and club barely differentiated Q. (S.) littoralis Salgado, 1999 

3 Parameres longer than median lobe, with club flattened and more or less 
pointed towards inner side 4 

3' Parameres as long as or slightly shorter than median lobe, with tip 

rounded and lacking distinguishable club 5 

4 8th, 9th and 10th antennal segments clearly longer than wide; apophysis 
of mesosternal carina reaching or just surpassing mid region of meso- 
sternum; parameres slightly surpassing tip of median lobe; distance 
between the two apical setae of club of parameres less then three times 
the distance between them and insertion point of lower seta 
Q. (S.) nietoi (Salgado, 1988) 

4' 8th antennal segment transverse, 9th and 10th segments slightly longer 
than wide; apophysis of mesosternal carina not reaching mid region of 
mesosternum; parameres widely surpassing tip of median lobe; distance 
between the two apical setae of club of parameres almost 5 times the 
distance between them and insertion point of lower seta 
Q. (S.) mermejaensis sp. n. 

5 3 setae of parameres inserted in apical region of club; mid-apical region 

of internal sac lacking developed fanerae . . . Q (S.) avicularis (Salgado, 1985) 
5' 3 setae of parameres not inserted in apical region of club, one inserted 
near base; mid apical region of internal sac with two or three groups of 
well-developed fanerae Q. (S.) diegoi (Salgado, 1989) 

BIOLOGICAL AND ECOLOGICAL OBSERVATIONS 

The following is a description of some of the characteristics of the four caves 
sampled in this study. 

Julio cave (UTM: 30TUN023803). This cave is situated 1000 m. in a limestone 
area from the Carboniferous period. It is surrounded by Corylus avellana L. and Fagus 
sylvatica L. as the predominating plant species, with a rocky area nearby. It is 800 m 
long and easy to enter and walk through. The first 50 m are highly eroded and there is 
a large clay substrate. Facing south-east, the temperature inside is 12° C (summer) with 
97-98% relative humidity above the first 50 m. Most of the catches were carried out 
randomly at between 50 and 150 m in heaps of organic plant material and scattered bat 
faeces and using attraction traps (cheese, fruit, liver). Q. (A.) incognitus sp. n. cohabits 
with Q. (Q.) longicornis. The populations of the latter species are also found in caves 
in the bordering area of the Sierra del Crespón. 

Prau cave (UTM: 30TUN055824). Situated at 815 m., also in limestone areas 
from the Carboniferous period. The mouth of the cave faces south and the surrounding 
area is predominated by Quercus robur L., Alnus glutinosa (L.) Gaertn. and Corylus 
avellana L. It is an active cave still in the process of formation and approximately 200 



TWO NEW TAXA OF LEPTODIRINI 



47 




Figs 13-15 
Quaestus (Q.) recordationis (Salgado, 1982): (13) aedeagus, dorsal view; (14) apical region of 
the paramere; (15) spermathecal complex. 



m long. A number of stalactites, stalagmites and outflows can be seen inside. The 
temperature is between 8° C (winter) and 11° C (summer), with 98.5% constant rela- 
tive humidity. 

As the ground is free from organic material and there are a number of gours, 
catches were only possible using attraction and pit fall traps, especially in the centre of 



48 



J. M. SALGADO COSTAS & J. FRESNEDA 




Figs 16-17 
Lateral view of the aedeagus: (16) Quaestus (Q.) jeannei (Coiffait, 1965); (17) Quaesîus (Q.) 
recordations (Salgado, 1982). 



the cave. From the biological perspective, this cave is of great interest as three troglo- 
bitic species of Leptodirini cohabit there: Q. (Q.) longicornis, Q. (A.) incognitus sp. n. 
and Q. (S.) mermejaensis sp. n., as well as Notidocaris franzi Jeannei, 1956, a mu- 
siculous species captured at the entrance. A very interesting member of the family 
Trechinae, Apoduvalius sp., morphologically very close to A. espanoli Salgado, 1996, 
is also found there. 



TWO NEW TAXA OF LEPTODIRINI 



49 




0.5 mm 



Figs 18-21 
Quaestus (Speogeus) mermejaensis sp. n.: (18) aedeagus, dorsal view; (19) apical region of the 
paramere; (20) spermathecal complex; (21) mesosternal carina. 



50 



J. M. SALGADO COSTAS & J. FRESNEDA 




E 
E 
in 



23 




Figs 22-23 
Quaestus (S.) mermejaensis sp. n.: (22) aedeagus, lateral view; (23) antenna. 



Carretera Cave (UTM: 30TUN056830). This cave is 400 m from the cave of 
the Prau. It is a small, very narrow gallery less than 25 m long. It is situated at 800 m 
alt. in limestone areas from the Carboniferous period. The entrance faces south-east, 
and is hidden mainly by Sambucus nigra L. and Quercus robur L. brambles and shrub 
branches. Because of its short length it is strongly influenced by atmospheric changes, 
so only a few specimens of Q. (A.) incognitus sp. n., Q. (S.) mermejaensis sp. n. and 
Q. (Q.) longicornis were captured using traps in the deepest parts. 



TWO NEW TAXA OF LEPTODIRINI 



51 



in 




Figs 24-26 
Quaestus (Speogeus) nietoi (Salgado, 1988): (24) aedeagus, dorsal view; (25) apical region of 
the paramere; (26) spermathecal complex. 



Rearco cave (UTM: 30TUN054846). Of all the caves, this one is situated 
furthest north at an altitude of 650 m. As in the previous cases the limestone area where 
the cave was formed belongs to the Carboniferous period. The surrounding vegetation 
is Crataegus monogyna Jacq., Corylus avellana L., Castanea sativa L. and Quercus 
robur L. The cave was formed from a joint, producing a very narrow gallery leading 
into a larger chamber, 2 m high at the most and from which a network of very narrow 
low galleries lead off. The accessible area is approximately 175 m long. As the cave 



52 



J. M. SALGADO COSTAS & J. FRESNEDA 




TWO NEW TAXA OF LEPTODIRINI 53 

entrance is very small, the physical factors are constant all the way through, with a 
temperature of 12° C and 98% relative humidity. 

The fauna were captured in the first few meters inside the cave in organic plant 
matter, and in the rest of the cave in small heaps of organic animal material, especially 
bat faeces. The cave is of great biological interest because, to date, it is the only one in 
the world where 4 troglobitic species belonging to the same genus, Quaestus (Q.) 
longicornis, Q. (Quaestus) sp., Q. (A.) incognitus sp. n. and Q. (S.) mermejaensis 
sp. n. have been located, which is possible because they are all at different stages of 
evolution and have different microniches; for example, the two species belonging to 
the subgenus Quaestus, Q. (Q.) longicornis and Q. (Quaestus) sp., besides having 
completely different genitalia, the first species has clearly evolved more than the 
second because the appendices, legs and antennae are much longer and much more 
slender. 

GEOGRAPHICAL DISTRIBUTION 

As already pointed out the two new taxa Q. (A.) incognitus and Q. (S.) 
mermejaensis are found in caves in the eastern foothills of Sierra Mermeja and cohab- 
it in the Prau, Carretera and Rearco caves. The two new species also cohabit with Q. 
(Quaestus) sp. and Q. (Q.) longicornis which is more widespread, though the morpho- 
logical differences amongst them are very evident. Also, Q. (S.) mermejaensis has 
similar morphological characters to Q. (S.) nietoi, which inhabits a nearby karstic 
region and is much more widely distributed. It would be interesting to carry out a bio- 
geographical analysis to obtain a better understanding of their genetic and geographical 
isolation in relation to the present distribution of the different species (fig. 27). 

The various degrees of evolution may be considered as multistage processes in 
which several events occurred and gave rise to non-allopatric and allopatric speciation. 
Non-allopatric speciation occurred as a result of a diverging process of adaptation 
during the colonization of hypogeous environments by a population via gradual steps 
or stages with changes brought about by very different factors (Wilson, 1989; Bush, 
1994). In the case of allopatric speciation, reproductive isolation was possible as a 
result of great climatic changes such as glaciation during the Pleistocene (Jeannel, 
1942; Galân, 1993; Trajano, 1995) and the search for shelter (Belles, 1987; Haffer, 
1994), or the fragmentation of the habitat in isolated karstic areas (Belles & Martinez, 
1980; Reveillet, 1980; Salgado, 1995), which obviously occurred in this study area. 

An examination of the species cohabiting in the caves described in this study, 
or morphologically proximate species inhabiting neighbouring areas, revealed that Q. 
(Quaestus) sp., which is very widespread, cohabits with Q. (Speogeus) nietoi, Q. (S.) 



Fig. 27 
Distribution map of the species: 1, Quaestus (Asturianella) incognitus sp. n.; 2, Quaestus (Speo- 
geus) mermejaensis sp. n.; 3, Quaestus (Quaestus) longicornis (Salgado, 1989); 4, Quaestus 
(Quaestus) sp.; 5, Quaestus (Speogeus) avicularis (Salgado, 1985); 6, Quaestus (S.) amicalis 
amicalis (Salgado, 1984); 7, Quaestus (S.) amicalis dilatatus (Salgado, 1984); 8, Quaestus (Q.) 
luctuosus (Salgado, 1984); 9, Quaestus (Q.) jeannei (Coiffait, 1965); 10, Quaestus (S.) nietoi 
(Salgado, 1988); 11, Quaestus (Q.) recordationis (Salgado, 1982). 



54 J- M. SALGADO COSTAS & J. FRESNEDA 

avicularis, Q. (S.) amicalis dilatatus, Q. (Quaestus) longicornis, Q. (Q.) luctuosus, and 
now with Q. (S.) mermejaensis sp. n. and Q. (A.) incognitus sp. n. in various caves. Of 
all the species mentioned, Q. (Quaestus) sp. is the least evolved, as shown by its mor- 
phological characters; the differentiating characters and genetic isolation may have 
occurred because the hypogeous environment was colonized in more recent periods. 
This theory is supported by the fact that the species is much more dispersed in karstic 
areas isolated by existing barriers which can be reached through the superficial under- 
ground compartment (MSS or Milieu Souterrain Superficiel, Juberthie et al., 1981) as 
demonstrated by 3 catches recently made on the upper karst of Munera (12-IX-1999), 
Pandanes (29-VI-2001) and Coballes (31-VH-2002). 

Q. (Q.) longicornis is only known from the deep underground environment. 
Data on 3 caves in the Sierra del Crespón (Salgado, 1989) already existed and the 4 
new ones from the Sierra Mermeja can now be added. Catches made in the caves show 
that there must have been a connection between the populations of these mountain 
ranges via fissures in the deep underground environment, the only way of establishing 
links between the populations of these fairly or very evolved species before the pres- 
ent barriers became established. The species cohabits with Q. (S.) avicularis and Q. 
(Quaestus) sp., in the Sierra del Crespon, with no impediments as they belong to dif- 
ferent evolutionary stages, and in Sierra Mermeja with the two new taxons studied in 
this paper, and Q. (Quaestus) sp. 

With regard morphological characters and proximity of colonization area, Q. 
(S.) nietoi is closest to Q. (S.) mermejaensis sp. n. Both belong to the same evolution- 
ary stage and are therefore not found in the same area or cave. Q. (S.) nietoi is much 
more widely distributed than Q. (S.) mermejaensis, and is known from various caves 
in the karstic area between the Aller and Nalón river basins but limited by the Crespón, 
Mermeja and Ques sierras. At present both species are found in the deep underground 
compartment as they have never been located in the superficial underground environ- 
ment through which they could migrate. Differentiation therefore occurred as a result 
of isolation by deep sandstone strata barriers and overthrusts which stopped the gene 
flow. 

ACKNOWLEDGEMENTS 

This paper was funded by the "Fauna Ibèrica" project (REN 2001-1956-C17- 
09). Our thanks go to "Fauna Ibèrica" for allowing the publication of the graphs 
included in the paper. 

REFERENCES 

Belles, X. 1977. Notas sobre Speocharis minos Jeannel y otros Catópidos recogidos en cuevas 

de la provincia de Burgos. Graellsia 31: 119-124. 
Belles, X. 1987. Fauna cavernicola i intersticial de la Peninsula Ibèrica i les illes Balears. 

C.S.I.C., Editorial Moll, Mallorca, 207 pp. 
Belles, X. & Martinez, A. 1980. La ecologia y la especiación de los Bathysciinae (Col. 

Catopidae) en la region del Penedés (Cataluna, Espana). Mémoires de Biospéologie 7: 

221-233. 
Bush, G. 1994. Sympatric speciation in animals: new wine in old bottles. Trends in Ecology & 

Evolution 9(8): 339-364. 



TWO NEW TAXA OF LEPTODIRINI 55 



Galan, C. 1993. Fauna hipógea de Guipüzcoa: su ecologia, biogeografìa y evolución. Munibe 
45: 3-1663. 

Haffer, J. 1994. Avian speciation in tropical South America. Nuttall Ornithology Club, 
Cambridge, 390 pp. 

Jeannel, R. 1942. La genèse des faunes terrestres. Eléments de biogéographie. Presses 
Universitaires, Éditions Gallimard, Paris, 514 pp. 

Juberthie, C, Bouillon, M. & Delay, B. 1981. Sur l'existence d'un milieu souterrain super- 
ficiel en zone calcaire. Mémoires de Biospéologie 8: 77-94. 

Newton, A.F. 1998: Phylogenetic problems, current classification and generic catalogue of 
world Leiodidae (including Cholevinae). In: Giachino, P.M. & Peck, S.B. (eds). Phylo- 
geny and evolution of subterranean and endogean Cholevidae (=Leiodidae Cholevinae). 
Proceedings of XX International Congress of Entomology. Firenze, 1996. Atti Museo 
Regionale di Scienze Naturali, Torino: 41-178. 

Peck, S.B. 1998. Phylogeny and evolution of subterranean and endogean Cholevidae (=Leio- 
didae Cholevinae) an introduction. In: Giachino, P.M. & Peck, S.B. (eds). Phylogeny 
and evolution of subterranean and endogean Cholevidae (=Leiodidae Cholevinae). 
Proceedings of XX International Congress of Entomology. Firenze, 1996. Atti Museo 
Regionale di Scienze Naturali, Torino: 11-40. 

Perreau, M. 2000. Catalogue des Coléoptères Leiodidae Cholevinae et Platypsyllinae. 
Mémoires de la Société Entomologique de France 4: 1-460. 

Reveillet, P. 1980. Les coléoptères souterrains du Vercors (France, SE). Essai de biogéogra- 
phie. Mémoires de Biospéologie 7: 173-186. 

Salgado, J.M. 1982. Nuevos Bathysciinae (Coleoptera: Catopidae) de grupo Speocharis 
jeannei. Boletin de Ciencias Naturales, I.D.E.A. 30: 49-58. 

Salgado, J.M. 1984. Estudio sobre el "grupo Speocharis occidentalis Jeannel, 1911" (Coleo- 
ptera, Catopidae). Mémoires de Biospéologie 11: 257-264. 

Salgado, J.M. 1985. Un nuevo subgénero de Bathysciinae en cuevas Asturianas (Coleoptera, 
Catopidae). Nouvelle Revue d'Entomologie (N.S.) 2 (3): 261-265. 

Salgado, J.M. 1988. Nuevo Speocharis del "grupo occidentalis" (Col., Catopidae). Mémoires 
de Biospéologie 15: 61-66. 

Salgado, J.M. 1989. Nuevos datos sobre la distribución de especies del "grupo Speocharis 
occidentalis" (Col., Catopidae). Mémoires de Biospéologie 16: 125-130. 

Salgado, J.M. 1993. Los Bathysciinae (Col., Cholevidae) Cantâbricos: "Sección Speocharis". 
Mémoires de Biospéologie 20: 221-230. 

Salgado, J.M. 1995. Fauna troglobia de la Cordillera de Sueve (Asturias, Espana). Mémoire de 
Biospéologie 22: 129-137. 

Salgado, J.M. 2000. A new revision and taxonomic position for the Cantabrian Leptodirini: sec- 
tions Quaestus and Speonomidius (Coleoptera: Cholevidae). Annales de la Société 
Entomologique de France (N.S.) 36 (1): 45-59. 

Traiano, E. 1995. Evolution of tropical troglobites: applicability of the model of Quaternary cli- 
matic fluctuations. Mémoires de Biospéologie 22: 203-209. 

Wilson, D. 1989. The diversification of simple gene pools by density and frequency-dependent 
selection. In: Otte, E. & Endler, J.A. (eds). Speciation and its consequences. Sinaver 
Assoc. Inc. Publish.: 366-385. 



Revue suisse de Zoologie 111 (1): 57-62; mars 2004 



Description of the first apterous genus of Sciomyzidae (Diptera), 
from Nepal 

Laszló PAPP 

Department of Zoology, Hungarian Natural History Museum, Baross u. 13, Budapest, 

H-1088 Hungary. 

E-mail: lpapp@zoo.zoo.nhmus.hu 



Description of the first apterous genus of Sciomyzidae (Diptera), from 
Nepal. - Apteromicra gen. n. (type species A. parva sp. n.) is described and 
illustrated from Nepal. This is the first known apterous species within 
Sciomyzidae, its systematic position is discussed in detail. 

Key- words: Diptera - Sciomyzidae - Sciomyzini - Apteromicra - taxonomy 
- apterous flies - Nepal. 

INTRODUCTION 

Reduction and loss of wings among the dipterous flies are rather common and 
those species have always provoked interest among dipterists. Hackman (1964) re- 
ferred to aptery as "...known from more than 20 dipterous families" but since then such 
reduced-winged or apterous species have been discovered in additional families [e.g. 
Drosophilidae, Papp (1979)]. When discussing the environments in which short- 
winged or apterous Diptera are found, Hackman (1964) referred principally to high 
altitude Diptera. Other environments (and congruent Diptera groups) are: Diptera in 
arctic, subarctic and subantarctic mainland habitats; nival Diptera; Diptera of oceanic 
islands; Diptera on sea shores; marine Diptera; Diptera in various terricolous and hypo- 
geous habitats; Diptera in nests of Hymenoptera and termites; and parasites on warm- 
blooded animals. 

In the course of my visit to the Muséum d'histoire naturelle Genève in 1986 I 
found a small apterous fly among extensive collections, which coleopterists of the 
museum had made by sieving and subsequent funnelling in the Oriental region. I 
borrowed that fly but it was untouched for years. When I decided to study it early in 
2002, immediately I found it peculiar, and its family relegation questionable, although 
it is obviously an "acalyptrate" fly. 

Keys to families of adult Diptera employ characteristics of wings extensively. 
Consequently, reduced- winged or apterous flies have to be dealt with separately. In the 
key for the Palaearctic Diptera families, Papp & Schumann (2000) keyed 23 families 
of those flies in the last part of the key. Among the 23 families there are only six 
families of the "acalyptrates". Our fly does not belong to any of them. I also failed to 



Manuscript accepted 18.08.2003 



58 L. PAPP 

identify it to a family with the aid of general or regional keys. Of course, the reason is 
rather simple: all of those keys are for families where wing reduction already has been 
known. I had to use only body characteristics other than those of wings to judge its rel- 
egation among higher flies. My conclusion was that this small fly is the first apterous 
representative of the family Sciomyzidae. 

The type specimen is deposited in the Muséum d'histoire naturelle Genève, 
Switzerland (MHNG). 

TAXONOMY 

Apteromicra gen. n. 

Type species: Apteromicra parva sp. n. 

Diagnosis. Small apterous species with extremely long dorsal pre-apical setae 
on each tibia (Fig. 1) and with extremely long postocellar seta. 

Frons only slightly narrowed apically. Face nearly straight in lateral view, eyes 
ovoid. No fronto-orbital spot. Ptilinal suture normal (slightly open with a herniated part 
of ptilinal sack on the holotype of the only known species). Genal height 0.24 mm, 
height of eye 0.30 mm, ratio 0.815. Two pairs of latero- and reclinate fronto-orbitals: 
anterior pair almost completely lateroclinate, posterior pair's direction is less than 45° 
to the longitudinal body axis. Short fronto-orbital setulae from medial vertical seta to 
antennal bases: 1+2 + 3(1 short fronto-orbital between medial vertical and posterior 
fronto-orbital; 2 setulae between the two fronto-orbitals and 3 setulae between anterior 
fronto-orbital and antennal base). No true interfrontal setulae, 8 frontal setulae anterior 
to anterior fronto-orbital seta. Midfrontal area anterior to ocelli not separable from 
other parts of frons and bare. Postocellar pair extremely long, slightly divergent, in 
occipital position. No vibrissa. Clypeus narrowly U-shaped, not concealed but small 
and broadly separated from other sclerotized parts of head. Proboscis medium long, not 
strongly sclerotized (proboscis not folded). 

Thorax much reduced in volume, probably not more than the volume of head, 
since definitely narrower than head. Both pro- and metathorax relatively enlarged, 
probably due to the loss of wings, and proportional reduction of mesothorax. 
Mesonotal suture present as a black line from behind posterior notopleural seta (from 
level of posterior margin of anepisternum) as far as medially to presuturai seta; meta- 
pleural callus weak but discernible. Prosternum subtriangular, bridging the pro- 
episterna. Pro-episternum with a medium long pro-episternal seta above the base of 
fore coxa. Anepisternum and anepimeron separated by a membraneous area. Posterior 
spiracle behind level of postalar seta. One shorter postpronotal, 2 notopleural, an 
extremely strong presuturai, 2 pairs of dorsocentral and 1 pair of probably strong 
postalar setae (postalar broken off on both sides on the holotype but on its right side 
also a setal base of a supra- alar is discernible). Two pairs of scutellars, apical pair dor- 
sally directed and cruciate (Fig. 1), basal pair is very long and is in lateral position. A 
distinct pro-epimeral seta and a large katepisternal present, otherwise pleura bare. 

Hind coxa bare posterodorsally. Mid and hind femora each with a large antero- 
dorsal seta at apical 5/6. All tibiae with very long dorsal preapicals, mid tibia with 1 
anterior and posterior short apical each and 2 large thick ventroapical pairs. 



FIRST APTEROUS GENUS OF SCIOMYZIDAE 59 

Wings reduced to minute processes (tegula), this tegula with 1 minute setula. 

Abdominal spiracles 2-5 in membrane (Fig. 1). Female postabdomen simple 
(Figs 3-4). Terga and sterna 7 and 8 are small but not telescoped. Epiproct and 
hypoproct are small, cerei normal without any peculiar armature. Two pairs of globular 
spermathecae (Fig. 5). 

Etymology. The new genus does not belong to the close relatives of Pteromicra. 
Its apterous state and its relegation to the Sciomyzini are the reasons why I name it so. 

Discussion. Of course, one may question the relegation of this wingless fly to 
the family Sciomyzidae. If this is a sciomyzid, its pro-episternal seta designates its 
position definitely as a member of the tribe Sciomyzini. However, this seta is reduced 
in two genera of Sciomyzini (Atrichomelina Cresson and Pseudomelina Malloch) and 
distinctly present in three genera of Tetanocerini (Eutrichomelina Steyskal, Perilimnia 
Baker, and Shannonia Malloch). Nearly the full complement of head and thoracic setae 
found in most plesiomorphous Sciomyzidae are present in this species. The presence 
of long and thick pre-apical dorsal setae on all tibiae, which might be shorter in the 
ancestor, reduces strongly the possible number of the families, where it may belong to. 
Again, it has no vibrissal angle or vibrissae, which character excludes a number of 
families with strong dorsal preapicals on the tibiae. The shape of the prosternum 
(abridging the pro-episterna) and the 2 globular spermathecae seem to be strong 
synapomorphies, when searching for its true relatives. The male genitalia would have 
provided more evidence. However, the female postabdomen (Figs 3-4) does not show 
any true differences from that of the Sciomyzidae (cf. e.g. Rozkosny, 1998: fig. 35.36 
for Tetanocera): characteristics of the postabdomen exclude a relationship to the 
Tephritoidea (with their ovipositor) and to a number of families (or even super- 
families), where the postabdomen is telescopized. Contrary to the Heleomyzoidea (and 
allies, like the Chyromyidae), vibrissae are absent. In any case, it falls into the group 
with strong dorsal preapicals on the tibiae and without vibrissae. Contrary to the 
Dryomyzidae and allies (where the clypeus is large and prominent), its clypeus is 
narrowly U-shaped, small and broadly separated from the other sclerotized parts. 
Unlike Lauxaniidae, the postocellar setae are divergent and Apteromicra has only two 
spermathecae instead of three or four as in the Lauxaniidae). 

Diagnostic remarks. There are only two genera of Sciomyzini known in the 
Oriental region, Pherbellia Robineau-Desvoidy and Pteromicra Lioy. None of them is 
closely related to the new genus (cf. Rozkosny & Knutson, 1970). In the key Rozkosny 
(1998) published for the Palaearctic genera, one cannot find a closely related genus. 
Sciomyza Fallen with its double pre-apical setae on fore tibia (exception S. sebezhica 
Przhiboro, 2001), Ditaeniella Sack with its single fronto-orbital pair, haired pro- 
sternum and setulose anepisternum cannot be close relatives. The structure and arma- 
ture of head of Colobaea Zetterstedt are strikingly different. I think the strength and 
position (direction) of the anterior fronto-orbital, the arrangement of setae on the meso- 
notum and also the direction of the apical scutellars are features, which will be distinc- 
tive for this fly as a genus even after further studies and morphological comparisons. 

This discovery of an apterous Sciomyzidae suggests that, although few scio- 
myzids have been found in pitfall traps, such collections may indeed produce other 
new species of apterous Sciomyzidae. 



60 



L. PAPP 



Apteromicra parva sp. n. Figs 1-5 

Holotype female (MHNG): Nepal, W Thorong Pass, 4450-4350 m, sous les pierres 
[= under stones], 29. IX. 1983, leg. I. Lobi & A. Smetana, No. 19. 

Remarks. The specimen was kept in 70 % ethanol for 20 years. I judge the loss of colours 
rather moderate. The postabdomen was removed and treated with 10 % sodium-hydroxide. After 
washing in water and alcohol, it was placed in a plastic microvial with glycerine. This microvial 
contains also the spermathecae, which were removed from the abdomen during the preparation. 
The specimen itself is still preserved in alcohol. 

Description. Body length 2.37 mm, length of head 0.385 mm, length of thorax 
0.51 mm. 

Body dark grey, head greyish yellow, legs black, lateral part of abdominal 
sclerites light. 

Head as described in generic diagnosis, but in addition: 3 pairs of setulae 
between ocellar and postocellar; postgena with 2 pairs of long setae. Facial plate (prae- 
frons) comparatively narrow, less broad than both cheeks. No vibrissa, genal bristles 
not well arranged, 1 peristomal row of ca. 7 pairs (back to the postgenal seta) plus other 
6 genal setulae more dorsally on gena. Antennae black, decumbent (Fig. 2). Scape 
reduced to a short ring with minute setulae. Pedicel 0.06 mm long dorsally, with short 
setulae only. First flagellomere longer than high, broadly rounded apically. Arista sub- 
basal, 0.3 mm long, with minute dark cilia. Basal aristomere very short, hardly dis- 
cernible, second aristomere short, as broad as basal 1/3 of third aristomere. Palpus 
yellowish, normal, i.e. not thickened, with 1 long subapical ventral seta and other 5 




Fig. 1 
Apteromicra parva sp. n., holotype female (del. A. Szappanos) 



FIRST APTEROUS GENUS OF SCIOMYZIDAE 



61 




Figs 2-5 
Aptewmicra parva sp. n., holotype female. 2, head, semilateral view (del. A. Szappanos). 3, 
postabdomen, dorsal view. 4, same, lateral view. 5, spermathecae (drawn in water). Scale bars: 
0.1 mm (Fig. 5), 0.2 mm (Figs 3-4). 



shorter setulae. The specimen has the ptilinium partly exposed (most probably an indi- 
vidual abnormality, depicted also on Figs 1-2). 

Thorax dark grey, wholly microtomentose. Prosternum bare. Mesonotal setulae 
not arranged, 1 short intra-alar setula lateral to dorsocentral line, other short setulae on 
presuturai mesonotum: 1 beside postpronotal, ca. 9 within the dorsocentral lines and 5 
lateral to that lines. Notopleura and prescutellar area bare. 

Pre-apical seta on hind tibia 0.17 mm long, anterodorsal seta on hind femur 
0.19 mm. 

Abdominal setae mostly broken off on the holotype but marginal setae are not 
particularly strong (not much longer than discal ones). Abdominal spiracles 2-5 in 
membrane. Setae on epiproct asymmetrically placed. Hypoproct covered by dense 
short setulae. Cerci of normal size, simple, without any longer setulae (Fig. 3). Also a 



62 L. PAPP 

pair of internal sclerites present, supporting gonopore at their mesal ends [these are 
probably the subgenital sclerites of Rivosecchi (1992)]. Two spermathecae (Fig. 5); 
they are unequal in size, both are globular, central part involuted even in water (when 
functioning, probably their shape is actually globular). Spermathecal duct thick, thick- 
walled but not sclerotized (pigmented). 

ACKNOWLEDGEMENTS 

I would like to thank Dr Albert Szappanos for his excellent habitus figure. I am 
grateful to the former curators of the Geneva Museum (Dr Claude Besuchet and 
Dr Ivan Lobi) as well as to the present curator, Dr Bernhard Merz, for loan of the spe- 
cimen. I gratefully thank Dr Merz and the three reviewers of the manuscript for 
invaluable comments and suggestions, as well as Dr Lloyd V. Knutson for linguistic 
revision. 

REFERENCES 

Hackman, W. 1964. On reduction and loss of wings in Diptera. Notulae entomologicae 44: 

73-93. 
Papp, L. 1979. On apterous and reduced-winged forms of the families Drosophilidae, Ephydridae 

and Sphaeroceridae (Diptera). Acta zoologica hungarica 25(3-4): 357-374. 
Papp, L. & Schumann, H. 2000. 1.5. Key to families - adults (pp. 163-200). In: Papp, L. & 

Darvas, B. (eds): Contributions to a Manual of Palaearctic Diptera. Volume 1. Science 

Herald, Budapest, 978 pp. 
Rivosecchi, L. 1992. Diptera Sciomyzidae. Fauna d'Italia 30: 1-270. Bologna. 
Rozkosny, R. 1998. 3.35. Family Sciomyzidae (pp. 357-382). In: Papp, L. & Darvas, B. (eds): 

Contributions to a Manual of Palaearctic Diptera. Volume 3. Science Herald, Budapest, 

880 pp. 
Rozkosny, R. & Knutson, L. V. 1970: Taxonomy, biology, and immature stages of Palearctic 

Pteromicra, snail-killing Diptera (Sciomyzidae). Annals of the Entomological Society of 

America 63: 1434-1459. 



Revue suisse de Zoologie 111 (1): 63-76; mars 2004 



Specie nuove o poco note di Homalotini, Silusini, Bolitocharini, 
Diestotini e Autaliini della Cina e della Thailandia 
(Coleoptera, Staphylinidae)* 

Roberto PACE 

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

E-mail: pace.ent@tiscali.it 

New or little known species of Homalotini, Silusini, Bolitocharini, 
Diestotini and Autaliini from China and Thailand (Coleoptera, 
Staphylinidae). - In this paper are listed twenty-two species of the sub- 
family Aleocharinae, of which nine are described as new for the science. 
The species belong to the genera Stenomastax (1 species), Coenonica 
(3), Linoglossa (1), Neosilusa (2), Silusa (6), Leptusa (3), Neoleptusa 
(1), Phymatura (1), Pseudatheta (1), Diestota (1) and Autalia (2). Each 
new species is illustrated and compared to the related close species, on 
the basis of the form of aedeagus or spermatheca. The figures of the un- 
published aedeagus of Linoglossa chinensis are given. 

Key-words: Coleoptera - Staphylinidae - Aleocharinae - taxonomy - 
China. 

INTRODUZIONE 

Le tribù, i generi e alcune specie compresi nel presente lavoro sono già noti per 
la Cina. Sono specie fitodetriticole e fungicole, prevalentemente predatrici di larve. 
Dato che le raccolte di Aleocharinae effettuate nel lontano passato in Cina sono state 
assai rare, le specie riportate dalle recenti missioni entomologiche in Cina dell'insigne 
studioso di Staphylinidae Dr. Ales Smetana di Ottawa e del collega Michael Schtilke 
di Berlino, come prevedibile, sono in buona parte costituite da specie nuove per la 
scienza. A queste specie cinesi sono aggiunte due specie della Thailandia. 

Gli olotipi delle nuove specie sono conservati nel Museo di Storia Naturale di 
Ginevra (MHNG), nel Museo Zoologico dell'Università Humboldt di Berlino (MZB) 
e nel Museo di Storia Naturale di Londra (NHML). Paratipi e altri esemplari si con- 
servano in collezione Michael Schùlke di Berlino e del Museo Regionale di Scienze 
Naturali di Torino. 

METODO 

Il riconoscimento delle varie specie della sottofamiglia Aleocharinae, nel pre- 
sente lavoro, come in altri di questa serie, è effettuato mediante l'esame della forma 
dell'edeago e della spermateca, senza dimenticare i caratteri esterni dell'esoscheletro. 
Poiché le specie del genere Silusa Erichson, 1837, presentano un edeago di forma 



* 182° Contributo alla conoscenza delle Aleocharinae. 
Manoscritto accettato il 04.12.2002 



64 R. PACE 

piuttosto uniforme, le specie di questo genere sono distinte soprattutto per la forma del- 
l'armatura interna dell' edeago, nella sua porzione sporgente dall'orifìzio apicale. Le 
lamine laterali sono nettamente differenti da specie a specie. 

ELENCO DELLE TRIBÙ, GENERI E SPECIE 

HOMALOTINI 

Stenomastax platygaster (Kraatz, 1859) 

Homalota platygaster Kraatz, 1859: 33 

Stenomastax platygaster: Cameron, 1939: 172; Pace, 1992: 120 

19 es., Thailand, Chumphong prov., 27.III-14.IV. 1996, Pha To env., P. Prüdek leg. 

Distribuzione. Sri Lanka, India, Bengala, Vietnam, Bali, Australia. Già nota 
per la Thailandia. 

Coenonica ming Pace, 1993 

Coenonica ming Pace, 1993: 80 

2 es., China, Sichuan, Qingchong Shan, NW Chengdu, 650-700 m, 3-4. VI. 1997, leg. 
M. Schülke; 1 S, Sichuan, Gongga Shan, Moxi, 1300 m, 11. VII. 1996, leg. A. Smetana. 

Distribuzione. Specie finora nota solo dello Yunnan. 

Coenonica schuelkei sp. n. (cf. Descrizioni) 
Coenonica rugosissima sp. n. (cf. Descrizioni) 

Linoglossa (Axinocolya) chinensis Pace, 1998 Figg. 8-9 

Linoglossa (Axinocolya) chinensis Pace, 1998: 202 

1 6 , China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 2, 2800 m, 5. VII. 1998, 
leg. A. Smetana. 

Distribuzione. Il 6 di questa specie è stato raccolto nella località tipica, il 
Monte Gongga, unica località nota della specie. Viene qui data l'immagine dell' edea- 
go finora sconosciuto, figg. 8-9. 

Neosilusa ceylonica (Kraatz, 1857) 

Stenusa ceylonica Kraatz, 1857: 8 
Plagiusa ceylonica: Cameron, 1939: 167 
Neosilusa ceylonica: Pace 1984: 15; Pace 1993: 71 

12 es., China, Sichuan, Qingcheng Shan, NW Chengdu, 650-700 m, 18.V-3-4.VI. 1997, 
leg. M. Schülke. 

Distribuzione. Mascarene, India, Malesia, Cina, Giappone e Borneo. 

Neosilusa moultoni Cameron. 1920 

Neosilusa moultoni Cameron, 1920: 233; Pace, 1992: 235; Pace, 1998: 142 
Plagiusa moultoni: Bernhauer & Scheerpeltz. 1926: 540 

3 es., China, Sichuan, Qingcheng Shan, NW Chengdu, 650-700 m, 18.V-3-4.VI. 1997, 
leg. M. Schülke. 

Distribuzione. Singapore, Thailandia. Cina, Vietnam. 



SPECIE NUOVE O POCO NOTE DI HOMALOTINI ETC. 65 



SlLUSINI 

Silusa smetanai Pace, 1998 

Silusa smetanai Pace, 1998: 179 

1 ó\ China, W Sichuan, 20 Km N Sabdê, 3200 m, 13.VII.1998, leg. A. Smetana. 

Distribuzione. Finora nota solo in Cina, Gongga Shan. 

Silusa bodemeyeri (Eppelsheim, 1883) 

Leptusa bodemeyeri Eppelsheim, 1883: 252 
Silusa bodemeyeri: Pace 1989: 24 

1 3, China, Sichuan, Qingcheng Shan, Rückseite, 650-700 m, 18.V.1997, leg. M. 
Schülke. 

Distribuzione. Romania e Cina. 

Silusa shaanxiensis sp. n. (cf. Descrizioni) 
Silusa leptusoides sp. n. (cf. Descrizioni) 
Silusa schuelkei sp. n. (cf. Descrizioni) 
Silusa sichuanensis sp. n. (cf. Descrizioni) 

BOLITOCHARTM 

Leptusa (Aleteleptusa) chinensis Pace, 1997 

Leptusa (Aleteleptusa) chinensis Pace,1997:753 

32 es., China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 3, 3000 m, 
6.VIL1998, leg. A. Smetana; 1 9 , China, Sichuan, Gongga Shan, Hailuogou, in front of glacier, 
3000 m, 6.VII.1998, leg. A. Smetana; 1 9, China, N Yunnan, Xue Shan, nr. Zhongdian, 4000- 
4100 m, 23.VI.2850, leg. A. Smetana; 6 es., China, Sichuan, Emei Shan, 3000 m, 17.VII.1998, 
leg. A. Smetana. 

Distribuzione. Finora nota solo del Monte Gongga. 

Leptusa (Aleteleptusa) xiahensis Pace, 1997 

Leptusa (Aleteleptusa) xiahensis Pace, 1997: 755 

1 9, China, W Sichuan, 20 Km N Sabdê, 3200 m, 13.VII.1998, leg. A. Smetana; 4 es., 
China, W Sichuan, 15 Km W Kanding, 3250 m, 19.VII.1998, leg. A. Smetana. 

Distribuzione. Finora nota solo del Monte Gansu. 

Leptusa (Drepanoleptusa) sichuanensis Pace, 1997 

Leptusa (Drepanoleptusa) sichuanensis Pace, 1997: 757 

1 3, China, Sichuan, Gongga Shan, Hailuogou, Lake above Camp 2, 2700 m. 
4.VII.1998, leg. A. Smetana; 1 9, China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 
3, 3000 m, 6.VII.1998, leg. A. Smetana; 1 3, China, N Yunnan, Xue Shan, nr. Zhongdian, 4050 
m, 24.VI.1996, leg. A. Smetana. 

Distribuzione. Finora nota solo del Monte Gongga. 

Neoleptusa schuelkei sp. n. (cf. Descrizioni) 
Phymatura sichuanensis sp. n. (cf. Descrizioni) 



66 R. PACE 

Pseudatheta thailandensis Pace, 1992 

Pseudatheta thailandensis Pace, 1992: 239 

1 9, China, Shaanxi, Qin Ling Shan, Hua Shan ML, Valley 1200-1400 m, 118 Km E, 
Xian, sifted, 18-20.Vin.1995, leg. M. Schiilke. 

Distribuzione. Finora nota solo della Thailandia, provincia di Chiang Mai. 
Nuova per la Cina. 

DlESTOTlNI 

Diestota ming sp. n. (cf. Descrizioni) 

AUTALUNI 

Autalia rivularis (Gravenhorst, 1802) 

Aleochara rivularis Gravenhorst, 1802: 73 
Autalia rivularis : Thomson, 1859: 30 

1 es., China, Sichuan, Kanding, 2900 m, 2.VII.1996, leg. A. Smetana. 

Distribuzione. Olartica. 

Autalia limata Assing, 2001 Figg. 34-35 

Autalia microptera Pace, 1998, in litteris 
Autalia limata Assing, 2001: 912 

1 9, China, Sichuan, Emei Shan, 3000 m, 17. VII. 1996, leg. A. Smetana. 

Distribuzione. Finora nota solo del Sichuan. La forma della spermateca di 
questo esemplare coincide con quella figurata da Assing. Poiché questo autore non dà 
la figura dell'habitus e la sua descrizione è incompleta, viene qui data la figura «in 
to to» dell'esemplare. La specie è micro ttera, non atta al volo. 

DESCRIZIONI 

Coenonica schuelkei sp. n. Figg. 1-5 

Holotypus S, Thailand, Chumphong prov., 27.ni-14.IV 1996, Pha To env., P. Priidek 
leg.(MZB). 

Paratypi: 3 S â e 1 9 , stessa provenienza. 

Descrizione. Lungh. 2,9 mm. Avancorpo debolmente lucido, addome lucido. 
Corpo nero; antenne nere con i due antennomeri basali e l'undicesimo nero-bruni; 
zampe bruno-rossicce con femori neri. La punteggiatura del capo è netta, ma sul disco 
è fine e molto superficiale. La punteggiatura del pronoto è indistinta. La granulosità 
delle elitre è superficiale, quella dell'addome è fìtta. La reticolazione del capo è netta, 
quella del pronoto vigorosa e quella delle elitre e dell'addome è superficiale. Edeago 
figg. 2-3, spermateca fig. 4, sesto urotergo libero del 6 fig. 5. 

Comparazioni. La forma dell 'edeago della nuova specie è molto simile a quel- 
la dell' edeago di C. malayana Cameron, 1936, della Malaysia (holotypus S esami- 
nato). L'apice dell' edeago, infatti, è molto protratto nelle due specie, ma quello della 
nuova è ricurvo e non rettilineo come in malayana. L'armatura interna dell' edeago è 
notevolmente differente nelle due specie: più sviluppata nella nuova specie che in 
malayana. Più netti sono i caratteri differenziali esterni: corpo nero della nuova specie 
e in malayana bruno-rossiccio con elitre giallo-rossicce orlate posteriormente di bruno. 



SPECIE NUOVE O POCO NOTE DI HOMALOTINI ETC. 



67 







FlGG. 1-5 

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del S . 1-5: 
Coenonica schuelkei sp. n. 



Etimologia. La nuova specie è dedicata allo studioso di Staphylinidae Michael 
Schiilke di Berlino, per avermi affidato in esame la nuova specie. 



Coenonica rugosissima sp. n. 



Figg. 6-7 



Holotypus 9, China, SE Sichuan, Jinfo Shan, 29°01'N/107°14'E, 1750 m, 27.VI.1998, 
leg. A. Smetana, 1998 China Expedition J. Farac, D. Krâl, J. Schneide + A. Smetana (MHNG). 



68 R. PACE 

Descrizione. Lungh. 2,8 mm. Capo e pronoto opachi, elitre debolmente lucide, 
addome lucido. Corpo nero pece, con margine posteriore degli uroterghi, tranne il 
quarto libero, bruno-rossiccio; antenne brune; zampe gialle. La granulosità del capo e 
del pronoto è fìtta, quella delle elitre è fittissima. Il pronoto presenta un largo solco 
mediano sui due terzi posteriori. La reticolazione sull'addome è assente, tranne sugli 
uroterghi liberi quarto e quinto, dove le maglie di reticolazione sono molto trasverse ed 
estremamente superficiali. Spermateca fig. 7. 

Etimologia. L'aspetto rugoso dell'avancorpo dà il nome alla nuova specie. 

Comparazioni. La nuova specie per la forma della spermateca si presenta affine 
a C. mutata Pace, 1984, del Nepal. Se ne distingue per i caratteri dati nella seguente 
chiave. 

1 Corpo bruno-rossiccio, con capo e quarto urotergo libero bruni; occhi 

molto più corti delle tempie; capo coperto di granulosità grossolana; 
appendice prossimale della spermateca, rettilinea. Lungh. 2,2 mm. 

Nepal C. mutata Pace 

Corpo nero pece, con margine posteriore degli uroterghi, tranne il quarto 
libero, bruno-rossiccio; occhi più lunghi delle tempie; capo coperto di 
granulosità fine; appendice prossimale della spermateca, ricurva. Lungh. 
2,9 mm. Cina C. rugosissima sp. n. 

Silusa shaanxiensis sp. n. Figg. 10-13 

Holotypus a, China, Shaanxi, Qin Ling Shan, Hua Shan, Mt.-top, 1960-2000 m, forest, 
sifted, 19.VIII.1995, leg. M. Schülke (MZB). 

Paratypi: 4 SS e 2 9 9, stessa provenienza; 1 S, China, Shaanxi, Qin Ling Shan, 
Autoroute Km 93, S of Zhouzhi, 108 Km SW Xian, mountain forest, sifted, 1650 m, 1-2.IX. 
1995, leg. M. Schülke. 

Descrizione. Lungh. 2,8 mm. Corpo lucido e bruno-rossiccio, con uriti liberi 
quarto e tre quarti basali del quinto bruni; antenne bruno-rossicce, con i due anten- 
nomeri basali rossicci; zampe rossicce. La punteggiatura del capo e dei solchi trasversi 
basali degli uroterghi liberi quarto e quinto, è netta, quella delle elitre è profonda. La 
granulosità del pronoto e dell'addome è saliente e più fitta sulla fascia longitudinale 
mediana del pronoto, che presenta un'impressione mediana anteriore e una fossetta 
mediana trasversa basale. Assente è la reticolazione del corpo. Edeago figg. 11-12, 
spermateca fig. 13. 

Comparazioni. La nuova specie presenta una fossetta mediana posteriore del 
pronoto, come in S. cooteri Pace, 1998, pure della Cina, ma della provincia dello 
Zhejang. Tuttavia, la punteggiatura delle elitre della nuova specie è profonda, mentre 
è superficiale in cooteri. Evidenti sono le differenze nella forma dell' edeago. Apice 
spesso e corto nella nuova specie, sottile e lungo in cooteri, armatura laterale interna 
dell'edeago, sporgente esternamente, irregolarmente ovale nella nuova specie, mentre 
in cooteri è appuntita e profondamente incisa. 

Etimologia. La nuova specie prende nome dallo Shaanxi. 



SPECIE NUOVE O POCO NOTE DI HOMALOTINI ETC. 



69 




Figo. 6-13 
Habitus, spermateca ed edeago in visione laterale e ventrale. 6-7: Coenonica rugosissima sp. n.; 
8-9: Linoglossa chinensis Pace; 10-13: Silusa shaanxiensis sp. n. 



Silusa leptusoides sp. n. 



Figg. 14-17 



Holotypus ó\ China, W Sichuan, 20 Km N Sabdê, 29°35'N/102°23'E, 3200 m, 
13.VÜ.1998, leg. A. Smetana, 1998 China Expedition J. Farac, D. Krâl, J. Schneide + A. 
Smetana (MHNG). 



70 R- pace 



Paratypi: 1 6 e 2 9 9, stessa provenienza, ma anche in data lie 14.VII.1998; 1 S e 1 
9, China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 3, 3000 m, 6.VH.1998, leg. A. 
Smetana. 

Descrizione. Lungh. 2,8 mm. Corpo lucido e bruno, con margine posteriore dei 
tre uriti basali, bruno-rossiccio; antenne e zampe rossicce. La punteggiatura ombelicata 
del capo è superficiale. La granulosità del pronoto è distinta, quella delle elitre è 
saliente. Assente è la reticolazione del corpo. Spermateca fig. 15, edeago figg. 16-17. 

Etimologia. Il nome della nuova specie significa «dall'aspetto di Leptusa», 
perché a prima vista sembra una specie di Leptusa. 

Comparazioni. Per la forma dell'armatura interna dell'edeago, la nuova specie 
si presenta affine a S. cooteri Pace, 1998, pure della Cina, ma della provincia dello 
Zhejang. Se ne distingue per i caratteri dati nella seguente chiave. 

1 Quarto antennomero nettamente trasverso; occhi molto più corti delle 

tempie; elitre lunghe quanto il pronoto; piastre sporgenti dell'armatura 
interna dell'edeago poco profondamente incise e divise in un lobo 

stretto e uno tozzo. Lungh. 2,8 mm. Zhejang S. cooteri Pace 

Quarto antennomero debolmente trasverso; occhi poco più corti delle 
tempie; elitre più lunghe del pronoto; piastre sporgenti dell'armatura 
interna dell'edeago molto profondamente incise e divise in un lobo largo 
e uno stretto e lungo. Lungh. 2,8 mm. Sichuan S. leptusoides sp. n. 

Silusa schuelkei sp. n. Figg- 18-21 

Holotypus 6, China, W Sichuan, Erlang Shan, E Sutao-Kiao Pass, 3000 m, 19.V.1997, 
leg. M. Schülke (MZB). 

Paratypi: 23 es., stessa provenienza. 

Descrizione. Lungh. 2,1 mm. Corpo lucido e bruno-rossiccio, con pigidio 
rossiccio e quarto urite libero bruno; antenne e zampe rossicce. La punteggiatura del 
capo e la granulosità del pronoto e dell'addome sono distinte. La granulosità delle 
elitre è saliente. La reticolazione del capo, del pronoto e delle elitre è netta e quella 
dell'addome è distinta. Edeago figg. 19-20, spermateca fig. 21. 

Comparazioni. La nuova specie, per avere le elitre più corte del pronoto, è 
nettamente distinta da tutte le specie del genere Silusa della Cina e delle regioni vicine. 
In base alla forma della parte sporgente dell'armatura interna dell'edeago, la nuova 
specie si presenta affine a S. cooteri Pace, 1998 e a S. leptusoides sp. n, sopra descritta. 

Etimologia. La nuova specie è dedicata al suo abile raccoglitore, lo studioso di 
Staphylinidae Michael Schülke di Berlino. 

Silusa sichuanensis sp. n. Figg- 22-26 

Holotypus ó\ China, W Sichuan, 20 km N Sabdê, 3200 m, 29°35'N/102°23'E, 
15.VH.1998, leg. A. Smetana, 1998 China Expedition J. Farac, D. Krâl, J. Schneide + A. 
Smetana (MHNG). 

Paratypi: 8 es., stessa provenienza, ma anche in data 11-13-14.VH.1998. 

Descrizione. Lungh. 2,9 mm. Corpo lucido e bruno-rossiccio, con capo, lato 
esterni delle elitre, uriti liberi terzo, quarto e base del quinto bruni, uriti basali e pigidio 
giallo-rossicci; antenne brune, con i due antennomeri basali e la base del terzo rossicci 



SPECIE NUOVE O POCO NOTE DI HOMALOTINI ETC. 



71 




Figo. 14-21 
Habitus, spermateca ed edeago in visione laterale e ventrale. 14-17: Silusa leptusoides sp. n. ; 
18-21: Silusa schuelkei sp. n. 

e metà apicale dell'undicesimo giallo-rossiccia; zampe giallo-rossicce. La pun- 
teggiatura del capo è poco profonda. La granulosità del pronoto è superficiale, quella 
delle elitre è poco saliente. La reticolazione del capo è distinta, quella del pronoto è 
netta sulla fascia longitudinale mediana, ai lati è estremamente superficiale. La retico- 



72 



R. PACE 




Figg. 22-27 
Habitus, edeago in visione laterale e ventrale e spermateca a ingrandimento minore e maggiore. 
22-26: Silusa sìchuanensis sp. n.; 27: Neoleptusa schuelkeì sp. n. 



lazione delle elitre e dei quattro uroterghi basali è assente, quella quinto urotergo libero 
è molto trasversa ed estremamente superficiale, quella sul sesto urotergo libero è pure 
molto trasversa, ma distinta. Edeago figg. 23-24, spermateca fig. 25-26. 



SPECIE NUOVE O POCO NOTE DI HOMALOTTNI ETC. 73 

Comparazioni. L'edeago della nuova specie è simile a quello di S. bodemeyeri 
(Eppelsheim, 1883), presente anche in Cina. Se ne distingue per la sua armatura inter- 
na molto più robusta di quella della specie in comparazione, con parte apicale poco 
dilatata (molto dilatata in bodemeyeri). 

Neoleptusa schuelkei sp. n. Figg. 27-29 

Holotypus ó\ China, Shaanxi, Qin Ling Shan, Hua Shan, Mt.-S top, 1950-2000 m, 
19.VHI.1995, leg. A. Pütz (MZB). 

Descrizione. Lung. 2,8 mm. Corpo lucido e bruno, con elitre, margine 
posteriore dei tre uroterghi basali e pigidio rossicci; antenne brune, con i due anten- 
nomeri basali e la base del terzo rossicci; zampe giallo-rossicce. La punteggiatura del- 
l'avancorpo è distinta. La reticolazione del capo e del pronoto è molto svanita, quella 
delle elitre e dell'addome è superficiale. Edeago figg. 28-29. 

Comparazioni. Per la forma dell' edeago, la nuova specie è più simile a N. ru- 
gosa Pace, 1985, del Nepal, che a N. brunnea Cameron, 1939, dell'India (tipi esa- 
minati), che mostra una lamina ventrale dell'edeago. In visione ventrale l'apice 
dell'edeago della nuova specie è acutissimo, mentre quello di rugosa è assai largo. 

Etimologia. La nuova specie è dedicata allo studioso di Staphylinidae Michael 
Schùlke di Berlino, che me l'ha affidata in esame. 

Phymatura sichuanensis sp. n. Figg. 30-31 

Holotypus 9 , China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 2, 2800 m, 
29°35'N /102°00'E, 5.VII.1998, leg. A. Smetana, 1998 China Expedition J. Farac, D. Krâl, J. 
Schneide + A. Smetana (MHNG). 

Descrizione. Lungh. 2,9 mm. Corpo debolmente lucido e bruno, con pronoto 
bruno-rossiccio dal margine laterale e posteriore rossicci, con base delle elitre giallo- 
bruna e con i due uriti basali liberi e il pigidio rossicci; antenne rossicce con i tre 
antennomeri basali giallo-rossicci; zampe giallo-rossicce con femori bruni. La granu- 
losità del capo è fine, quella del pronoto è distinta. La punteggiatura delle elitre è ben 
visibile. La reticolazione dell'avancorpo è distinta, quella dell'addome è molto super- 
ficiale. Spermateca fig. 31. 

Comparazioni. Per la forma della spermateca, la nuova specie si presenta affine 
a P. smetanai Pace, 1998, pure delle Cina. Se ne distingue per gli occhi più sviluppati, 
per la presenza di una fossetta mediana posteriore del pronoto e per la profondissima 
introflessione apicale del bulbo distale della spermateca. 

Etimologia. La nuova specie prende nome dal Sichuan. 

Diestota ming sp. n. Figg. 32-33 

Holotypus 9, China, Bowring, 63-47, 541, 16.VI.1950 (NHML). 

Descrizione. Lungh. 2,1 mm. Corpo lucido e bruno-rossiccio; antenne brune 
con i tre antennomeri basali rossicci; zampe giallo-rossicce. La punteggiatura del capo 
è ombelicata e netta e sta solo sulla metà posteriore, in avanti la superficie è coperta 
di granulosità. Il pronoto presenta una netta granulosità, in avanti sta un debole e largo 
solco mediano, all' indietro due solchi uniti posteriormente da uno trasverso. Le elitre 



74 



R. PACE 




Figo. 28-33 
Edeago in visione laterale e ventrale, habitus e spermateca. 28-29: Neoleptusa schuelkei sp. n.; 
30-31: Phymatura sichuanensis sp. n.; 32-33: Diestota ming sp. n. 



presentano alcuni punti profondi sparsi su un fondo superficialmente punteggiato. I due 
uriti basali mostrano file trasverse di granuli. Solo il quarto urotergo libero presenta 
carene basali. Assente è la reticolazione del corpo. Spermateca fig. 33. 



SPECIE NUOVE O POCO NOTE DI HOMALOTINI ETC. 



75 




Figo. 34-35 
Habitus ed edeago. 34-35: Autalia limata Assing, 2001 (= Autalia microptera Pace, 1998, in 
litteris). 

Comparazioni. Per la forma della spermateca e per alcuni caratteri esterni, la 
nuova specie si presenta affine a D. castanea (Cameron, 1920), di Singapore (tipo 9 
esaminato). La parte mediana della spermateca della nuova specie è avvolta a matassa 
con ampie spire, mentre la porzione mediana della spermateca di castanea è avvolta 
a matassa con spire strette. Il pronoto della nuova specie è sinuato davanti agli angoli 
posteriori, mentre non lo è quello di castanea. 

Etimologia. La nuova specie prende nome dalla dinastia cinese Ming, che 
regnò dal 1368 al 1644. 



RINGRAZIAMENTI 

Per avermi affidato in studio il materiale oggetto del presente lavoro rivolgo i 
miei più cordiali ringraziamenti ai colleghi Dr. Ales Smetana di Ottawa, Michael 
Schiilke di Berlino e Guillaume de Rougemont di Londra. Per il prestito di tipi 
ringrazio il Dr. P. M. Hammond del Museo di Storia Naturale di Londra, il Dr. Lothar 
Zerche del D.E.I di Ebers walde, e il Dr. Manfred Uhlig del Museo Zoologico 
dell'Università Humboldt di Berlino. 



76 R- pace 



BIBLIOGRAFIA 



AssiNG, V. 2001. Review of Palaearctic Autalia Leach. IV. New species and additional records 

(Coleoptera, Staphylinidae, Aleocharinae). Revue suisse de Zoologie 108 (4): 911-917. 
Bernhauer, M. & Scheerpeltz, 0. 1926. Coleopterorum Catalogus, pars 82, Staphylinidae VI: 

499-988, Berlin. 
Cameron, M. 1920. New species of Staphylinidae from Singapore, Part in. Transactions of the 

Entomological Society of London 1920: 212-284. 
Cameron, M.1939. The Fauna of British India, including Ceylon and Burma. Coleoptera, 

Staphylinidae, 4 (Part 1). London, 410 pp. 
Eppelshem, E. 1883. Neue Staphyliniden der Österreichischen-ungarischen Monarchie und der 

angrenzenden Länder. Wiener Entomologische Zeitschrift 2: 251-255. 
Gravenhorst, J. L. C. 1802. Coleoptera Microptera Brunsvicensia nee non exoticorum quotquot 

exstant in collectionibus entomologorum Brunsvicensium in genera, familias et species 

distribuit. Brunsvigae: Carolus Reichard, 206 pp. 
Kraatz, G. 1857. Naturgeschichte der Insekten Deutschlands. Abteilung 1, Coleoptera. Vol. 2, 

Staphylinii. Nicolai, Berlin, 376 pp. 
Kraatz, G 1859. Die Staphyliniden-Fauna von Ostindien, insbesonders der Insel Ceylan. 

Archiv für Naturgeschichte 25: 1-196. 
Pace, R. 1984a. Aleocharinae delle Mascarene, parte I: tribù Myllaenini, Pronomaeini, Oligotini 

e Bolitocharini (Coleoptera Staphylinidae) (XLV Contributo alla conoscenza delle 

Aleocharinae). Revue suisse de Zoologie 91: 3-36, 159 figg. 
Pace, R. 1984b. Aleocharinae della Thailandia e della Birmania riportate da G. de Rougemont 

(Coleoptera Staphylinidae). (LIX Contributo alla conoscenza delle Aleocharinae). 

Bollettino del Museo civico di Storia naturale di Verona 11: 427-468, 139 figg. 
Pace, R. 1989. Monografia del genere Leptusa Kraatz (Coleoptera Staphylinidae) (LXXV 

Contributo alla conoscenza delle Aleocharinae). Memorie del Museo civico di Storia 

naturale di Verona (115), A 8: 1-307, 1696 figg. 
Pace, R. 1992a. Aleocharinae del Vietnam (Coleoptera, Staphylinidae). Nouvelle Revue 

d'Entomologie (N. S.) 9: 119-129. 
Pace, R. 1992b. Aleocharinae della Thailandia (Coleoptera Staphylinidae) (XCV Contributo al- 
la conoscenza delle Aleocharinae). Bollettino del Museo civico di Storia naturale di 

Verona 16: 227-268. 
Pace, R. 1993. Aleocharinae della Cina (Coleoptera Staphylinidae). Bollettino del Museo civico 

di Storia naturale di Verona 17: 69-126. 
Pace, R. 1997. Specie del genere Leptusa in Cina. Monografia del Genere Leptusa Kraatz: 

Supplemento VII (Coleoptera, Staphylinidae) (137° Contributo alla conoscenza delle 

Aleocharinae). Revue suisse de Zoologie 104: 751-760, 22 figg. 
Pace, R. 1998. Aleocharinae della Cina: Parte I (Coleoptera, Staphylinidae). Revue suisse de 

Zoologie 105: 139-220, 234 figg. 
Thomson, C. G. 1859. Skandinaviens Coleoptera, synoptikt bearbetade, I. Berlingska Bok- 

tryckeriet, Lund, 290 pp. 



Revue suisse de Zoologie 111 (1): 77-101; mars 2004 



Pauropoda (Myriapoda) from Vietnam 

(Pauropoda and Symphyla of the Geneva Museum XIII) 

Ulf SCHELLER 

Häggeboholm, Häggesled, S-53194 Järpas, Sweden. 



Pauropoda (Myriapoda) from Vietnam (Pauropoda and Symphyla of 
the Geneva Museum XIII). - Eight species of Pauropoda are reported from 
and near limestone hills in south Vietnam. Seven species from four genera 
are new to science and are described here: Allopauropus (D.) leptotarsus 
sp. n., A. (D.) linguatulus sp. n., A. (D.) barbatulus sp. n., A. (D.) absimilis 
sp. n., Pauropus asiaticus sp. n., Samarangopus campanulatus sp. n. and 
Sphaeropauropus lecongkieti sp. n. 

Key- words: Myriapoda - Pauropoda - taxonomy - new species - Vietnam - 
biogeography - soil fauna - limestone. 



INTRODUCTION AND BACKGROUND 

As far as I know only one pauropod species has previously been recorded from 
Vietnam, viz. Pauropus dawydoffi Remy. A single specimen was reported (Remy, 
1933) from central Vietnam (Dalat, in the tableland Lang Biang, alt. 1500 m a.s.). The 
species has not been found again either in Vietnam or elsewhere. 

The ecosystem of the limestone hills in the Hon Chong area is seriously 
threatened by irreversible damages caused by intensive limestone exploitation linked 
to the recent development of large cement plants. Though these hills and their fauna 
and flora certainly have a biological originality of high value they were till recently 
virtually unknown to science. The first results of a soil and cave fauna survey have 
shown that several endemic species of Collembola and Coleoptera are living there 
(Deuve, 1996; Deharveng & Bedos, 1995,1996; Bedos & Deharveng, 2000). 

Now data on Pauropoda can be added. Though it was not the primary aim of 
collecting to obtain pauropods, Drs Deharveng's and Bedos' material is of greatest 
value. The present study shows that the pauropods too have developed several endemic 
species there. Among 8 species reported, no less than 7 are new to science and one can 
hardly believe that they have a much wider distribution outside the area of these lime- 
stone hills. No comparable ecosystems exist neither in the neighbourhood nor else- 
where in Vietnam, where the next significant karstic area is located 750 km north-east 
of Hon Chong. Scattered limestone hills of the same type can be found 30-40 km 
further north near the border in Ha Tien and in Cambodia (near Kompong Trach). 
Calcareous areas with similar climatic conditions also exist in Thailand, but they are 



Manuscript accepted 30.10.2003 



78 U. SCHELLER 

lying in a different geographic area and support a largely different wildlife. The Hon 
Chong limestone hills are small and isolated, and evidence is growing that their litter 
and soil fauna contains endemic taxa of great interest. 

MATERIAL 

In 1995, 2000 and 2001 Drs Louis Deharveng and Anne Bedos, Université Paul 
Sabatier, Toulouse (now Muséum National d'Histoire Naturelle, Paris), assisted by 
Professor Le Cong Kiet from the University of Ho Chi Minn City (Department of 
Botany and Ecology), made Berlese extractions of litter and soil samples in south- 
western Vietnam. Their interest was focused on the Ha Tien - Hon Chong limestone 
hills in the Kien Giang Province. From the collected material 45 specimens of 
Pauropoda were sorted out, among which 8 species could be identified, 7 of them as 
new to science. 5 species belong to the Pauropodidae (genera Allopauropus and 
Pauropus) and 3 species to the Eurypauropodidae (genera Samarangopus and Sphae- 
ropauropus). The species new to science are described below: Allopauropus (D.) 
leptotarsus sp. n., A. (D.) linguatulus sp. n., A. (D.) barbatulus sp. n., A. (D.) absimilis 
sp. n., Pauropus asiaticus sp. n., Samarangopus campanulatus sp. n. and Sphaeropau- 
ropus lecongkieti sp. n. 

Unless stated otherwise, all material was collected by Drs Louis Deharveng and 
Anne Bedos. The material is deposited in the collections of the Department of 
Arthropods and Entomology I, Natural History Museum of Geneva. 

ABBREVIATIONS AND MEASUREMENTS 

Abbreviations: ad. ..., subad. ... and juv. ... = an adult, a subadult or a juvenile 
specimen with the number of pairs of legs indicated. 

Measurements: length of the body in mm and range of variation in adult 
paratypes given in brackets. Indication of absolute lengths are generally used only in 
the description of Eurypauropodidae and always applicated with urn. Otherwise the 
text refer the reader to relative lengths. 

SYSTEMATICS 

Order Tetramerocerata 
Pauropodedae 

One described species in the Pauropodidae, Pauropus dawydoffi Remy, was 
previously known from Vietnam (Remy, 1933). Five additional species, all new to 
science, with a total of 19 specimens, are here added. The high number of undescribed 
species in comparison with the low number of collected specimens indicates a very di- 
versified fauna. These 6 species belong to the genera Allopauropus and Pauropus, 2 of 
the most widespread genera in Pauropoda. Future collecting will certainly increase 
considerably the number of species in these genera, particularly in Allopauropus, but I 
suppose that not only many species will be discovered there, but also some additional 
genera to the list of Vietnam, e.g. Scleropauropus, Stylopauropus, Hemipauropus, 
Rabaudauropus, Cauvetauropus. Thus, those 6 species now known from Vietnam are 
only a small part of an expected rich fauna of Pauropodidae. 



PAUROPODA FROM VIETNAM 79 

In the studied collection another Pauropodidae species was found, which, how- 
ever, could not be described. A single ad. 9(a), of a species close to Allopauropus (D.) 
chichinii Remy, was collected from a limestone hill at Mo So near Hon Chong. 

Genus Allopauropus Silvestri, 1902 

Subgenus Decapauropus Remy, 1957 (Remy, 1957a) 

1. Allopauropus (D.) leptotarsus sp. n. Figs 1-11 

Type material. Holotype: ad. 9( 9 ), Vietnam, Ken Giang Province, Kien Luong, Hon 
Chong, Nui Binh Tri, secondary forest, Berlese extraction, 19.1.2000 (Loc. VIET-861). Para- 
types: 2 ad. 9(9), same data as holotype (Loc. VIET-858). 

Other material. Vietnam, Ken Giang Province, Kien Luong, Hon Chong, Nui Son Cha, 
under bush on limestone, litter, Berlese extraction, 1 ad. 9(9), 18.1.2000 (Loc. VIET-840). 

Total number. 4 specimens. 

Diagnosis. A. (D.) leptotarsus sp. n. is well delimited from all other species of 
Decapauropus by the following combination of distinctive characters: anal plate rhom- 
bic, with two straight parallel posterior appendages close to each other; pygidial 
tergum subrectangular, with a small median lobe above anal plate; tarsi of posterior 
legs very slender and setae of the pygidial tergum long, pointed, glabrous and directed 
posteriorly. From the above characters A. (D.) leptotarsus sp. n. seems to be close to 
species in the subgenus Allopauropus s. str., but the absence of the pygidial setae b 3 
assigns it to the subgenus Decapauropus. 

Etymology. From Greek leptos = thin, fine, slender (referring to the tarsi of the 
last pair of legs). 

Description 

Length. (0.88-) 0.92(-0.94) mm. 

Head. Tergal setae mostly lacking, those present showing thin cylindrical or 
subcylindrical setae of medium length or fairly long ones. Relative lengths of setae 
(paratype specimen), 1 st row: a x = 10, a 2 = 9; 2 nd row: a 1 = 10, a 2 = 13, a 3 = 11; 3 rd row: 
a l = 9, a 2 = ?; 4 th row: a x = a 4 = 10, a 2 = a 3 = 17; lateral group of setae: l 1 = l 3 = 15, U 
= ?. Length of temporal organs somewhat shorter than shortest interdistance. Head 
cuticle glabrous. 

Antennae. Segment 4 with 5 setae, all thin, cylindrical, distally tapering, densely 
striate. Relative lengths of setae: p = 100, /?' = (67-)73(-75), p" = (48-)50(-52), p'" = 
(12-)13, r= 13(-14). Tergal branch t thin, slender, (5.2-)6.0(-6.1) times as long as great- 
est diameter and (1.3-)1.6(-1.7) times as long as sternal branch s, which is 2.2(-3.1) 
times as long as its greatest diameter; anterior corner strongly truncate. Seta q similar 
to seta r of 4 th segment, with high insertion point, almost straight, (0.5-)0.6 of length 
of s. Relative lengths of flagella (base segments included) and base segments alone: 
F x = 100, bs 1 = 4;F 2 = 30(-33), bs 2 = 2; F 3 = (78-)80(-81), bs 3 = 4. F x 3.4(-3.5) times 
as long as t, F 2 and F 3 1.1(-1.2) and (2.9-)3.0(-3.1) times as long as s respectively. 
Distal calyces of F l conical, those of F 2 and F 3 very small and not examined in detail. 
Distal part of flagella axes somewhat widened but only just below calyx. Globulus g 
only very little longer than wide, capsule flattened; width of g 0.8 of greatest diameter 
of t. Antennae glabrous. 



80 



U. SCHELLER 




Figs 1-11 
Allopauropus (D.) leptotarsus sp. n., holotype, ad. 9(9). 1, head, median and right part, tergal 
view; 2, right antenna, tergal view; 3, collum segment, median and left part, sternal view; 4, ter- 
gile VI, posterior part; 5, seta on coxa of leg 9; 6, seta on coxa of leg 8: 7, seta on trochanter of 
leg 8; 8, tarsus of leg 9; 9, pygidial tergum, tergal view; 10, pygidium, posteromedian part, 
sternal view; 11, anal plate, lateral view. Scale a: Figs 4-8; b: Figs 1-3, 9-11. 



PAUROPODA FROM VIETNAM 8 1 

Trunk. Setae of collum segment furcate. Primary branch thick, cylindrical, 
densely annulate; secondary branch rudimentary and glabrous. Sublateral seta 2.7C-2.8) 
times as long as submedian one; sternite process in anterior part very narrowly rod- 
shaped and without apical incision. Appendages much wider in proximal than in distal 
half, distal caps hemispherical. Process and appendages glabrous. 

Setae on tergite I as on head, on II similar but somewhat tapering, on VI pointed 
and glabrous. 4+4 setae on tergite I, 6+6 on II-IV, 6+4 on V and 4+2 on VI. Submedian 
posterior setae on tergite VI (0.8-)0.9 of their interdistance. 

Bothriotricha. Relative lengths of bothriotricha: T x = 100, T 2 = (110-)113(-114), 
T 3 =108(-119), T 4 = 7(124-128), T 5 = 168(-181). Axes thin, simple, straight, those of T 3 
only a little thicker than the others. Pubescence very short depressed-oblique. 

Legs. Setae on coxa and trochanter of leg 9 similar to each other, furcate, 
branches cylindrical, striate, blunt; secondary branch somewhat shorter than primary 
one. These setae on legs 1-8 with rudimentary glabrous secondary branches, setae on 
coxa thicker than those on trochanter. Tarsus of leg 9 very slender, (6.2-)7.3(-7.4) times 
as long as greatest diameter, distal half subcylindrical. Proximal seta thin, pointed, with 
short oblique pubescence, its length 0.4 of length of tarsus and (4.0-)4.3 times as long 
as distal seta; the latter cylindrical, blunt, with short oblique pubescence. Cuticle of 
tarsus almost glabrous. 

Pygidium. Tergum. Almost rectangular with small semicircular lobe between st 
just above anal plate. Relative lengths of setae: a x = 10, a 2 = (10-)11, a 3 = (22-)24, st 
= 2(-3). All these setae almost straight, tapering, pointed, glabrous; a x and a 2 somewhat 
converging, a 3 somewhat diverging, st strongly converging. Distance a x - a x 0.8 of 
length of a x \ distance a x - a 2 3.0 times longer than distance a 2 - a 3 ; distance st - st 2.3(- 
2.4) times as long as st and 0.7 of distance a x - a x . Cuticle glabrous. 

Sternum. Posterior margin between b x indented and with a broad triangular lobe 
projecting backwards below anal plate. Relative lengths of setae (a x = 10): b x = 
12(-13), b 2 = 5. These setae thin, tapering, distally striate; b x (1.2-)1.3 times as long as 
interdistance, b 2 l.l times as long as distance b x - b 2 . 

Anal plate directed upwards-backwards, rhombic, somewhat longer than wide, 
lateral corners rounded, 2 thin posteromedian, parallel appendages close to each other. 
The latter tapering, directed posteriorly, their length 0.7 of length of plate. Plate and 
appendages glabrous. 

2. Allopauropus (D.) linguatulus sp. n. Figs 12-21 

Type material. Holotype: ad. 9( 9 ), Vietnam, Kien Giang Province, Kien Luong, Hon 
Chong, Nui Hang Tien, under bush on limestone, soil sample, Berlese extraction, 19.1.2000 (Loc. 
VIET-864). Paratypes: 3 ad. 9(1 o\ 2?), 1 juv. 3, same data as holotype. Kien Giang Province, 
Kien Luong, Hon Chong, Nui Son Cha, under limestone rock shelter, soil sample, Berlese 
extraction, 1 ad. 9(a), 18.1.2001 (Loc. VIET-840). 

Total number. 6 specimens. 

Diagnosis. Only a few species in Allopauropus have a linguiform anal plate 
without appendages. Among them, A. linguatulus sp. n. may be closest to A. ligulosus 
Hagino from Japan (Hagino, 1991), A. andriai Remy and A. palifer Remy, both from 
Madagascar (Remy, 1956a). Distinctive characters in relation to A. ligulosus are the 
shape of the bothriotricha (e.g. T 3 with thin subcylindrical axis in A. linguatulus sp. n., 



82 



U. SCHELLER 




Figs 12-21 
Allopauropus (D.) linguatulus sp. n.: 12-16, 18-21, holotype, ad. 9(9), 17, paratype, ad. 9(a). 
12, head, median and right part, tergal view; 13, right antenna, tergal view; 14, collum segment, 
median and left part, sternal view; 15, tergite VI, posteromedian part; 16, T 3 ; 17, genital papillae 
and seta on coxa of leg 2, anterior view; 18, seta on coxa of leg 9; 19, tarsus of leg 9; 20, 
pygidium, posterior part, tergal view; 21, anal plate, lateral view. Scale a: Fig. 16; b: Figs 12, 14, 
15, 17-20; c: Figs 13, 21. 



proximal 2/3 of axis club-shaped in A. ligulosus) and the st (thin and tapering in A. lin- 
guatulus sp. n., not davate in A. ligulosus). A. linguatulus, sp. n. is distinguished from 
A. andriai Remy by the shape of the bothriotrix T 3 (axis thin and tapering in A. lin- 
guatulus sp. n., with distal end widened in A. andriai) and by the dissimilar margins of 
the anal plate (glabrous, not with fringe of distinct pubescence hairs). The new species 
is well delimited from A. palifer Remy too. 



PAUROPODA FROM VIETNAM 83 

The anal plate has a glabrous margin in A. linguatulus sp. n., but a fringe of dis- 
tinct pubescence hairs in A. palifer and the st are thin and cylindrical, not davate. 
Moreover is the shape of the setae a x on the pygidial tergum thin and tapering in A. lin- 
guatulus sp. n., and short and davate in A. palifer. 

Etymology. From Latin lingua = tongue (referring to the tongue-like shape of 
the anal plate). 

Description 

Length. (0.65-)0.89 mm. 

Head. Tergal setae of medium length, median ones subcylindrical, annulate, 
blunt; a 3 in 2 nd row and lateral group of setae striate, pointed. Relative lengths of setae, 
1st r0 w: a x = 10, a 2 = 10(-11); 2°* row: a i = \\,a 2 = 23(-25), a 3 = 17(-20); 4* row: a x 
= (13-) 16, a 2 = 14(-19), a 3 = ?(19), a A = 17; lateral group of setae: l x = 27, l 2 = 20, l 3 = 
19. Temporal organs about 0.8 of shortest interdistance; small aperture at posterior 
margin. Head cuticle glabrous. 

Antennae. Segment 4 with 5 setae, all but r and/?"' subcylindrical, annulate; r 
straight, thin, cylindrical, striate; /?"' a rudimentary knob only. Relative length of setae 
(paratype): p = 100, /?' = 36, p" = 40, p'" = 1, r = 68. Tergal branch t somewhat 
fusiform, (2.5-)2.6 times as long as greatest diameter and (1.0-)1.2 times as long as 
sternal branch s, which is 1.7(-1.8) times as long as greatest diameter; anterior corner 
distinctly truncate. Seta q cylindrical, annulate-striate, pointed, (1.1-)1.3(-1.5) times as 
long as s. Relative lengths of flagella (base segments included) and base segments 
alone: F l = 100, bs 1 = 6; F 2 = 36(-37), bs 2 = 4;F 3 = (85-)88, bs 3 = 6. F 3 thinner than 
F v F 2 thinner than F 3 . F x 5.3 times as long as t, F 2 and F 3 (1.9-)2.3 and 4.7 times as 
long as s, respectively. Distal calyces somewhat flattened. Distal part of flagella axes 
only slightly widened. Globulus g 1.2(-1.3) times as long as greatest diameter, with 
(lO-)llbracts, capsule flattened; width of g 0.9(-1.0) of greatest diameter of t. Antennae 
glabrous. 

Trunk. Setae of collum segment simple or with minute rudimentary secondary 
branch. Primary branch very thin, striate. Sublateral seta (3.3-)3.4 times as long as sub- 
median one; sternite process anteriorly narrow, with small incision. Appendages 
obliquely conical, caps flat; process and appendages almost glabrous. 

Tergal setae similar to median setae on tergal side of head; 4+4 setae on tergite 
I, 6+6 on II-IV, 6+4 on V and 4+2 on VI. Submedian posterior setae on VI almost 0.4 
of their interdistance. 

Bothriotricha. Relative lengths of bothriotricha: 7, = 100, T 2 = 83(-88), T 3 
=117(-123), T 4 = 126(-133), T 5 = ?(193). Their axes thin, simple, straight, those of T 3 
only a little thicker than the others. Pubescence almost erect, strongest on T 3 . 

Genital papillae (paratype). Base wide, distally strongly tapering, distal half 
narrow, subcylindrical; they are 1.8 times as long as widest part, seta 0.4 of length of 
papilla. Seta on coxa of leg 2 similar to other coxal setae on anterior legs. 

Legs. Setae on coxa and trochanter of leg 9 similar to each other, furcate, 
branches subcylindrical, striate-annulate, blunt; secondary branch somewhat shorter 
and thinner than primary one. These setae seem to be simple on legs 1-8 and without 
rudimentary secondary branch. Tarsus of leg 9 slender, tapering, somewhat curved, 



84 U. SCHELLER 

(3.5-)3.7 times as long as greatest diameter. Proximal seta thin, cylindrical, striate, 
blunt, length 0.2(-0.3) of length of tarsus and 1.2(-1.3) times as long as distal seta; the 
latter cylindrical, striate, blunt. Cuticle of tarsus glabrous. 

Pygidium. Tergum. Posterior margin rounded, with triangular lobe between st 
just above anal plate. Relative lengths of setae: a x = 10, a 2 = 6, a 3 = ?(8-9), st = 4(-5). 
fl l5 a 2 and a 3 almost straight, thin, tapering, striate distally; st glabrous, tapering, curved 
inwards and converging; a 2 thinnest and inserted almost straight anterior of a 3 . 
Distance a x - a x 0.9 of length of a 7 ; distance a x - a 2 twice longer than distance a 2 - a 3 ; 
distance st - st 2.6(-3.1) times as long as st and 0.8(-0.9) of distance a x - a v Cuticle 
glabrous. 

Sternum. Posterior margin between b x with broad and low lobe below anal plate. 
Relative lengths of setae (a x = 10): b x = 17(-18), b 2 = 8(-9). These setae thin, tapering, 
striate distally. b x 1.1 (-1.2) times as long as their interdistance, b 2 0.8 of distance 
b x - b 2 . 

Anal plate directed backwards-downwards, narrowest anteriorly, linguiform, 
1.1 times as long as wide, glabrous; no appendages. 

3. Allopauropus (D.) barbatulus sp. n. Figs 22-31 

Type material: Holotype: ad. 9( 2 ), Vietnam, Kien Giang Province, Kien Luong, Hon 
Chong, Nui Binh Tri, secondary forest in area without limestone, soil sample, Berlese extraction, 
19.1.2000 (Loc. VIET-853). 

Total number. 1 specimen. 

Diagnosis. A. barbatulus sp. n. is well defined by the following combination of 
characters: 1. anal plate almost cordate, with two short-stalked and ball-shaped 
appendages, one at each posterolateral corner; 2. distal seta on tarsus of leg 9 much 
longer than proximal seta; 3. pygidial tergum with thin subsimilar a-setae and short 
davate st. This new species may be related to A. rhopalophorus Remy from Algeria 
(Remy, 1947) by the shape of the anal plate and some of the setae of the pygidial 
tergum. The two species are easily distinguishable by the shape of the pygidial setae 
a± (long and thin in A. barbatulus sp. n., short and blunt in A. rhopalophorus Remy) 
and by the shape of the appendages of the anal plate (ball-shaped and with short pu- 
bescence, versus not subcylindrical with a few long pubescence hairs only). More 
distant relationships presumably exist with the following species, e.g., A. compatruelis 
Remy & Rollet from Madagascar (Remy & Rollet, 1960), A. scitulus Remy from 
Madagascar too (Remy, 1956a) and A. chartoni Remy from Réunion (Remy, 1956b). 

Etymology. Diminutive of Latin barbatus = bearded (referring to the hairy anal 
plate). 

Description 

Length. 0.76 mm. 

Head. Tergal setae short or of medium length. Relative lengths of setae, 1 st row: 
a x = 10, a 2 = 11; 2 nd row: a x = 9,a 2 = 23, a 3 = 14; 3 rd row: a x = \\, a 2 = 16; 4 th row: a x 
= 11, a 2 = 22, fl 3 = 26, fl 4 = 20. Lateral group of setae: l x = 20, l 2 and / 3 not studied. 
Temporal organs 1.2 times as long as their shortest interdistance; small aperture close 
to posterior margin. Head cuticle glabrous. 



PAUROPODA FROM VIETNAM 



85 




Figs 22-31 
Allopauropus (D.) barbatulus sp. n., holotype, ad. 9( 9 ). 22, head, median and right part, tergal 
view; 23, left antenna, sternal view; 24, collum segment, median and left part, sternal view; 25, 
tergite VI, right posterior part; 26, T 4 ; 27, seta on coxa of leg 9; 28, seta on trochanter of leg 9; 
29, tarsus of leg 9; 30, pygidium, posterior part, sternal view; 31, anal plate, tergal view. Scale 
a: Figs 22, 24-26; b: Figs 27-31; c: Fig. 23. 



Antennae. Segment 4 with 4 setae, /?' not studied, p subcylindrical, tapering 
distally, annulate, /?" cylindrical, striate, /?'" a rudimentary knob, relative lengths of 
setae: p = 100, /?' = ?, /?" = 33. Tergal branch t somewhat fusiform, 2.2 times as long 
as greatest diameter and 1 .2 times as long as sternal branch s. The latter 1 .4 times as 



86 U. SCHELLER 

long as greatest diameter, anterodistal corner truncated. Seta q as seta p of 4 th segment 
but much thinner, 1.6 times as long as s. Relative lengths of flagella (base segments 
included) and base segments alone: F 1 = 100, bs x = 8; F 2 = 27, bs 2 = 4; F 3 = 77, bs 3 = 
6. F l 5.1 times as long as t, F 2 and F 3 1.7 and 4.8 times as long as s respectively. Distal 
calyces low, distal part of flagella axes widened, fusiform. Globulus g 1 .2 times as long 
as greatest diameter, ~ 9 bracts, capsule somewhat longer than wide; width of g 0.9 of 
greatest diameter of t. Antennae glabrous. 

Trunk. Setae of collum segment simple, cylindrical, blunt, densely annulate. 
Sublateral seta twice longer than submedian seta; sternite process narrow anteriorly 
and with small V-shaped anterior incision. Appendages wide, obliquely subcylindrical, 
caps flat. Process and appendages glabrous. 

Setae on tergites thin, short, similar to those on tergal side of head, 4+4 setae on 
tergite I, probably 6+6 on II-IV, 4+2 on VI. Posteromedian setae on tergite VI pointed, 
annulate, 0.2 of their interdistance. 

Bothriotricha. All bothriotricha except T 4 lost or broken. T 4 = 55 urn, with thin, 
simple, straight axes, pubescence of straight, simple, almost erect hairs, which are 
longest just outside the middle. 

Legs. Seta on coxa of leg 9 very thin, simple, cylindrical, striate; seta on 
trochanter furcate, main branch as on coxal seta, secondary branch very short and thin, 
striate. These setae simple on legs 1-8. Tarsus of leg 9 tapering, 2.5 times as long as 
greatest diameter. Proximal seta very thin, cylindrical, blunt, striate, curved inwards. 
Distal seta somewhat davate, striate, its length 0.3 of length of tarsus and 1.9 times as 
long as proximal seta. Cuticle of tarsus glabrous. 

Pygidium. Tergum. Posterior margin rounded, but with slight indentation below 
setae a x . Relative lengths of setae: a x = 10, a 2 = 7, a 3 = 15, st = 2. a x , a 2 and a 3 very 
thin, tapering, striate distally, diverging; st short, somewhat davate, curved inwards 
and converging. Distance a x - a x almost as long as a x and somewhat longer than dis- 
tance a x - a 2 , which is twice longer than distance a 2 - a 3 , distance st - st 3 times longer 
than st and 0.6 of distance a x - a x . Cuticle glabrous. 

Sternum. Posterior margin between b x almost straight. Relative lengths of setae 
(a x = 10): b x = 23, b 2 = 13. These setae thin, tapering, striate distally. b x 0.9 of their 
interdistance, b 2 as long as distance b x -b 2 . 

Anal plate narrow at base, broadly spatulate with rounded posterolateral 
corners. Posteromedian margin almost straight but with small indentation on tergal 
side, somewhat cordate in dorsal view. Plate 1 . 1 times as long as wide, posterolateral 
corners each with a short-stalked subspherical appendage. Appendages diverging, 1/3 
of length of plate. Plate with short pubescence, indistinct and irregular on sternal side, 
arranged in 6 distinct longitudinal rows on tergal side; pubescence longest and erect on 
appendages. 

4. Allopauropus (D.) absimilis sp. n. Figs 32-37 

Type material: Holotype: ad. 9( $ ), Vietnam, Kien Giang Province, Kien Luong, Hon 
Chong, Nui Hang Tien, under bush on limestone, soil sample, Berlese extraction, 19.1.2000 (Loc. 
VŒT-868). 

Other material. Same data as holotype, 1 juv. 3. 

Total number. 2 specimens. 



PAUROPODA FROM VIETNAM 



87 




Figs 32-37 
Allopauropus (D.) absimilis sp. n., holotype, ad. 9(5). 32, head, median and right part, tergal 
view; 33, left antenna, tergal view; 34, seta on coxa of leg 9; 35, seta on trochanter of leg 9; 36, 
tarsus of leg 9; 37, pygidium, posteromedian and left part, sternal view. Scale a: Figs 34-36; b: 
Figs 32, 33, 37. 

Diagnosis. A. absimilis sp. n. may be a close relative to Remy's A. consociatus 
from Madagascar (Remy, 1956a). The anal plates have the same groundplan and the 
antennal globuli are similar in both species. They can be distinguished by the shape of 
the posterior processes of the anal plate (0.5 of the length of plate in A. absimilis sp. 
n., as long as plate in A. consociatus Remy), by the shape of the seta on trochanter of 
leg 9 (simple, not furcate) and by some setae on the pygidial tergum (a{>a x - a x , not 
a l <a l - a x ; st pointed, not blunt). 

Etymology. From Latin absimilis = unlike (referring to the processes and ap- 
pendages of the anal plate). 



88 u. scheller 

Description 

Length. 0.81 mm. 

Head. Most setae cylindrical, striate; all of those studied blunt, except a 3 in 2 rd 
tergal row and l x which are tapering. Relative lengths of setae (a 2 in 1 st row = 10, a x 
lacking,): 2 nd row: a x = 9, a 3 = 10; 3 rd row: a x = a 2 = 9; 4 th row: a x = a 4 = 11, a 2 = a 3 = 
14; lateral group of setae: l x = 2l. Temporal organs 1.4 times as long as their shortest 
interdistance. Head cuticle glabrous. 

Antennae. Antennae in bad condition. Seta p cylindrical, tapering distally, 
striate, 1 = 40, p" = 10 p.m. Tergal branch t thin, slender, 5.3 times as long as greatest 
diameter and 1.8 times as long as sternal branch s which is about 2.4 times as long as 
its greatest diameter; anterior corner distinctly truncate. Flagellum F 3 , 1 = 83 (base 
segment included), base segment alone 1 = 4 urn. F 3 4.6 times as long as s. Globulus g 
1.5 times as long as greatest diameter; width of g 1.6 times as long as greatest diameter 
of t. Antennae glabrous. 

Trunk and bothriotricha not available for study. 

Legs. Seta on coxa of leg 9 furcate, branches similar to each other, cylindrical, 
blunt, striate; seta on trochanter simple, similar to main branch of coxal seta. These 
setae on legs 1-8 simple, seta on coxa thickest, seta on trochanter longest. Tarsus of leg 
9 4.5 times as long as greatest diameter, very slender in distal 2/3. Proximal seta thin, 
tapering, pointed, with minute pubescence in distal half; proximal seta 0.3 of length of 
tarsus and twice longer than distal seta; the latter subcylindrical, tapering, pointed, 
striate. Cuticle of tarsus faintly granular. 

Pygidium. Tergum. Posterior margin evenly rounded, but with distinct semicir- 
cular lobe between st above anal plate. Relative lengths of setae: a x = 10, a 2 = 11, a 3 = 
17, st = 9. These setae tapering, pointed, almost glabrous; a x , a 2 and st converging, a 3 
diverging. Distance a x - a x 1.4 times as long as a x ; distance a x - a 2 3.8 times as long as 
distance a 2 - a 3 ; distance st - st 1.8 times as long as st and 1.1 times as long as distance 
a x - a x . Cuticle glabrous. 

Sternum. Posterior margin between b x with 3 inner lobes, median one lowest 
and situated below anal plate. Relative lengths of setae (a x = 10): b x = 30, b 2 = 8 and 
9. These setae thin, tapering, striate distally. b x 1.2 times as long as their interdistance, 
b 2 0.7 of distance b x - b 2 . Cuticle glabrous. 

Anal plate 1 . 1 times as wide as long, narrowest anteriorly, with convex lateral 
margins and two posterior triangular lobes separated by a broadly V-shaped incision. 
Each lobe with a subcylindrical and tapering process protruding backwards, processes 
almost 0.5 of length of plate. Besides, two short subparallel appendages protruding 
backwards-downwards from inner sternal side of posterior lobes; length of appendage 
0.5 of length of posterior processes. Plate and appendages glabrous. 

Genus Pauropus Lubbock, 1 867 

5. Pauropus asiaticus sp. n. Figs 38-46 

Type material. Holotype: ad. 9( 9 ), Vietnam, Kien Giang Province, Kien Luong, Hon 
Chong, Nui Hang Tien, under bush on limestone, litter, Berlese extraction, 19.1.2000 (Loc. VI- 
ET-865). Paratypes: Same data as holotype, 1 ad. 9(9) (Loc. VIET-863). Ibidem, soil sample, 
Berlese extraction, 3 ad. 9(c?), 1 subad. 8(9), 19.1.2000 (Loc. VIET-864). 

Total number. 6 specimens. 



PAUROPODA FROM VIETNAM 



89 




Figs 38-46 
Pauropus asiaticus sp. n., holotype, ad. 9(9). 38, head, median and right part, tergal view; 39, 
right antenna, tergal view; 40, collum segment, median and left part, sternal view; 41, seta on 
coxa of leg 9; 42, seta on trochanter of leg 9; 43, tarsus of leg 9; 44, right genital papilla, outer 
lateral view; 45, pygidium, median and left side, sternal view; 46, anal plate, lateral view. 
Pubescence only partly drawn in Fig. 40. Scale a: Figs 41-43; b: Figs 39, 40, 44; c: Figs 38, 
45,46. 



90 U. SCHELLER 

Diagnosis. P. asiaticus sp. n. closely resembles P . forficularis Serieller from Sri 
Lanka (Serieller, 1970). Distinctive characters are e.g. the shape of both the lateral 
branches and the posteromedian appendages of the anal plate (the former subcylindri- 
cal and long in P. asiaticus sp. n., tapering and short in P . forficularis Serieller; the lat- 
ter lanceolate in P. asiaticus sp. n., claw-like in P. forficularis). The setae of the py- 
gidial tergum are glabrous in P. asiaticus sp. n., but with a distinct pubescence in P. 
forficularis, and the pygidial setae 03 are 1.5 times as long as the a 2 in P. asiaticus sp. 
n., almost of the same length in P. forficularis. The new species may also be close (but 
to a less degree) to P. wieheorum Remy from Mauritius (Remy, 1959) and to P. diffi- 
cilis Remy from Pondichéry (Remy, 1961). 

Etymology. Latinized adjective of Asia. 

Description 

Length. (0.98-)1.02(-1.42) mm. 

Head. Most tergal setae lacking, those studied indicating long blunt glabrous 
setae. a 2 in 1 st and 2 nd rows and a x in 3 rd row somewhat davate; a 3 in 2 nd row and a 4 in 
4 th row cylindrical. Temporal organs covering almost whole the posterior half of lateral 
side of head, their length 0.8 of their shortest interdistance. A very small pore in 
posterior part of temporal organ close to insertion point of l v Head cuticle and temporal 
organs glabrous. 

Antennae. Segment 4 with 6 cylindrical setae, most of them lacking. In a 
paratype specimen p" = 13 urn, /?'" = 2 and u = 1 urn. Tergal branch t very slender, 
subcylindrical, 7.0(-7.1) times as long as its greatest diameter and (1.2-)1.3 times as 
long as sternal branch s which is very little davate, (3.9-)4.3 times as long as its 
greatest diameter. Seta q cylindrical, tapering distally, indistinctly striate, 0.9 of length 
of s. Relative lengths of flagella (base segments included) and base segments alone: F 1 
= 100, bs y = 6(-7); F 2 = (63-)79(-82), bs 2 = 6; F 3 = (78-)86, bs 3 = 6(-7); F x 2.3(-2.5) 
times as long as t, F 2 and F 3 (2.2-)2.4 and (2.6-)2.7 times as long as s respectively. 
Globulus g 1.5 times as long as wide, with 9(-ll) bracts, capsule with flattened bottom, 
width of g 0.9 of greatest diameter of t. Antennae glabrous. 

Trunk. Setae of collum segment furcate, both branches with minute pubescence; 
primary branch foliform, (6.5-)6.6 times as long as broad, secondary branch rudimen- 
tary, subcylindrical. Sublateral setae 1.4(-1.5) times as long as submedian ones. 
Sternite process with narrow anterior part with distal incision; appendages low with flat 
caps; process and appendages with distinct pubescence. 

Setae on tergite I lanceolate, glabrous; 4+4 setae on I, 6+6 on II-IV, ? on V and 
VI. 

Bothriotricha. Most bothriotricha lost or broken. Relative lengths of them: T x - 
100, T 2 = 101, T 3 = (107-108), T A = (156), T 5 = ?. Bothriotricha with thin straight axes 
with short pubescence, for the greatest part oblique but distally erect. 

Genital papillae (paratypes). Genital papillae with almost straight sides, nar- 
rowly conical, 2.1-2.2 times as long as greatest diameter, glabrous. Apical seta short, 
0.1-0.2 of length of papilla. Seta on coxa of leg 2 as coxal setae on other anterior legs. 

Legs. Posterior legs long. Setae on coxa and trochanter of leg 9 furcate, with 
minute pubescence, main branch thickest, fusiform; secondary branch somewhat 



PAUROPODA FROM VIETNAM 91 

davate (or cylindrical), blunt, protruding from a point 1/4 above the base of seta; 
secondary branch 0.8 of length of primary branch. More anteriorly these setae with a 
rudimentary cylindrical glabrous secondary branch. Tarsus of leg 9 strongly tapering, 
slender, 5.5(-6.0) times as long as greatest diameter. Setae with short oblique pubes- 
cence, proximal seta tapering, pointed; distal seta somewhat davate, blunt. Proximal 
seta 0.4 of length of tarsus and 4.4 times as long as distal seta. Cuticle of tarsus with 
minute pubescence. 

Pygidium. Tergum. Posterior part broadly triangular, between a x si small lobe 
with median incision. Relative lengths of setae: a x = 10, a 2 = 12(-13), a 3 = 17(-18), st 
= 6. All setae glabrous, all but st lanceolate, somewhat curved inwards, st broadly 
lanceolate, bent outwards and converging; a x and a 2 directed posteriorly, a 3 diverging. 
Distance a x - a x 1.2 times as long as a x ; distance a x - a 2 1.3 times as long as distance 
a 2 - a 3 \ distance st - st (1.4-)1.5 times as long as st and 0.8 of distance a x - a x . 

Sternum. Posterior margin between b x broadly indented and with low rounded 
median bulge below anal plate. Relative lengths of setae (a x = 10): b x = 47, b 2 = 15, b 3 
= 24. b x tapering, pointed, glabrous; b 2 lanceolate, somewhat curved inwards, glabrous; 
b 3 subcylindrical and with minute pubescence most distally. b x 1.7 times as long as 
their interdistance, b 2 2.0(-2.1) times as long as distance b x - b 2 ; b 3 0.9 of distance 
b 3 - b 3 . 

Anal plate broadest anteriorly, this part with two posterolateral branches 
directed backwards-outwards and somewhat curved inwards, their length 0.7 of total 
length of plate with appendages. Posteromedian part of plate subrectangular, with 
posterior V-shaped cleft separating two branches, each with a distal, posteriorly di- 
rected, somewhat lanceolate appendage with short erect pubescence. Length of 
appendages almost 0.5 of length of lateral branches. 

EURYPAUROPODIDAE 

In most collections of Pauropoda Eurypauropodidae are poorly represented. The 
occurrence of 3 species, 2 of them new to science, in this material from Vietnam, indi- 
cates a proportionally high species diversity. This is in accordance with previous 
reports from south Asia, from Thailand (Serieller, 1995) and from Nepal (Serieller, 
2000), which show that Eurypauropodidae, at least locally, may be almost as diverse 
as Pauropodidae. A further indication in this direction are two other Samarangopus 
species occurring in this collection which are unfortunately not in the best condition 
and therefore not described. One, 1 ad. 9(6) and 1 juv. 5, from a limestone hill at Hon 
Chong, Nui Hang Tien, (Loc.VIET-866), is close to S. longipenes Serieller from 
Borneo (Serieller, 2001). The other, a juv. 6 specimen, from Hon Chong, Mo So, (Loc. 
VIET-014), is close to S. segniter Scheller (Scheller et al., 1994) from Borneo. 

Eurypauropodinae 

Genus Samarangopus Verhoeff, 1934 

6. Samarangopus campanulatus sp. n. Figs 47-64 

Type material. Holotype: ad. 9(9) Vietnam, Kien Giang Province, Kien Luong, Hon 
Chong, Mo So, under bush on limestone, humus on rock, Berlese extraction, 18.XII.1994 (Loc. 
VIET-018, leg. Deharveng & Le Cong Kiet). Paratypes: Same data as holotype, 6 ad. 
9(2 c* ,4 9), 1 subad. 8(5). 



92 



U. SCHELLER 




ìxU-^y^Jj^- cujjuxìJ*2£i>- 



Figs 47-55 
Samarangopus campanulatus sp. n., holotype, ad. 9(2). 47, body, tergal view, tergites I-VI 
showing the symmetric pattern of depressions surrounded by raised cuticle; 48, body, lateral 
view; 49, right antenna, sternal view; 50, collum segment, median and left part, sternal view; 51, 
tergite I, part of anterior margin, tergal view; 52, tergite I, central part; 53, tergite I, right 
posterolateral corner; 54, tergite I, median part of posterior margin; 55, tergite II, right antero- 
lateral corner, tergal view. Scale a: Figs 51-55; b: Figs 49-50. 



PAUROPODA FROM VIETNAM 93 

Other material. Vietnam, Kien Giang Province, Kien Luong, Hon Chong, Mo So, un- 
der bush on limestone, soil sample, Berlese extraction, 1 ad. 9(6), 18.XII.1994 (Loc. VIET-013, 
leg. Deharveng & Le Cong Kiet). Ibidem, Hon Chong, Batai hill, under bushes, soil sample, 
Berlese extraction, 1 ad. 9(9), 1 juv. 3, 21. XII. 1994 (Loc. VIET-028, leg. Deharveng & Le Cong 
Kiet). Ibidem, Hon Chong, Nui Son Cha, under bush on limestone, litter, Berlese extraction, 1 
ad. 9(6), 1 juv. 5, 18.1.2000 (Loc. VIET-836). Ibidem, Hon Chong, Nui Hang Tien, under bush 
on limestone, soil sample, Berlese extraction, 1 ad. 9(a), 1 juv. 5, 19.1.2000 (Loc. VIET-866). 

Total number. 15 specimens. 

Diagnosis. S. campanulatus sp. n. is most similar to the Bornean S. segniter 
Scheller (Scheller et al., 1994) and to S. doiinthanonaeus Scheller from north-western 
Thailand (Serieller, 1995). Distinctive characters in relation to S. segniter Serieller are: 
the shape of the pygidial setae st (straight and longer than a x in S. campanulatus sp. n., 
curved outwards and very short in S. segniter Serieller), the shape of the posterior 
margin of the pygidial sternum (straight in S. campanulatus sp. n., with posteromedian 
lobe with median incision in S. segniter) and the shape of the pygidial setae b 2 (thin 
tapering in S. campanulatus sp. n., broad lanceolate in S. segniter). Distinctive 
characters in relation to S. doiinthanonaeus are: the number of large protuberances on 
the posterior margin of tergite VI (one protuberance behind the pit of bothriotrix T 5 in 
S. campanulatus sp. n., two in S. doiinthanonaeus Serieller), the shape and pubescence 
of the bothriotrix T 3 (proportionally large distal swelling with minute pubescence in S. 
campanulatus sp. n., indistinct swelling and distinct pubescence in S. doiinthanonaeus) 
and the length of the pygidial setae st (about half of the length of the anal plate in S. 
campanulatus sp. n., as long as the anal plate in S. doiinthanonaeus). 

Etymology. From Latin campana = bell (referring to the shape of the large 
marginal setae on the anterior tergites). 

Description 

Length. (0.74-)0.84(-0.90) mm. 

Head (holotype only). Vertex: no setae. Tempus: not studied. Index of setae, lat- 
eral row, 3 setae, lp x = ?, lp 2 = 20, lp 3 = 14, lp x - lp 2 =?, lp 2 - lp 3 = ?, lp x - lp x = 16; an- 
terior row, 3 setae, la x = 8, la 2 = 17, la 3 = 21, la x - la 2 = 7, la 2 - /a 3 = la x - la x = 13 urn. 
Distance la x - la x /la x - la 2 = 1.9. Peristomal setae: ipe not ascertained, pe x = 9,pe 2 = 14, 
pe 3 =13, pe 4 = 14, pe 5 = 20, pe x - pe 2 = 3, pe 2 - pe 3 = 2, pe 3 - pe 4 = 5, pe 4 - pe 5 =11, 
pe x - pe x = 16 |im. 

Antennae. Antennae almost glabrous; chaetotaxy of segments 1-4: 2/2/2+g73; 
setae thin, cylindrical, striate, their lengths on segment 4: p = 18(-20), /?' = (15-) 16, p" 
= 13(-16) urn; no /?'", u and r. Sternal branch s with distinct anterior indentation at 
level of F 2 , anterior margin = (11-)13, posterior margin = (19-)20(-21), of base = 
6(-7), maximum = 11(-12), q thin, cylindrical, annulate-striate, 1 = 10(-16) u,m. 
Posterior margin/length of g 1.3(-1.7), posterior margin/maximum = 1.8(-1.9), 
maximum 0/0 of base = (1.7-)1.8. Tergal branch fusiform, widest in distal half, 1 = 20 
(-24), of base = 5, maximum = 7(-8) urn; pore not ascertained; length of 
^/maximum = (2.5-)2.7. Globulus g, 1 = (12-)14(-15), maximum = 7(-8) urn; length 
/maximum = (1.8-)2.0; number of bracts (10-)13, their length = 7, capsule spherical, 
= 3 (im. Relative lengths of flagella (base segments included): F x = 100, F 2 = 
42(-49), F 3 = (79)82(-85). Lengths of base segments: bs x = 10(-13), bs 2 = 7(-8), bs 3 = 



94 



U. SCHELLER 



,&■ <3 ® ® il 

^ 6 ° 




Figs 56-64 
Samarangopus campanulatus sp. n.: 56-57, 59-64, holotype, ad. 9(9), 58, paratype ad. 9(3). 
56, tergite VI, median and right posterolateral part; 57, T 3 ; 58, genital papillae, anterior view; 59, 
seta on coxa of leg 9; 60, seta on trochanter of leg 9; 61, tarsus of leg 1; 62, tarsus of leg 9; 63, 
femur of leg 1 with appendage; 64, pygidium, sternal view. Pubescence only partly drawn in Figs 
61-63. Scale a: Figs 56-58; b: Figs 59-64. 



PAUROPODA FROM VIETNAM 95 

11(-12) um. F x (3.6-)4.1 times as long as t, F 2 and F 3 (1.8-)2.1 and (3.3-)3.4 times as 
long as s, respectively. F 2 thinner than F x and F 3 . Calyces of F x largest, conical, those 
of F 2 and F 3 smaller, subhemispherical. 

Trunk. Setae of collum segment similar to each other, furcate. Branches 
tapering, pointed; main branch, striate; secondary branch rudimentary, glabrous; both 
setae 1 = 13(-14) urn. Sternite process broad and low, with anterior V-shaped incision. 
Appendages directed posteriorly, barrel-shaped, caps flat and with collar. Process and 
appendages with minute pubescence, caps glabrous. 

Tergites. A single row of campanulate protuberances on anterior and lateral mar- 
gins of tergite I, on lateral margins of II- V and on posterior margin of VI. Central part 
of all tergites with small cylindrical cuticular structures with distal candleflame-like 
vesicle surrounded by circular, funnel-shaped, transparent collar. Surface between 
these organs coarse. 

Tergites I- VI with a symmetric pattern of depressions surrounded by raised 
cuticle with the above mentioned subcylindrical structures (Figs 47, 52, 54, 56). 
Number of campanulate marginal protuberances: I (27-)31; II, 1 small - Z\ - 1 small + 
(8-) 9; m, 5(6) - T 2 - 1 small + (6-)7; IV, 5(-6) - T 3 - (4-)5; V, 7 - 7/ 4 - 4; VI, (5-)6 - T 5 
- 1. Length/width ratio of tergites: I = (0.5-)0.6, II and V = 0.4(-0.5), III and VI = 0.5, 
IV = 0.5(-0.6). 

Bothriotricha. All bothriotricha but 7/ 3 curled distally and with very thin axes; 
these glabrous except for a minute pubescence on their distal third. T 3 with thicker axes 
and distal 2/5 increasing in width, forming a davate distal end-swelling with minute 
pubescence. Relative lengths of bothriotricha: T x = 100, T 2 = 62(-105), T 2 = 28(-53), T 4 
= ?(72-82), T 5 = (72-)81(-100). 

Genital papillae. Base segments cylindrical. Length of papillae = 39(-48) um, 
greatest = 16(-20) urn, length of seta = 20 um. Proximal part of genital papillae sub- 
cylindrical, distal part conical, seta (0.4-)0.5 of length of papilla, which is 2.4 times as 
long as greatest diameter. Cuticle glabrous. Coxal seta of leg 2 as on leg 1, length = 
17(-20) urn. 

Legs. All legs 5-segmented. Seta on coxa and trochanter of leg 9 subsimilar to 
each other, very thin, furcate, striate, with glabrous base; length of secondary branch 
0.7-0.8 of primary one. More anteriorly these setae with rudimentary pointed glabrous 
secondary branches. Tarsi short, tapering, those of leg 9 1.8(-1.9) times as long as 
greatest diameter, with two tergal setae, both pointed and glabrous. Proximal seta (13-) 15, 
distal one 9(-10) urn. Proximal setae 0.4(-0.5) of length of tarsus and (1.3-)1.7 times as 
long as distal seta. Cuticle of tarsus with minute pubescence. No proximal seta on 
tarsus of leg 1. All legs with large main claw and small setose anterior secondary claw; 
on leg 9 the former reaching 0.5 of the length of tarsus. On anterior side of femur of 
leg 1 a blunt appendage with short pubescence, length = 4(-5) urn. 

Pygidium. Tergum. Posterior margin between the lateral digitiform appendages 
with 5 lobes, a triangular median one and two rounded ones on each side of it. Setae 
glabrous, a x and a 2 somewhat davate, the former curved inwards and the latter almost 
straight; a 3 straight, cylindrical, somewhat tapering, diverging; st straight, lanceolate. 
Lengths of setae: a x = a 2 = (5-)6, a 3 = (10-)12(-14), st = 9(-12) urn. Distance a x - a x = 
9(-ll), a 2 -a 2 = (28-)29(-30), fl 3 - a 3 = (34-)37(-39), a x -a 2 = 9(-10), a 2 - a 3 = 5, st - st 



96 U. SCHELLER 

= 9(-12) firn. Distance a x - a x 1.5(-2.0) times as long as a lt distance a x - a 2 (1.8-)2.0 
times as long as distance a 2 - a 3 ; distance st - st (1.1-)1.3 times as long as st and (1.1-)1.3 
times as long as distance a x - a x . Cuticle glabrous. 

Sternum. Posterior margin with shallow indentations just inside b x and in 
between a broad low lobe with straight posterior margin. Setae thin, tapering, distally 
striate, b 2 and b 3 pointed. Lengths of setae: b x = (32-)37, b 2 = 15 and 17(-19), b 3 = (11- 
)12 urn . Distance b x - b x = (34-)36, b 2 -b 2 = (47-)50, b 3 - b 3 = (21-)22(-23), b x - b 2 = 
(18-)20, b 2 -b 3 = (13-)14 urn. Distance b x - b x (0.9-)1.0 of the length of b x , b 2 0.7-0.8 
of distance b x - b 2 , b 3 0.5 of distance b 3 - b 3 . 

Anal plate twice as wider than long, widest in the middle, widest part about 1.5 
times as wide as distal part; broadest part forming indistinct posterolateral corners, 
from there two short, thin, cylindrical, blunt, pubescent appendages protruding 
obliquely backwards; appendages 0.3 of length of plate; posterior 3/5 of plate divided 
into two tapering branches by a narrow V-shaped incision, each branch provided with 
two appendages: a submedian short straight tapering glabrous one and a stalked 
bladder of triangular shape in sternal view. Bladder- shaped appendages 0.6 of length 
of plate. Plate glabrous, bladder- shaped appendages with short erect pubescence. 

Sphaeropauropodinae 

Genus Sphaeropauropus Silvestri, 1930 

7. Sphaeropauropus malayus Silvestri, 1930 

Sphaeropauropus malayus Silvestri, 1930: 229-231, figs 3-4. 

Material. Vietnam, Kien Giang Province, Kien Luong, Hon Chong, Mo So, under bush 
on limestone, soil sample, Berlese extraction, 1 ad. 9(9), 18.XII.1994 (Loc. VIET-014, leg. 
Deharveng & Le Cong Kiet). 

Total number. 1 specimen. 

General distribution. S. malayus has been reported from Java (Silvestri, 1930; 
Scheller, 1998), Réunion (Remy, 1956b) and the Philippines (Remy, 1957b). However, 
only on the base of the descriptions, none of the specimens reported by Remy seems 
to be completely identical with the type specimens from Java (Serieller, 1998) and only 
an examination of Remy' s specimens can decide about their conspecificity with S. 
malayus. 

8. Sphaeropauropus lecongkieti sp. n. Figs 65-84 

Type material. Holotype: ad. 9{S), Vietnam, Kien Giang Province, Kien Luong, Hon 
Chong, Mo So, under bush on limestone, soil sample, Berlese extraction, 18.XII.1994 (Loc. VI- 
ET-020, leg. Deharveng & Le Cong Kiet). Paratypes: Vietnam, Kien Giang Province, Kien 
Luong, Hon Chong, Batai, under bush on limestone, soil sample, Berlese extraction, 1 ad. 9( 9 ), 
1 subad. 8(9), 18.1.2000 (Loc. VIET-844). 

Other material. Vietnam, Kien Giang Province, Kien Luong, Hon Chong, Batai, under 
bushes,l subad. 8(9), 1 juv. 5, 21.XJI.1994 (Loc. VIET-028, leg. Deharveng & Le Cong Kiet). 
Ibidem, same date, 1 subad. 8(9) (Loc. VIET-029. leg. Deharveng & Le Cong Kiet). Ibidem, 
Hon Chong, near pagoda, under bushes, Utter, Berlese extraction, 1 ad. 9(9), 1 subad. 8(c?), 
19.1.2000 (Loc. VIET-846). 

Total number. 8 specimens. 

Diagnosis. S. lecongkieti sp. n. is close to S. brevi globulatus Serieller from 
Nepal (Scheller, 2000) by similarities in the shape of the tuft-like setae and the 



PAUROPODA FROM VIETNAM 97 

pubescence on the tergites and in the general shape of the anal plate. Good distinctive 
characters are: the shape of the setae ipe 2 (simple and davate in S. lecongkieti sp. n., 
thin and furcate distally in S. brevi globulatus), the proportion between the length of the 
base segment of the F 3 and the length of the sternal branch s (0.5, not 0.8), and the 
shape of the antennal globulus g (2.3 times as long as greatest width, not 1.2; 8-9 
bracts, not 10-13; capsule spherical, not flattened). The two species may also be 
distinguished by the peculiar structure in the anterior part of the tergites V and VI (Figs 
73-75), occurring in S. lecongkieti sp. n., but not in S. breviglobulatus. 

Etymology. Dedicated to one of the collectors, Professor Le Cong Kiet (Ho Chi 
Minn City). 

Description (holotype only) 

Length. 0.85 mm. 

Head. All setae but ipe 2 cylindrical, thin, pointed, striate; ipe 2 davate, bent in- 
wards, with short pubescence. Vertex: no setae. Tempus: te not identified, no fungiform 
organ. Frons: frontal pore not identified. Index of frontal setae: median row, 3 setae, 
mp = 20, mm = 25, ma =10, mp - mm =10, mm - ma = 21 urn; lateral row, 3 setae, lp x 
= 22, lp 2 = 20, lp 3 = 24, lp x - lp 2 = 12, lp 2 - lp 3 = 13, lp i - lp x = 19 urn; anterior row, 3 
setae, la x = 16, la 2 = 19, la 3 = 17, la x - la 2 = 10, la 2 - la 3 = 22, la x - la x = 15 um. Distance 
la x - la x lla x - la 2 = 1.4. Peristomal setae, ? in number, ipe x and lb not ascertained, ipe 2 
= 9, pe x = 13, pe 2 = 17, pe 3 = 20, pe 4 = 16, pe 5 = 25, pe x - pe 2 = 4, pe 2 - pe 3 = 2, pe 3 - 
pe 4 = ?, pe 4 - pe 5 =18, pe x - pe x = 20, pe 5 - pe 5 = 68 urn. 

Antennae. Cuticle of segments 1-4 and branches t and s sparsely granular, g and 
flagella glabrous. Chaetotaxy of segments 1-4: 2/2/2/3. Setae subcylindrical, tapering, 
annulate; g' not ascertained, setae on segment 4, p = 24, p' = 22, p" = 16 urn. Sternal 
branch with distinct anterodistal truncation at the level of F 2 , anterior margin =17, pos- 
terior margin = 26, of base = 8, maximum = 14, q = 16 urn. Anterior margin/length 
of g = 0.9; anterior margin/length of q = 1.1; anterior margin/posterior margin = 0.7; 
anterior margin/maximum = 1.2; maximum 0/0 of base = 1.7. Tergal branch t 
fusiform, length = 33, of base = 5, maximum 0=11 urn, length oft/maximum = 
3.0. Globulus g straight, 1 = 20, maximum = 9 urn; with 8 or 9 bracts, their length = 
9 urn; capsule subspherical, = 5 urn. Relative lengths of flagella (base segments in- 
cluded): F x = 100, F 2 = 49, F 3 = 83. Length of base segments: bs x = 22, bs 2 = 9, bs 3 = 
16 urn. F x 3.2 times as long as t, F 2 and F 3 1.7 and 2.9 times as long as s respectively. 
Calyces helmet-shaped, those of F x largest. 

Trunk. Setae of collum segment furcate; main branch cylindrical, striate-annu- 
late, secondary branch rudimentary, conical, pointed, glabrous; length of submedian 
ones = 25, lateral ones = 20 urn; sterni te process broad, anterior part divided into two 
rounded lobes separated by a shallow median incision; appendages short with thick 
round caps, = 20 urn. 

Tergites with tuft-like setae with davate stalk. Cuticle between setae with very 
small sessile organs in the centre of round glabrous spots surrounded by sparse but dis- 
tinct pubescence. Pubescence longest on lateral parts of tergites. 

Setae on outer margin of latero sternal furrows resembling tuft-like setae but 
with thinner axes, some anterior ones even pointed. All setae in inner row pointed and 
with shorter oblique pubescence. 



98 



U. SCHELLER 




Figs 65-72 
Sphaeropauropus lecongkieti sp. n., holotype, ad. 9(6). 65, head, median and left part, tergal 
view; 66, left antenna, sternal view; 67, collum segment, median and left part, sternal view; 68, 
tergite I, central part; 69, tergite I, near posterolateral corner; 70, tergite I, posterior margin; 71, 
tergite II, right laterosternal furrow, sternal view; 72, tergite VI, posteromedian margin, tergal 
view. Scale a: Figs 65, 67, 71; b: Figs 66, 68-70, 72. 



Anteromedian part of tergite V with wedge-shaped area, provided with densely 
packed transverse bands of small circular spots. A similar but smaller area on anterior 
part of tergite VI. 

Bothriotricha. All but T 3 with very thin axes and distinct pubescence, the latter 
erect on distal halves, hairs branched at least on T 5 . T 3 with thicker axes and distal 
ovoid swelling, its length 1/4 of length of bothriotrix; only distal part of T 3 with minute 



PAUROPODA FROM VIETNAM 



99 




Figs 73-84 
Sphaeropauropus lecongkieti sp. n., holotype, ad. 9(a). 73, tergite V, wedge-shaped cuticular 
pattern on anteromedian part; 74, detail of the cuticular pattern in Fig. 73 in higher magni- 
fication; 75, tergite VI, wedge-shaped cuticular pattern on anteromedian part; 76, T 3 ; 77, T 5 ; 78, 
genital papillae and seta on coxa of leg 2, anterior view; 79, seta on coxa of leg 9; 80, seta on 
trochanter of leg 9; 81, tarsus of leg 1; 82, tarsus of leg 9; 83, femur of leg 1 with appendage; 
84, pygidium, sternal view. Pubescence only partly drawn in Figs 81 and 82. Scale a: Figs 73, 
75; b: Figs 76-83; c: Fig. 84; d: Fig. 74. 

but dense pubescence of oblique hairs. Relative lengths of bothriotricha: T x = 100, T 2 
= T 5 = 95, T 3 = 75, T 4 = 90. 

Genital papillae. Proximal half subcylindrical, distal half conical, length = 51, 
greatest = 22 jim; length of seta = 20 |im. Papillae 2.6 times as long as greatest 



100 U. SCHELLER 

diameter, seta 0.4 of length of papilla. Cuticle glabrous. Coxal seta of leg 2 as on 
leg 1 , length = 27 um. 

Legs. All legs 5-segmented. Setae on coxa and trochanter of leg 9 furcate. Main 
branch thin, striate-annulate, on coxa blunt and on trochanter pointed; secondary 
branch rudimentary, curved, pointed, glabrous. These setae on legs 1-8 of similar shape 
but coxal setae thinner. 

Tarsi of leg 9 slender, tapering, somewhat bow-shaped, 4.1 times as long as 
greatest diameter, minutely pubescent; two tergal setae, both tapering, pointed, 
glabrous. Proximal seta = 22, distal seta = 8 urn, proximal seta 0.3 of length of tarsus 
and 2.8 times as long as distal seta. No proximal seta on tarsus of leg 1. On anterior 
side of femur of leg 1 a triangular blunt appendage with short pubescence; length of 
appendage = 5 urn. 

All legs with almost straight main claw and small setose anterior secondary 
claw. Base of main claw with distinct pubescence, other parts glabrous. Main claw of 
leg 1 = 13, secondary claw = 8 urn; main claw of leg 9 = 14, secondary claw = 9 urn; 
main claw 0.2 of length of tarsus in all legs. 

Pygidium. Tergum. Posterior margin rounded and with protruding lobe between 
a x . Setae subsimilar to each other, subcylindrical, curved inwards, with very short pu- 
bescence. a x , a 2 and a 3 cylindrical, blunt, converging; st tapering, directed posteriorly. 
Index of tergal setae: a x = 9, a 2 = a 3 = 7; a x - a x = 16, a 2 - a 2 = 30; a 3 - a 3 = 50, st = 13, 
st - st = 14 urn; st - st/a x - a x = 0.9, a x la x - a x = 0.6, a x la x - a 2 = 1.1, a x - a x la 2 - a 3 = 2.0. 
Tergum glabrous. 

Sternum. Posterior margin between b x evenly rounded, setae tapering, with 
short oblique pubescence or striate distally. Index of sternal setae: b x = 43, b 2 = 15, b 3 
= 16, b x -b x = 38, b 2 - b 2 = 43, b 3 - b 3 = 28, b x - b 2 = 11 urn; b x /b x - b x = 1.1, b x lb 2 = 
2.9, b 2 lb x - b 2 = 1.4, \lb 3 - b 3 = 0.6. 

Anal plate pentagonal and with 2 cylindrical branches protruding backwards 
from posterior part of sternal side. Each branch with posterolateral-posterosternal trun- 
cation and two distal appendages: one of them tergal submedian and tapering, the other 
submedian and davate. The latter 2.3 and 3.1 times as long as its greatest width and 
1/3 of length of plate. Anal plate glabrous, davate, with distinct erect pubescence. 

ACKNOWLEDGEMENTS 

The author is indebted to the collectors, Drs Louis Deharveng and Anne Bedos 
(Paris), and Professor Le Cong Kiet (Ho Chi Minh City). The two former have initiated 
and kindly supported this study. 

Anne Bedos could participate in the field sampling of January 2000 thanks to 
the grant "Bourse Germaine Cousin" of the Société Entomologique de France. 

REFERENCES 

Bedos, A. & Deharveng, L. 2000. Un nouveau Collembole Neanurinae du Sud du Vietnam, 
Blasconura batai sp. n., avec une clé des espèces du genre (Collembola: Neanuridae). 
Revue suisse de Zoologie 107: 351-357. 

Deharveng, L. & Bedos, A. 1995. Lepidonella lecongkieti n. sp., premier Collembole caverni- 
cole du Vietnam (Collembola, Insecta). Bulletin de la Société Entomologique de France 
100: 21-24. 



PAUROPODA FROM VIETNAM 101 



Deharveng, L. & Bedos, A. 1996. Rambutsinella, a new genus of Entomobryidae (Insecta: 

Collembola) from Southeast Asia. The Raffles Bulletin of Zoology 44: 279-285. 
Deuve, T. 1996. Description d'un coléoptère troglobie du genre Eustra, découvert dans un karst 

du Vietnam méridional (Paussidae). Revue Française d'Entomologie (N. S.) 18: 23-6. 
Hagino, Y. 1991. New species of the family Pauropodidae (Pauropoda) from Central Japan. The 

Canadian Entomologist 123: 1009-1045. 
Lubbock, J. 1867. On Pauropus, a new type of Centipede. Transactions of the Linnean Society, 

London 26: 181-190, pi. 10. 
Remy, P. A. 1933. Un Pauropode nouveau d'Indochine: Pauropus Dawydofß n. sp. Faune des 

Colonies Françaises 5: 189-193. 
Remy, P. A. 1947. Liste de Pauropodes d'Algérie, avec description de formes nouvelles. Bulletin 

de la Société des Sciences de Nancy 6: 66-84. 
Remy, P. A. 1956a. Pauropodes de Madagascar. Mémoires de l'Institut Scientifique de Mada- 
gascar 10: 101-229. 
Remy, P. A. 1956b. Pauropodes de l'île de la Réunion. Mémoires de l'Institut Scientifique de 

Madagascar 11: 127-166. 
Remy, P. A. 1957a. Pauropodes des États-Unis (Tennessee, North Carolina). Revue Française 

d'Entomologie 24: 82-87. 
Remy, P. A. 1957b. Pauropoda. In: Bernice, P. Bishop Museum (ed.). Insects of Micronesia 4: 

1-12. 
Remy, P. A. 1959. Pauropodes de l'île Maurice. 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 9: 1-24. 
Remy, P. A. & Rollet, G. 1960. Pauropodes de la côte de Madagascar. Bulletin de la Société 

d'Histoire Naturelle de la Moselle 38: 201-236. 
Scheller, U. 1970. The Pauropoda of Ceylon. Entomologica scandinavica, Supplementum 1: 

5-97. 

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

Scheller, U. 1998. Redescriptions of two Javanese species in Eurypauropodidae (Myriapoda, 
Pauropoda). Bolettino del Laboratorio di Entomologia Agraria "Filippo Silvestri" 54: 
145-154. 

Scheller, U. 2000. Eurypauropodidae from the Nepal Himalaya (Myriapoda, Pauropoda). 

Senckenbergiana biologica 80: 101-126. 
Scheller, U. 2001. Pauropoda (Myriapoda) from Sabah (East Malaysia) (Pauropoda and 

Symphyla of the Geneva Museum XI). Revue suisse de Zoologie 108: 949-986. 
Scheller, U., Brtnck, P. & Enckell, P. H. 1994. First record of Pauropoda (Myriapoda) on 

Borneo. Stobaeana 1: 1-14. 

Silvestri, F. 1902. Ordo Pauropoda. In: Berlese, A. Acari, Myriapoda et Scorpiones hucusque 

in Italia reperta. X: fase. 93, 95, 96. 
Silvestri, F. 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. Pauropoda. Bronn's Klassen und Ordnungen des Tierreichs 5, II, 3: 

121-200. 



Revue suisse de Zoologie 111 (1): 103-110; mars 2004 



A re-evaluation of the generic assignment of Bradypodion spinosum 
(Matschie, 1892) and some considerations on the genus 
Rhampholeon Günther, 1874 

Colin R. TILBURY 1 & Jean MARIAUX 2 

1 112A 3 rd Avenue, Fairland, Johannesburg 2195, South Africa. 

E-mail: sacoltil@mweb.co.za 
2 Dept des Invertébrés, Muséum d'histoire naturelle, CP 6434, CH-1211 Geneva 6, 

Switzerland. E-mail: jean.mariaux@mhn.ville-ge.ch 



A re-evaluation of the generic assignment of Bradypodion spinosum 
(Matschie, 1892) and some considerations on the genus Rhampholeon 
Günther, 1874. - The generic assignment of the dwarfed African chameleon 
Bradypodion spinosum is re-evaluated with regard to phenotype, soft tissue 
morphology, cranial osteology and DNA analysis. All methods used indicate 
that the species is unequivocally a member of the genus Rhampholeon to 
which it is formally transferred here: Rhampholeon spinosus (Matschie) 
comb. nov. Comparison of DNA with other species of the genus indicates 
that the genus Rhampholeon, as currently composed, is not monophyletic 
and consists of at least two separate lineages. 

Key-words: Bradypodion spinosum - Rhampholeon - Chamaeleonidae - 
taxonomy - molecular analysis - Usambara Mountains - Tanzania. 

INTRODUCTION 

Prior to 1892, only two species of Rhampholeon had been described viz. 
kerstenii (Peters, 1868) and spectrum (Bucholz, 1874). Although both were initially 
placed within the genus Chamaeleo, Günther erected the genus Rhampholeon in 1 874 
with spectrum as the type species for the genus. Both of these species have bicuspid 
claws. During a stay in the Eastern Usambara station of Derema in 1891, L. Conradt 
assembled a small collection of mammals and reptiles from which five new species of 
chameleons were described (Matschie, 1892) including two further new species of 
Rhampholeon named as Chamaeleon (Brookesia) brevicaudatus and Chamaeleon 
(Brookesia) temporalis. The latter, although possessing simple claws, has a typical 
Brookesiinae body habitus and it is not difficult to understand why it was allocated to 
Brookesia. Lastly there was a single specimen of a rather strange small chameleon with 
simple claws, an unusual ovo-globular rostral projection and prominent spines 
arranged in irregular rows along the dorsal keel and on the limbs. This chameleon was 
aptly named Chamaeleon spinosus, and all subsequent authors and taxonomists have 
treated it either as a member of the genus Chamaeleo (Werner, 1902; Mertens, 1966), 



Manuscript accepted 23.12.2003 



104 C. R. TILBURY & J. MARIAUX 

or more recently as a member of the genus Bradypodion [as B. spinosum] (Broadley 
& Howell, 1991; Klaver & Böhme, 1986; Uetz, 2003). However, a number of 
observations made us question the generic allocation of this species. The available 
evidence, as well as new characters pertinent to this question, are reviewed below. 

MATERIAL AND METHODS 

Specimens of B. spinosum examined 

[BMNH: The Natural History Museum (London, UK), MHNG: Muséum d'Histoire 
Naturelle de Genève (Geneva, Switzerland), MNHN: Muséum National d'Histoire 
Naturelle (Paris, France), PEM-R: Port Elizabeth Natural History Museum, (South 
Africa)]. 

BMNH 95.4.8.3 (FF), Usambara, Tanzania 

BMNH 1974.522 (FF), Mazumbai Estate, Western Usambara, Tanzania 

BMNH 1900. 12. 12.2 (MM), Ambangulu, Eastern Usambara, Tanzania 

BMNH 1971.159 (FF), Amani, Eastern Usambara, Tanzania 

MHNG 2609.067 (FF), Amani, Eastern Usambara, Tanzania 

MHNG 2620.032 (FF), Eastern Usambara, Tanzania (skeleton prepared) 

MHNG 2620.033 (FF), Eastern Usambara, Tanzania 

MHNG 2620.034 (FF), Mazumbai Estate, Western Usambara, Tanzania 

MHNG 2620.035 (MM), Mazumbai Estate, Western Usambara, Tanzania 

MHNG 2620.036 (MM), Mazumbai Estate, Western Usambara, Tanzania 

PEM-R 5738 (FF), Herkulu estate, Western Usambara Mtns, Tanzania 

Material used for the molecular analysis and EMBL/Genbank accession numbers: 

Bradypodion tavetanum (Steindachner, 1891) 

MHNG 2612.095, Bundunki, Uluguru, Tanzania. (AJ609593) 

Bradypodion oxyrhinum Klaver & Böhme, 1988 

MHNG 2612.096, Bundunki, Uluguru, Tanzania. (AJ609594) 

Bradypodion fischeri (Reichenow, 1887) 

MHNG 2609.080, Amani, Eastern Usambaras, Tanzania. (AJ609592) 

Bradypodion spinosum (Matschie, 1 892) 

MHNG 2609.067, Amani, Eastern Usambaras, Tanzania. (AJ609595) 

Bradypodion spinosum 

MHNG 2620.032, Eastern Usambara, Tanzania. (AJ609600) 

Rhampholeon sp. 

MHNG 2624.074, Kindoroko FR, North Pares, Tanzania. (AJ609601) 

Rhampholeon spectrum (Buchholz, 1874) 

MNHN 3511, Cameroon. (AJ609599) 

Rhampholeon uluguruensis (Tilbury & Emmrich, 1996) 

MHNG 2617.099, Mkungwe, Uluguru, Tanzania. (AJ609602) 



GENERIC ASSIGNMENT OF BRADYPODION SPINOSUM 1 05 

Rhampholeon kerstenii (Peters, 1868) 

MHNG 2611.010, Captive breeding. (AJ609598) 

Rhampholeon brevicaudatus (Matschie, 1892) 

MHNG 2609.064, Amani, Eastern Usambaras, Tanzania. (AJ609597) 

Sceloporus undulatus (Bosc & Daudin, 1801) (L28075) 

For osteological observation, the skeleton of a female specimen from the 
Museum of Natural History of Geneva (MHNG 2620.032) was cleaned by Dermestes 
sp. larvae for one week, then whitened with H 2 2 . 

DNA extractions were made out of liver or muscle samples with the DNeasy 
Tissue kit (Qiagen) according to the manufacturer instructions. A 0.56kbp fragment of 
the 16S rDNA was amplified using the universal primers L2510 and H3059 designed 
by Palumbi et al. (1991). PCR conditions were as follows: Initial denaturation 94° for 
3 minutes, then (93°, 45 seconds; 55°, 45 seconds; 72°, 1 minute) x 37 cycles, and final 
extension 72° for 5 minutes. PCR products were checked for size on 1% agarose gel 
and purified on QIAquick columns (Qiagen). PCR fragments were directly sequenced 
for both strands using the BigDye cycle sequencing kit (Applied Biosystems), and an 
ABI 377 automated sequencer. Sequences were aligned with Sequencher™ v4.1.2 
(Gene Codes Corp.), and minor corrections were done by hand. The final matrix was 
analyzed with PAUP* v4.0bl0 (Swofford, 2002). Base composition heterogeneity was 
tested with the same software. Exhaustive parsimony analyses were performed on the 
whole matrix, and on a reduced dataset comprising the Chamaeleonidae only (without 
Sceloporus) with the following settings: Uninformative characters excluded, characters 
unordered and unweighted, gaps treated as missing, multistate treated as uncertainty. 
Branch support was estimated with lO'OOO bootstrap pseudoreplicates. 

RESULTS AND DISCUSSION 

MORPHO- ANATOMICAL AND ECOLOGICAL OBSERVATIONS 

Bradypodion spinosum is a dwarfed, short tailed, cryptically coloured, and rare 
inhabitant of the Afro-montane forests of the Eastern and Western Usambara 
Mountains of northern Tanzania (Fig. 1). The chameleons live in the lower strata of the 
deep forest biotope where they may be found at perch heights from a few cm to 3 
meters elevation. Over part of their distribution (in the Eastern Usambaras) B. spino- 
sum is sympatric with Rhampholeon temporalis. A superficial glance at this chameleon 
reveals some features that are not seen in the typical Rhampholeon. It has a network of 
interstitial skin that divides the flanks into clumps or islands of tubercles. In the genus 
Rhampholeon, this character is only seen in R. gorongosae Broadley, 1971. The 
tubercles on the flanks are not composed of sheets of stellate tubercles as is seen in 
other species of Rhampholeon, but rather of polygonal tubercles. The occiput of the 
skull has a rather prominent casque and the tail is relatively long for the typical 
Rhampholeon, averaging around 40% of the total length. It also has simple claws, a 
feature common to all species of the Chamaeleo and Bradypodion genera and to date 
only described in Rhampholeon temporalis. 



106 



C. R. TILBURY & J. MARIAUX 




Fig. 1 
Rhampholeon spinosus comb, n., female. Herkulu Tea Estate, Western Usambaras, Tanzania. 
(Photo. C. Tilbury) 

Bradypodion spinosum has however some morphological features unlike any 
seen in the genus Bradypodion (sensu lato) but reminding one of Rhampholeon (sensu 
lato). A clearly demarcated ridge of inter-orbital tubercles is present. In addition, the 
gular region is adorned with posteriorly diverging rows of low conical tubercles. 
Whilst there are a few species of Chamaeleo (e.g. tempeli, laterispinis) and one species 
of Rhampholeon {kerstenii) that demonstrate this gular arrangement, there are no mem- 
bers of the east African Bradypodion that have any trace of gular ornamentation. Closer 
examination of the scalation of the lower flanks shows that in areas there are small 
zones of stellate tubercles. In addition the rostral process is unlike any other rostral 
processes seen outside the Rhampholeon Type II group (Rieppel, 1987). Where they 
occur among the genus, the rostral process is formed by a cluster of small conical tu- 
bercles around a soft tissue cushion that projects forward off the snout as a flexible 
process. Occasional specimens of spinosum may also have weak axillary dermal 
dimples equivalent to the better formed mite pockets of other species of Rhampholeon, 
and finally all specimens examined have low palmar and plantar spines. These are only 
seen in the Rhampholeon type II species (sensu Rieppel, 1987). 

Klaver (1979) provided a valuable insight into the relationship of spinosum to 
the Brookesiinae when he commented that the simple sac-like a-diverticulate lungs of 
spinosum were "truly Rhampholeon like". A single male specimen of spinosum 
(MHNG 2620.036) was found with everted hemipenes. These demonstrate an acaly- 
culate hemipenal truncus and a complete absence of any apical structures - a condition 
not described outside the Brookesiinae genera. It is possible that the hemipenes are not 
fully everted and so further comment on the apex is withheld. 

OSTEOLOGICAL CHARACTERS 

The skull of Bradypodion spinosum demonstrated the following osteological 
features. The nasals are paired and form the dorsal margin of the external nares. The 
nasal process of the pre-maxilla meets the frontal bone just anteriorly to the nasals. The 
external nares reach the frontal bone. There is no fontanelle between the nasal and the 
prefrontal. The parietal is triangular and resembles that of Rhampholeon platyceps as 
illustrated by Rieppel (1987, Fig. 9E). The latter bone clearly lacks a supratemporal 



GENERIC ASSIGNMENT OF BRADYPODION SPINOSUM 107 

process, but the squamosal presents a normal dorsal process closing the upper temporal 
fossa posteriorly. These characters were all used by Rieppel (1987) to define taxa of the 
Rhampholeon type II group. The supratemporal itself is absent from the specimen we 
observed. Although this bone is present in most of the Rhampholeon type II species, 
exceptions occur like in R. spectrum (Rieppel, 1987). The presence of the splenial 
could not be ascertained. Additionally it was observed that the orbitae are bordered by 
three bones (i.e. including the frontal), a character typical of the Brookesiinae 
according to Klaver & Böhme (1986). 

Molecular phylogeny 

Partial DNA sequences from the 16S mitochondrial rDNA were obtained for 10 
taxa and aligned with an outgroup belonging to the Iguanidae (Phrysonomatinae). Out 
of 558 aligned characters, 62 were excluded for alignment difficulties and, among the 
496 remaining ones, 82 were parsimony informative (70 when Sceloporus was 
excluded). No compositional bias was detected in the dataset. Pairwise uncorrected 
distances between the outgroup and ingroup taxa vary between 22.3 and 25.8%. 
Interspecific distances within the ingroup fluctuate between 4.3 and 13.7%. There are 
5 substitutions (4 transitions and 1 transversion), representing 1 % divergence between 
both B. spinosum samples. The parsimony analysis of the 10 chameleons sequences 
resulted in a single most parsimonious tree (Fig. 2) with a length of 158 (CI 0.601, RI 
0.606). In this tree B. spinosum is the sister-group of Rampholeon sp. a new species 
from the neighbouring North Pare Moutains. This relationship is well supported. Both 
species forms a monophyletic group with two Rhampholeon (II) taxa although boot- 
strap support is much lower for this node. Interestingly this group of species includes 
R. spectrum, the type species of Rhampholeon. The other Rhampholeon (belonging to 
type I) group together, and so do the Bradypodion taxa, however all these nodes are 
weakly supported by bootstrap values. When the tree is rooted with the Sceloporus 
sequence, its topology remains the same and the root branches between the 
Rhampholeon (type II) and the Bradypodion/ Rhampholeon (type I) clades (Fig. 2). 
However, because of saturation concerns with the use of the available outgroup we 
prefer to discuss our results on the basis of an unrooted tree. In any case this does not 
change the relationships of B. spinosum with Rhampholeon. Despite the obvious cau- 
tion we should exercise because of the weak bootstrap support for some branches, 
these additional molecular results confirm the conclusions we derived from morpho- 
anatomical analyses. They allow us to obtain two important conclusions: a) B. spino- 
sum is definitely more closely related to (some) Rhampholeon species than to other 
Bradypodion, therefore its transfer to Rhampholeon is fully justified; and b) The genus 
Rhampholeon is most probably not monophyletic and comprises at least 2 separate 
lineages concordant with the Type I and Type II groupings. This idea has already been 
suggested by several authors on the basis of diverse morphological characters (Bauer, 
1997; Rieppel, 1987; Tilbury, 1992; Tilbury & Emmrich, 1996, Townsend & Larson, 
2002), and is now supported by a new class of characters. The taxonomic decisions and 
corresponding nomenclatural acts needed to reflect this situation will be published 
elsewhere on the basis of a more comprehensive dataset (Matthee et al., in prep), 
except for the new combination proposed below. 



108 C. R. TILBURY & J. MARIAUX 



B. spinosum 
B. spinosum { = R. spinosus comb. n. 

= R. spinosus comb. n. 



R. uluguruensis (II) 

R. sp, (!!) ^ 

/?. spectrum (II) 




R oxyrhinum . 

R. brevicaudatus (I) 

/?. kerstenii (I) 

ß. fischeri 

B. tavetanum 5 changes 

Fig. 2 
Unrooted phylogram showing the phylogenetic relationships of Rhamp ho! e on spinosus comb. n. 
based on a parsimony analysis of partial 16S rDNA sequences. Numbers on branches are boot- 
strap values (lO'OOO repeats). B = Bradypodion, R = Rhampholeon, (I)= Rhampholeon type I, 
(II)= Rhampholeon type II. The dotted line indicates the position of the root when Sceloporus 
undulatus is used as outgroup. 

CONCLUSIONS 

On the basis of its general phenotype, rostral process, simple lung morphology, 
presence of accessory palmar spines, row of enlarged inter-orbital tubercles, acalycu- 
late hemipenal truncus, osteological skull structure, and particular DNA characteris- 
tics, B. spinosum clearly does not belong in the genus Bradypodion (sensu lato), but 
rather in the genus Rhampholeon [Type II (Rieppel, 1984; Tilbury, 1992)]. Whilst it is 
apparent that spinosum belongs within the Brookesiinae subfamily, its relationships 
within the genus Rhampholeon are by no means clear yet. By virtue of its acalyculate 
hemipenal truncus it is likely to be more closely related to the species around R. 
boulengeri and uluguruensis. Compared to the sympatric species R. temporalis, the 
exaggerated development of the rostral process, the small gular spines and the flank 
scalation of spinosum, more reminiscent of the other chameleon genera, may indicate 
differentiation due to character displacement (Brown & Wilson, 1956). The gular 
spines are very reminiscent of those found in R. kerstenii. The flank scalation, and 



GENERIC ASSIGNMENT OF BRADYPODION SPINOSUM 109 

simple claws of spinosum are morphological features that appear to bridge the generic 
gap between Rhampholeon and Chamaeleo and may be important clues in the complex 
evolutionary history of the Chamaeleonidae. The molecular data reported here do not 
allow us to confirm this assumption, however preliminary analyses of more complete 
molecular datasets tend to support this idea (JM, unpublished data). 

We conclude with the transfer of B. spinosum to the genus Rhampholeon 
Günther, 1874, and propose the following new combination: Rhampholeon spinosus 
(Matschie, 1852) comb. nov. 

ACKNOWLEDGEMENTS 

We are grateful to the Government of the United Republic of Tanzania through 
its Commission for Science and Technology for the Research permits granted to JM 
(RCA 97/128), as well as to Kim Howell, Gamba Nkwengulila (Dar es Salaam) and 
Joe Berraducci (Arusha) for their support. Ivan Ineich (MNHN, Paris) kindly provided 
the R. spectrum tissue, and M. & A. Grimm, the R. kerstenii tissue. JM was supported 
by a grant of the Swiss National Research Foundation (3100-55710.98) and by the 
Museum of Natural History of Geneva. 

REFERENCES 

Bauer, A. M. 1997. Peritoneal pigmentation and generic allocation in the Chamaeleonidae. 
African Journal of Herpetology 46: 117-122. 

Broadley, D. G. & Howell, K. M. 1991. A checklist of the reptiles of Tanzania, with synoptic 
keys. Syntarsus 1: 1-70. 

Brown, W. L. & Wilson, E. O. 1956. Character displacement. Systematic Zoology 5: 49-64. 

Klaver, C. J. J. 1979. A review of Brookesia systematics with special reference to lung mor- 
phology. Bonner zoologische Beiträge 30: 163-175. 

Klaver, C. J. J. & Böhme, W. 1986. Phylogeny and classification of the Chamaeleonidae 
(Sauria) with special reference to hemipenis morphology. Bonner zoologische Mono- 
graphien 22: 1-64. 

Matthee, C. A., Tilbury, C. R. & Townsend, R. A phylogenetic review of the African genus 
Rhampholeon (Chamaeleonidae: Brookesiinae) with notes on the environmental factors 
responsible for diversification within the genus. (In prep). 

Matschie, P. 1892. Über eine kleine Sammlung von Säugethieren und Reptilien, welche Herr L. 
Conradt aus Usambara (Deutsch-Ostafrika) heimgebracht hat. Sitzungsberichte der 
Gesellschaft naturforschender Freunde zu Berlin 7: 101-110. 

Mertens, R. 1966. Chamaeleonidae. Das Tierreich, Berlin 83: 1-37. 

Palumbi, S., Martin, A., Romano, S., McMillan, W. O., Stice, L. & Grabowski, G. 1991. The 
simple fool's guide to PCR, version 2.0. University of Hawaii, Honolulu. 

Rieppel, O. 1987. The phylogenetic relationships within the Chamaeleonidae, with comments on 
some aspects of cladistic analysis. Zoological Journal of the Linnean Society 89: 41-62. 

Swofford, D. L. 2002. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). 
Version 4. Sinauer Associates, Sunderland, Massachusetts. 

Tilbury, C. R. 1992. A new dwarf forest chameleon (Sauria: Rhampholeon Günther 1874) from 
Malawi, Central Africa. Tropical Zoology 5: 1-9. 

Tilbury, C. R. & Emmrich, D. 1996. A new dwarf forest chameleon (Squamata: Rhampholeon 
Günther 1 874) from Tanzania, East Africa with notes on its infrageneric and Zoogeo- 
graphie relationships. Tropical Zoology 9: 61-71. 



1 10 C. R. TILBURY & J. MARIAUX 



Townsend, T. & Larson, A. 2002. Molecular phylogenetics and mitochondrial genomic evolu- 
tion in the Chamaeleonidae (Reptilia, Squamata). Molecular Phylogenetics and 
Evolution 23: 22-36. 
Uetz, P. 2003. The EMBL reptile database, [http://www.reptile-database.org] 
Werner, F. 1902. Prodromus einer Monographie der Chamäleonten. Zoologische Jahrbücher 
(Systematik) 15: 295-460. 



Revue suisse de Zoologie 111 (1): 111-120; mars 2004 



Redescription of Brooksiella praeputialis and Goezeella siluri 
(Eucestoda: Proteocephalidea), parasites of Cetopsis coecutiens 
(Siluriformes) from the Amazon and proposition of 
Goezeella danbrooksi sp. n. 

Alain DE CHAMBRIER K Amilcar A. REGO 2 & Jean MARIAUX ! 

1 Muséum d'histoire naturelle, Département des Invertébrés, PO Box 6434, 
CH-1211 Geneva 6, Switzerland. 

2 Institute Oswaldo Cruz, Department of Helminthology, PO Box 926, 
21045-900, Rio de Janeiro, Brazil. 

E-mail : alain.dechambrier@mhn.ville-ge.ch 



Redescription of Brooksiella praeputialis and Goezeella siluri (Eu- 
cestoda: Proteocephalidea), parasites of Cetopsis coecutiens (Siluri- 
formes) from the Amazon, and proposition of Goezeella danbrooksi 

sp. n. - Brooksiella praeputialis (Rego, Santos & Silva, 1974) and Goe- 
zeella siluri Fuhrmann, 1916 from the Amazonian siluriform fish Cetopsis 
coecutiens (Siluriformes: Cetopsidae) are redescribed. Goezeella piramutab 
Woodland, 1933 is considered as a synonym of Goezeella siluri. Goezeella 
siluri sensu Brooks & Rasmussen, 1984 becomes Goezeella danbrooksi 
sp. n., differing from G siluri by the vitelline follicles position (only dorsal, 
not lateral), size (much smaller in type material) and shape (slightly wider 
posteriorly); by the position of vaginal sphincter (which is terminal in 
G siluri) and by the number of testes. 

Key- words: Eucestoda - Monticelliidae - Brooksiella praeputialis - 
Goezeella siluri - Goezeella danbrooksi sp. n. - catfish parasites - taxonomy. 

INTRODUCTION 

Two proteocephalidean tapeworms, Goezeella siluri Fuhrmann, 1916 and 
Brooksiella praeputialis (Rego, Santos & Silva, 1974) have been described from the 
"candiru", Cetopsis coecutiens (Cetopsidae) from the Amazon River. During a field 
expedition to Amazonia conducted by the Natural History Museum, Geneva (MHNG) 
and the Institute Oswaldo Cruz (IOC), we were able to collect both cestodes species in 
excellent condition. Due to some confusion in citations or redescription of both species 
(Woodland, 1933; Freze, 1965; Rego et al, 191 A; Rego, 1975), we give here a brief 
redescription of Brooksiella praeputialis and Goezeella siluri, and clarify a number of 
issues linked to the synonymy of these taxa. 



Manuscript accepted 21.08.2003 



112 A. DE CHAMBRIER ETAL. 

MATERIAL AND METHODS 

The hosts were collected by fishermen from the Amazon river near Itacoatiara, 
Brazil, in 1992 and 1995; they were dissected and examined for parasites immediate- 
ly after death. The gut was dissected along its entire length. The worms were fixed 
immediately after dissection with hot 4% neutral formaldehyde solution, stained with 
Mayer's hydrochloric carmine solution, dehydrated in an ethanol series, cleared with 
eugenol (clove oil) and mounted in Canada balsam. Pieces of the strobila were em- 
bedded in paraffin wax, cross sectionned (thickness 12-15 urn), stained with Weigert's 
hematoxylin and counterstained with 1% eosin B (de Chambrier, 2001). Eggs were 
studied in distilled water. All measurements are given in micrometers (um) unless 
otherwise stated. 

Abbreviations used in descriptions are as follows: x = mean, n = number of 
measurements, CV = coefficient of variability, OV = ovary width versus proglottis 
width ratio (in %), PP = position of genital pore (cirrus pore) as % of proglottis length, 
PC = cirrus-pouch length versus proglottis width ratio (in %). CHIOC = the 
Helminthological Collection of the Institute Oswaldo Cruz; MHNG = Natural History 
Museum, Geneva, INVE = Geneva Museum Invertebrate Collection, USNM = U.S. 
National Museum. 

RESULTS 

Brooksiella praeputialis (Rego, Santos & Silva) Figs 1-5, 9-10 

Amphoteromorphus praeputialis Rego, Santos & Silva, 1974: 200. 
Amphoteromorphus praeputialis; Brooks & Rasmussen, 1984: 751. 
Brooksiella praeputialis; Rego, Chubb & Pavanelli, 1999: 354. 

Type-host: Cetopsis coecutiens Spix & Agassiz (Siluriformes: Cetopsidae); vernacular 
name : Candiru. 

Other host: Pseudocetopsis othonops Eigenmann (Siluriformes: Cetopsidae). 

Materiel studied: Brooksiella praeputialis (Rego, Santos & Silva, 1974), syntypes 
CHIOC 26.421, 31.047a-d, 31.048 a-c and 26.524. 

Other material : Itacoatiara, Amazonas State, Brazil, MHNG INVE 19357 (14.09.1992); 
INVE 21875, 21876 (07.10.1995); INVE 22059, 22088, 21994, 21995, 21996 (16.10.1995). 

Site of infestation: Intestine. 

Type locality: Rio Amazonas, Maicuru, Para State, Brazil. 

Distribution: Amazon River. 

Redescription (based on 10 specimens) 

Proteocephalidae, Zygobothriinae. Cestodes up to 12-22 mm long, 1,785 wide, 
with numerous transverse grooves in immature and mature proglottides, less marked in 
gravid proglottides. Strobila acraspedote, anapolytic, consisting of 46 to 51 (n = 7) 
proglottides : 25-36 immature (up to appearance of spermatozoa in vas deferens), 2-4 
mature (up to appearance of eggs in uterus), 7-15 pregravid (up to appearance of hooks 
in oncospheres), and up to 8 gravid proglottides. Proliferation zone very short. 

Immature, mature, pregravid proglottides wider than long (Fig. 3), gravid 
proglottides slightly wider than long or longer than wide. Metascolex slightly conical 
910-1370 (x = 1145, n = 10) in diameter, much wider than neck (Figs 1, 9-10), with 
antero-laterally directed, embedded, uniloculate suckers, 275-395 (x = 325, n = 40, CV 



REDESCRIPTION OF BROOKSIELLA PRAEPUTIALIS 1 1 3 

= 9%) in diameter. Suckers with a well-developed circular musculature situated in 
upper marginal region (Fig. 1). External surface of metascolex bearing numerous 
wrinkles (Figs 9-10). 

Internal longitudinal musculature weakly developed, represented by fine 
bundles of separated muscular fibres (Figs 4-5) forming anastomoses. Osmoregulatory 
canals overlapping vitelline follicles and testes (Figs 3-5). Ventral canals wider than 
dorsals. Presence of a posterior transverse canal and of secondary canals situated 
posteriorly and ending laterally beneath the tegument. Within scolex, canals form a 
dense network (Fig. 1). 

Testes medullary, forming 2 compact fields in two or three layers, not overlap- 
ping cirrus sac, vagina and vas deferens (Fig. 3), numbering 193-292 (x = 244, n = 5, 
CV = 19%); aporal testes 98-150 in number (x = 123); 17-25 preporal testes (x = 21); 
141 postporal testes 75-122 (x = 98). Testes spherical to oval, 45-60 in diameter, 
representing 2.5 - 4% of proglottis width. Testes reaching laterally almost the margin 
of proglottides, but not reaching not the posterior one (Fig. 3), overlapping vitelline 
follicles, present also in gravid proglottides. 

Cirrus sac elongate to pyriform, with tapered distal part, thick- walled, 250-385 
long and wide (Fig. 3); PC 15-23% (x =19 %, n = 36, CV = 11%). Cirrus up to 65% of 
cirrus pouch length. Vas deferens strongly coiled, situated between proximal part of 
cirrus sac and midline of proglottides, occupying up to 32 % of proglottis width, wider 
in its basal part (Fig. 3). 

Genital atrium deep, pores alternating irregularly, situated anteriorly; PP = 12- 
26% (x = 18%, n = 24). Genital ducts passing between osmoregulatory canals. 

Ovary medullary, bilobate, highly follicular in pre-gravid proglottides (Figs 
3-4), with numerous dorsal and ventral outgrowths scarcely reaching dorsal cortex 
(Fig. 4), 880-1210 wide, OV =55-69 % (x = 59%; n = 34; CV = 7%). Mehlis' glands 
100-150 in diameter (Fig. 3). 

Vitelline follicles cortical, arranged in two lateral ventral field near margins of 
proglottides, more numerous posteriorly, occupying almost all proglottis length, not 
interrupted at level of cirrus sac ventrally, with some isolated follicles dorsal to cirrus 
sac. Vitelline fields of ventral follicles overlapping testes (Figs 3-5). 

Vaginal canal forming small seminal receptacle antero-dorsal to ovarian isth- 
mus, terminal part of vaginal canal (pars copulatrix vaginae) muscular (Fig. 3). Vagina 
anterior 26(%) or posterior 74(%) (n = 126) to cirrus-sac. Vaginal canal strongly curved 
before reaching seminal receptacle. 

Primordium of uterine stem medullary, present in immature proglottides. 
Formation of uterus of type 1 (see de Chambrier et al, 2004b): in immature proglot- 
tides, uterine stem straight, occupying entire length of proglottis, formed by longitu- 
dinal thick column of chromophilic cells along midline of proglottides. Lumen of 
uterus appearing in first mature proglottides (Figs 3-5); diverticula (lateral branches) 
formed before first eggs appear in uterine stem. In pregravid proglottides, eggs 
completely filling uterine stem and thick-walled diverticula. In gravid proglottides, 
diverticula occupying up to 62 % of proglottis width; 9-17 (n = 26) medullary lateral 
branches on each side. Longitudinal ventral uterine opening observable in some 
terminal proglottides. 



114 



A. DE CHAMBRIER ETAL. 




Figs 1-2 
Brooksiella praeputialis (Rego, Santos & Silva, 1974), INVE 22059. 1. Frontal section of the 
scolex, showing the concentration of gland(?) cells and dense network of osmoregulatory canals 
beneath the apical part. Note also the muscular fibres situated in the anterior part of the sucker. 
2. Eggs drawn in distilled water. Abbreviations: cm, internal circular musculature; em, em- 
bryophore; gc, gland cells; oc, osmoregulatory canals; oe, outer envelope; on, oncosphere. Scale- 
bars: 1 = 500 urn; 2 = 20 urn. 

Eggs with two polar projections 15- 20 long (Fig. 2). Embryophore spherical, 
bilayered, 25-29 x 22-24; internal layer containing granular material and paired polar 
chambers, 22-26 x 13-16; oncosphere spherical to oval, 10-13 in diameter, with 3 pairs 
of hooks, 4-5 long (Fig. 2). 



Remarks 

Brooks & Rasmussen (1984) commented on the confusion of figures by Rego 
et al. (1974). We re-examined the original material and agree with Brooks & 
Rasmussen: figures 23, 24, 26 and 27 of Rego et al. (1974) are of B. praeputialis (No 
CHIOC 31047a, 31047d respectively) and not Monticellia siluri as stated. Figures 
30-32 (CHIOC 31048a) and figure 33 (CHIOC 31048c) correspond to the syntype 
material of B. praeputialis. 



REDESCRIPTION OF BROOKSIELLA PRAEPUTIALIS 



15 




Figs 3-5 
Brooksiella praeputialis (Rego, Santos & Silva, 1974). 3. INVE 21996, mature proglottis, dor- 
sal view. 4-5. INVE 22059, pregravid proglottis, cross sections at level of ovary and at level of 
middle of the proglottis. Note the follicular structure of ovary and the lateral and ventral posi- 
tion of vitelline follicles. Abbreviations: do: dorsal osmoregulatory canal; gm, Mehlis glands; lm, 
internal longitudinal musculature; ov: ovary: po : transverse osmoregulatory canal; sc : second- 
ary canal; te: testes; ut: uterus; vc: vaginal canal; vi: vitellaria; vo: ventral osmoregulatory canal. 
Scale-bar: 500 um. 



Goezeella siluri Fuhrmann 



Figs 6-8 



Goezeella siluri Fuhrmann, 1916: 386. 

Goezeella piramutab Woodland, 1933: 488, syn. n., see discussion below. 
Monticellia siluri; Woodland, 1925: 714. 
Goezeella siluri; Freze, 1965: 508. 
Monticellia siluri; Rego, Santos & Silva, 1974: 195. 
Goezeella siluri; Brooks & Deardorff, 1980: 15. 
Goezeella siluri; Rego, Chubb & Pavanelli, 1999: 348. 

[Not Goezeella siluri; Brooks & Deardorff, 1980: 15]. See Goezeella danbrooksi sp. n., de- 
scribed below. 
[Not Goezeella siluri; Brooks & Rasmussen, 1984: 750]. See discussion below. 



Type-host: Cetopsis coecutiens (Siluriformes: Cetopsidae). 

Other host: Pinirampus pirinampu (Spix &Agassiz) (Siluriformes: Pimelodidae). 



116 



A. DE CHAMBREER ETAL. 




Figs 6-8 
Goezeella siluri Fuhrmann, 1916. 6. Scolex, type material showing the biloculate suckers. 7. 
INVE 19858, egg drawn in distilled water. 8. Cross section at level of ovary, showing the ven- 
tral position of vitelline follicles, after Fuhrmann (1931, Fig. 275). Abbreviations (according to 
Fuhrmann): Im, internal longitudinal musculature; n, longitudinal nerve; ve, ventral osmoregu- 
latory canal. Scale-bars: 6 = 1000 um: 7 = 20 um (8 = no scale). 

Materiel studied: Goezeella siluri, syntype material, 2 whole mount slides, 14 cross- 
sections slides, Coll. Institut de Zoologie, Neuchâtel. Other material: Itacoatiara, Amazonas 
State, Brazil, MHNG FNVE 19858, 13.09.1992. 

Site of infestation: Intestine. 

Type locality: "Amazonas", Brazil. 

Distribution: Amazon River. 



Description 

Strobila somewhat contracted, 77 mm long, 2025 in width. About 47 immature, 
10 mature and 143 (pregravid and gravid) proglottides. Scolex with metascolex, 1825 
in diameter, with biloculated suckers 435 long and 300 width (Fig. 6). Internal longi- 
tudinal musculature developed, represented by fine bundles of separated muscular 
fibres (Fig. 8) forming anastomoses. 



REDESCRIPTION OF BROOKSIELLA PRAEPUTIALIS 



117 




Figs 9-10 
Bwoksiella praeputialis (Rego, Santos & Silva, 1974), INVE 22088. Scanning electron micro- 
graphs. 9. Scolex, apical view. 10. Scolex, lateral view. Scale-bar: 100 pm. 



About 380-433 small testes in one cortical dorsal field and in one layer. Deep 
genital pore irregularly alternating, anterior, PP = 10-16%. Cirrus pouch elongated to 
pyriform, PC = 13-17% (x = 14%, n = 9), cirrus occupying between 30-50% of cirrus 
pouch length. Vagina always anterior to cirrus pouch (n = 36), with powerful muscular 
terminal sphincter. Ovary cortical, bilobate, follicular, with numerous dorsal out- 
growths, OV = 75-79%. 

Ventral cortical vitelline follicles lateral, occupying large triangular field, wider 
posteriorly, overlapping ovary and cirrus pouch, running from anterior to posterior 
margin of proglottis. Ventral cortical uterus of type 2 (see de Chambrier et al., 2004b, 
for definition), with 14-21 thin digitate diverticula with thick walls. Eggs oval (in 
whole mounts), embryophore bilayered, about 20 x 16; oncosphere spherical to oval, 
about 10 in diameter (Fig. 7). 

Goezeella danbrooksi sp. n. 

Goezeella siluri; Brooks & Deardorff, 1980: 15. 
[Not Goezeella siluri Fuhrmann, 1916: 386]. 

Type-host: Ageneiosus cancanus (Siluriformes: Ageneiosidae). 

Holotype (examined): 1 unique specimen with some proglottides cut in serial sections, 
USNM Helminthological collection No. 74498 (2 slides), No. 74544, (19 slides, serial sections). 
In our knowledge, no other material of this species cited in literature. 

Site of infestation: Small intestine near juncture of stomach and intestine. 

Type locality: Magdalena River, vicinity of San Cristobal, Bolivar Province, Colombia. 

Remarks 

Brooks & Deardorff (1980) studied, in a detailed description, Goezeella siluri 
in Ageneiosus cancanus Steindachner from Columbia. De Chambrier & Vaucher 
(1999, p. 184) noted that the species described by these authors apparently did not 
belong to G. siluri. When looking at figures 2 to 5 of Brooks & Deardorff (1980), we 
observed that their material differed from the type material of G. siluri (see Fig. 8, 



118 A. DE CHAMBRIER ET AL. 

present work) in several aspects: the position of the vitelline follicles (only dorsal, not 
lateral), relative size (much smaller in type material), and shape (slightly wider 
posteriorly, see Fuhrmann, 1916, Fig. 4). Goezeella siluri sensu Brooks & Deardorff 
also differs in the position of vaginal sphincter (terminal in G siluri), by the size of the 
cirrus / cirrus pouch ratio, and by the number of testes (183-380 versus 380-433). 
Furthermore, the taxon described by Brooks & Deardorff parasitizes a different family 
of hosts (Ageneiosidae). 

In our opinion, Goezeella siluri sensu Brooks & Deardorff represents a distinct 
species and we name it Goezeella danbrooksi sp. n. in honour of Prof. Daniel R. 
Brooks from Toronto, Canada. 

DISCUSSION 

Fuhrmann (1916) in his description of G siluri, observed oval eggs without 
polar projections. Woodland (1933) in his expedition in Amazonian river in 1931, 
found in the type host Cetopsis coecutiens (field number Amaz 174) twenty three 
worms he determined as G siluri. He first thought that he had two different species, as 
six specimens fitted Fuhrmann's description (thick proglottides, similar transverse 
sections) but the remaining specimens included much thinner proglottides. He finally 
decided to consider that all the material belonged to a single species because the "eggs 
of both types are identical in shape and size". In fact, Woodland observed and drew 
only those from B. praeputialis (his Fig. 18, compare with Fig. 2, present work). 

The material described by Woodland (1933) as Goezeella siluri (BMNH 
1964.12.15.174-183) is in fact a mixture of two species: G. siluri Fuhrmann, 1916 with 
biloculate suckers (Fig. 6, present work) and Brooksiella praeputialis (Rego, Santos & 
Silva, 1974) with uniloculate suckers (Figs 1, 9-10, present work). The shape described 
by Woodland (1933) as "cobra like" due to the hood shaped neck (Woodland's Fig. 12) 
corresponds with Brooksiella praeputialis (Woodland's plate 30, Figs 10, 11, 12, 14, 
17, 18) and the other figures correspond with Goezeella siluri (Figs 9, 13, 16). Of his 
six whole mounted slides, three are a mixture (slides number 1, 3, 6), one is G. siluri 
(slide number 4) and two are B. praeputialis (slides number 2, 5). The cross sections 
numbered G' 1-6 and G" 1-3 are of G. siluri (Woodland's Fig. 16) and those G 1-2, Gc 
1-7 and G'" 1-2 correspond to B. praeputialis (Woodland's Fig. 17). 

In the same paper, Woodland (1933) considered Goezeella piramutab, a para- 
site of Brachyplaty stoma vaillanti, as a new species. He based his decision to separate 
G. piramutab from G. siluri only on the differences on eggs shape and size. He 
suggested that a complete diagnosis must await the examination of further material. In 
fact, Woodland equivocally compared eggs from B. praeputialis (Woodland's Fig. 18) 
and eggs of G piramutab (Woodland's Fig. 22). In our opinion, the eggs figured by 
Woodland (Fig. 22) belong to Goezeella siluri (see Fig. 7, present work). The other 
eggs (Woodland's Fig. 18) are typical of B. praeputialis (compare with Fig. 2, present 
work). As the argument to separate G. piramutab from G siluri by their eggs shape and 
size is no longer valid, we consider Goezeella piramutab as a synonym of Goezeella 
siluri. 

Brooks & Rasmussen (1984) found a cestode they determined as B. praeputialis 
in a new host, Pseudocetopsis othonops in Venezuela but did not give a description. 



REDESCRIPTION OF BROOKSIELLA PRAEPUTIALIS 1 19 

Judging by their figures (5, 6, 7), we believe that it may correspond to another species 
: the diameter of scolex is smaller (705 urn versus 1145 urn in mean), the suckers seem 
to be deeply embedded and the testes are less numerous according to their figures 
(6-7, p. 751) (64-178 versus 205 - 342). 

The shape of the ovary of Brooksiel la praeputialis is particular. It is highly folli- 
cular and is similar to that of Goezeella siluri. This is an unusual observation for the 
Proteocephalidea, although this ovarian shape is also present in Manaosia braco- 
democa Woodland, 1935 and Electrotaenia malapteruri (Fritsch, 1886) (see de 
Chambrier, 2003; de Chambrier et al, 2004a). 

Brooksiella praeputialis shows some similarities with the species of Rudol- 
phiella: a metascolex, a wide band of ventral vitelline follicles, a follicular ovary, type 
1 uterine formation and eggs with a similar structure including polar formations and 
bilayered embryophores. Furthermore, in a recent molecular analysis, two Rudol- 
phiella species clustered with Brooksiella praeputialis in a strongly supported clade 
(see de Chambrier et al., 2004b: fig. 1. Rudolphiella possesses cortical vitelline 
follicles and testes, and medullar ovary and uterus, whereas in Brooksiella only the 
vitelline follicles are cortical. Both genera belong to two different subfamilies, 
Rudolphielliinae and Zygobothriinae, which are characterised by a different arran- 
gement of the genital organs relative to the internal longitudinal musculature. However 
we have recently expressed doubts about the importance of this type of character and 
have suggested that other characters may prove to be useful in suprageneric classi- 
fications of the Proteocephalidea (de Chambrier et al, 2004b). 

ACKNOWLEDGEMENTS 

The authors thank David I. Gibson and Eileen Harris for loan of material. They 
also thanks anonymous referees who provided judicious comments on an earlier 
version of this manuscript, Jean Wuest (Geneva) for the SEM photography, Gilles Roth 
(Geneva) for finalising the drawings and the fishermen in Itacoatiara, whose colla- 
boration in providing freshwater fishes was greatly appreciated. 

REFERENCES 

Brooks, D.R. & Deardorff, T.L. 1980. Three proteocephalid cestodes from Columbian siluri- 
form fishes, including Nomimoscolex alovarius sp. n. (Monticelliidae: Zygobothriinae). 
Proceedings of the Helminthological Society of Washington 47: 15-21. 

Brooks, D.R. & Rasmussen, G. 1984. Proteocephalid cestodes from Venezuelan catfish, with a 
new classification of the Monticelliidae. Proceedings of the Biological Society of 
Washington 97: 748-760. 

de Chambrier, A. 2001. A new tapeworm from the Amazon, Amazotaenia yvettae n. gen., n. sp., 
(Eucestoda: Proteocephalidea) from the siluriform fishes Br achyplaty stoma filamento- 
sum and B. vaillanti (Pimelodidae). Revue suisse de Zoologie 108 (2): 303-316. 

de Chambrier, A. 2003. Redescription of Manaosia bracodemoca Woodland, 1935 and 
Paramonticellia itaipuensis Pavanelli & Rego, 1991 (Eucestoda, Proteocephalidea) par- 
asites of Sorubim lima (Siluriforme, Pimelodidae). Folia Parasitologica 50: 121-127. 

de Chambrier, A. & Vaucher, C. 1999. Proteocephalidae et Monticelliidae (Eucestoda: 
Proteocephalidea) parasites de poissons d'eau douce du Paraguay avec descriptions d'un 
genre nouveau et de dix espèces nouvelles. Revue suisse de Zoologie 106(1): 165-240. 



120 A. DE CHAMBRER ETAL. 



de Chambrter, A., Scholz, T. & Ibraheem, M. H. 2004a. Redescription of Electrotaenia 

malopteruri (Fritsch, 1886) (Cestoda: Proteocephalidae), a parasite of Malapterurus 

electricus (Siluriformes: Malapteruridae) from Egypt. Systematic Parasitology 57(2): 

97-109. 
de Chambrier, A., Zehnder, M. P., Vaucher, C. & Mariaux, J. 2004b. The evolution of the 

Proteocephalidea (Platyhelminthes, Eucestoda) based on an enlarged molecular phy- 

logeny, with comments on their uterine development. Systematic Parasitology 57(3). [In 

press]. 
Freze, V.l. 1965. Essentials of cestodology. Vol. V Proteocephalata in fish, amphibians and 

reptiles. Moski'a: Izdatel'stvo "Nauka", 538 pp. (In Russian: English translation, Israel 

Program of Scientific Translation, 1969, Cat. No. 1853, v + 597pp). 
Fuhrmann, O. 1916. Eigentümlische Fischcestoden. Zoologischer Anzeiger 46(13): 385-398. 
Fuhrmann, O. 1931. Dritte Klasse des Cladus Plathelminthes. Cestoidea. In: Kükenthal & 

Krumbach. Handbuch der Zoologie. Walter de Gruyter & Co., Berlin und Leipzig. Vol. 

2: 141-416. 
Rego A.A. 1975. Estudos de cestóides de peixes do Brasil. 2. a nota: revisào do gènero Monti- 

cellia La Rue, 1911 (Cestoda, Proteocephalidae). Revista brasileira de Biologia 35(3): 

567-586. 
Rego, A.A., Chubb, J.C., Pavanelli, G.C. 1999. Cestodes in South American freshwater fishes: 

keys to genera and brief description of species. Revista brasileira de Zoologia 16: 

299-367. 
Rego, A.A., Dos Santos, J.C. & Silva, P.P. 1974. Estudos de cestóides de peixes do Brasil. 

Memorias do Instituto Oswaldo Cruz (Rio de Janeiro) 72: 187-204. 
Woodland, W.N.F. 1925. On some remarkable new Monticellia-like and other cestodes from 

Sudanese siluroids. Quarterly Journal of Microscopical Science 69(4): 703-729. 
Woodland, W.N.F. 1933. On two new cestodes from the Amazon siluroid fish Brachy- 

platy stoma vaillanti Cuv. and Val. Parasitology 25: 485-490. 



Revue suisse de Zoologie 111 (1): 121-137; mars 2004 



Salaria economidisi, a new species of freshwater fish from 
Lake Trichonis, Greece, with comments on variation in 
S. fluviatilis (Teleostei: Blenniidae) 

Maurice KOTTELAT 

Route de la Baroche 12, Case postale 57, CH-2952 Cornol, Switzerland 
(address for correspondence), and Department of Biological Sciences, 
National University of Singapore, Kent Ridge, Singapore 119260. 
E-mail: mkottelat@dplanet.ch 

Salaria economidisi, a new species of freshwater fish from Lake 
Trichonis, Greece, with comments on variation in S. fluviatilis 
(Teleostei: Blenniidae). - Salaria economidisi, new species, is distin- 
guished from S. fluviatilis, its hypothesised closest relative, in having a 
longer head, more teeth in both jaws, a simple orbital cirrus, a much shorter 
lateral line, and a distinctive colour pattern of 3-5 rows of bold black dots 
on the cheek. 

Key-words: Blenniidae - Salaria - Greece - Lake Trichonis - taxonomy. 

INTRODUCTION 

The family Blenniidae includes a large number of small benthic fishes. The vast 
majority are marine and occur in tropical areas. A few species are known to enter 
brackish or fresh waters in Southeast Asia (e.g. Phenablennius heyligeri (Bleeker, 
1859); see Springer & Smith-Vaniz, 1972: 70), India, Africa (Omobranchus species; 
see Springer & Gomon, 1975), and South America (e.g., Lupinoblennius paivai (Pinto, 
1958); see Sazima & Carvalho-Filho, 2003). The genus Salaria, as usually recognised, 
includes 3 species of which S. fluviatilis (Asso, 1801) is known only from freshwaters. 
It has been recorded under a variety of synonyms (e.g., Blennius vulgaris Pollini, 1816, 
Salarias varus Risso, 1827, B. cagnota Valenciennes, 1836, B. lupulus Bonaparte, 
1840, B. alpestris Blanchard, 1866; see Kottelat, 1997) from drainages around the 
northern Mediterranean shore from Israel to Portugal and in Morocco and Algeria. 

Salaria fluviatilis inhabits portions of streams with swift water and rocks 
varying in size from small stones to large boulders (Freeman et al, 1990); it also 
occurs on rocky shores of lakes (Elmiger, 2002). It spawns under stones and the male 
defends the eggs (Wickler, 1957). The larvae are pelagic (as are those of marine 
blennies) and the young settle in benthic habitats when they reach about 15 mm. The 
pelagic larvae apparently are one of the factors limiting the distribution of the species. 
Larvae from riverine populations are drifted by the current and need to reach a quiet 
section of the stream or a lake where they can spend their pelagic time, and then need 



Manuscript accepted 03.11.2003 



122 M. KOTTELAT 

to be able to return to the riverine stretch. Although sometimes stated in the literature, 
there is no evidence that this pelagic period can be spent in estuaries or brackish water. 
In the Mediterranean climatic zone, streams and rivers severely shrink in summer, and 
many survive only as a succession of pools connected by trifles of water. Water in these 
pools is moving very slowly, probably providing suitable habitats for the larvae. 

All freshwater populations of European blennies have been referred to S. fluvia- 
tilis (Kottelat, 1997; Perdices et al, 2000), but they have rarely been compared. 
Examination of the population from Lake Trichonis (a large natural lake in western 
Greece) shows that it is a species with a distinctive morphology, colour pattern and 
ecology. The species is described here and some populations of 5. fluviatilis are 
discussed. 

The generic nomenclature of Bath (1977, 1996) is followed here. Bock & 
Zander (1986: 142) united the genera Lipophrys and Salaria and used Lipophrys Gill, 
1896 as the valid name. As discussed by Kottelat (1997), Salaria Forsskâl, 1775 is the 
oldest valid name. The argument of similarity of the generic names Salaria and 
Salarias Cuvier, 1816 is irrelevant to the discussion (ICZN art. 56.2 is explicit). 

MATERIAL AND METHODS 

All measurements were made point to point with a dial calliper. Standard length 
is measured from the tip of the upper lip to the end of the hypural complex, total length 
to the posteriormost point of the caudal fin, head length to the posterior most point of 
the opercle, pre-anal length to the origin of the first anal spine. Body depth is measured 
at the origin of dorsal fin, depth of caudal peduncle at the narrowest point of the cau- 
dal peduncle. Length of caudal peduncle is measured from the base of the last anal-fin 
ray to the end of the hypural complex, at mid-height. Snout length is measured from 
tip of upper lip to the nearest point of margin of orbit. Eye diameter is the greatest 
diameter of the eye ball. The eye-rictus distance is measured from the posterior 
extremity of upper lip to the nearest point of the orbit. Interorbital width is the width 
of the bony area. Terminology of cephalic lateral line canals is derived from that of 
Bath (1977). Lateral line pore count starts with the pore immediately posterior to the 
upper extremity of the gill opening. Sequence of description generally follows the tem- 
plate of Springer & Williams (1994). Notation for dorsal and anal fin ray counts 
follows Hubbs & Lagler (1947). 

Examined material is in: MHNG, Muséum d'histoire naturelle, Genève; CMK, 
author's collection. 

DESCRIPTION 

Salaria economidisi sp. n. Figs 1-2 

Holorvpe. MHNG 2641.89, 60.8 mm SL; Greece: Lake Trichonis east of Panetolio; 
38°35'28.1"N 21°27'51.2"E; M. Kottelat & P. S. Economidis. 6-7 October 2001. 

Paratopes. MHNG 2641.90, 10; CMK 16968, 158; 19.5-55.9 mm SL; same data as holo- 
type. - CMK 13428, 5, 29.9-44.0 mm SL; Lake Trichonis; P. S. Economidis, 1 Nov 1996. 

Diagnosis. Salaria economidisi is distinguished from the marine S. paro (Risso, 
1810) and S. basilisca (Valenciennes, 1836) in having XII-XIII,16-17 dorsal rays (vs. 
XII,22-25) and 11,18-19 anal rays (vs. 11,23-26; Bath, 1977; Whitehead et al, 1986). It 



SALARIA ECONOMIDISI 



123 



Fig. 1 
Salaria economidisi; 
Greece: Lake Tricho- 
rds, a, paratype, fe- 
male, 48.4 mm SL, 
CMK 16968; b, male, 
holotype, 60.8 mm 
SL, MHNG 2641.89; 

c, male, paratype, 50.9 
mm SL, CMK 16968; 

d, male, paratype, 55.9 
mm SL, CMK 16968. 



Fig. 2 
Salaria economidisi; 
Greece: Lake Tricho- 
rds, eastern end, at 2- 
3 m depth on gently 
sloping sand-gravel 
beach; August 1995 
(photograph by U. 
Schliewen). 







124 M. KOTTELAT 

is distinguished from the freshwater S. fluviatilis in having a longer head (29.1-33.1 % 
SL, vs. 25.3-30.6), 25-30 teeth in the upper jaw (vs. 16-24), 20-27 in the lower jaw (vs. 
16-20), orbital cirrus simple (vs. branched), anterior continuous canal of lateral line 
with 5-9 simple pores (vs. usually 12-22, rarely 6-8) continued by a series of 1-3 short 
bi-pore tubes (vs. 4-12), and a pattern of 3-5 rows of bold black dots on the cheek, run- 
ning obliquely from eye backwards and downwards, the space between the lower two 
rows being greyish and spaces between the upper rows paler (vs. cheek with a broad 
diagonal band of tiny dots from lower edge of eye downwards backwards; Figs 3, 4a). 
Description. Selected morphometric data of holotype and 9 paratypes in 
Table 1. Dorsal fin. XII-XIII,16-17 (XIII, 16), modally XIII, 16, total 28-30; XIV spines 
in a single specimen out of 3 1 specimens examined for this character. Spinous portion 
somewhat shallower than segmented-ray portion; posterior membranous attachment on 
dorsal edge of caudal peduncle anterior to caudal-fin base. 

Table 1. Morphometric data of holotype (60.8 mm SL) and 9 paratypes oï Salaria economidisi. 
M: males, F: females. 





M 


M 


M 


M 


M 


M 


M 


M 


F 


F 


SL(mm) 


60.8 


59.9 


52.1 


50.9 


49.7 


47.8 


42.8 


39.9 


48.4 


41.3 


TL (mm) 


73.7 


69.2 


63.4 


62.4 


60.2 


59.0 


52.0 


43.9 


58.5 


50.7 


in % of standard length 






















Head length 


29.1 


29.4 


30.5 


31.0 


30.0 


33.1 


32.7 


32.3 


31.2 


29.5 


Pre-Anal length 


54.6 


51.9 


55.5 


57.0 


55.7 


54.2 


55.8 


57.6 


55.8 


57.9 


Body depth 


24.3 


23.0 


25.5 


26.3 


23.1 


22.6 


24.5 


23.1 


24.8 


23.7 


Depth of caudal peduncle 


9.4 


9.7 


10.4 


9.6 


8.2 


9.2 


8.9 


8.8 


9.7 


11.4 


Length of caudal peduncle 


13.0 


10.7 


11.7 


12.0 


10.7 


13.4 


9.3 


11.8 


11.0 


9.7 


in % of head length 






















Snout length 


33.9 


33.0 


31.4 


34.2 


31.5 


31.6 


32.9 


31.0 


28.5 


32.0 


Eye diameter 


22.6 


21.6 


20.1 


24.7 


23.5 


18.4 


25.7 


22.5 


23.2 


24.6 


Eye-rictus distance 


22.6 


22.7 


22.6 


22.8 


23.5 


22.8 


20.7 


22.5 


21.9 


25.4 


Interorbital width 


18.1 


17.0 


15.1 


13.3 


16.8 


13.3 


13.6 


14.0 


15.9 


23.0 



Anal fin 11-111,17-19 (11,18), modally 11,17, three spines in a single specimen. 
Posterior membranous attachment on ventral edge of caudal peduncle anterior to cau- 
dal-fin base, male with a large "ear or onion-like glandular formation" (sensu Bath, 
1977: 208) on anterior face of each anal spine; "leaflet- shape" glands at tip of anal rays 
in specimens over 50-55 mm SL. 

Pectoral-fin rays 13-14 (13), modally 14. Pelvic-fin segmented rays 3. Caudal- 
fin segmented rays 13, 9 branched. 

Cirri. Orbital cirrus much shorter than eye diameter, simple (out of 40 speci- 
mens examined for this character, one 30 mm SL has a short side branch and one 36 
mm SL has two short side branches); cirrus missing in specimens less than 30 mm SL. 
Anterior nostril with a short point along posterior margin, never developed as a cirrus. 

Lateral line. Continuous canal anterodorsally with 5-9 simple pores, extending 
posteriorly to a point between vertical from bases of 3rd to 7th spine, then continuing 
posteriorly as a series of 1-3 short, disjunct, horizontally bi-pored tubes in skin; poste - 
riormost tube in area between verticals from bases of 4th to 9th dorsal-fin spines. 



SALARIA ECONOMIDISI 



125 



Fig. 3 
Salaria economidisi; Greece: Lake 
Trichords. Head pattern; a, paratype, 
female, 48.4 mm SL, CMK 16968; b, 
male, holotype, 60.8 mm SL, MHNG 
2641.89; e, male, paratype, 50.9 mm 
SL, CMK 16968; d, male, paratype, 
55.9 mm SL, CMK 16968. 




126 M. KOTTELAT 

Cephalic pores. Supratemporal canal with 3 pores. Preoperculo-mandibular 
canal with 11 pores. Orbital canal with 8 infraorbital pores (9 in two specimens) and 5 
supraorbital pores. 

Upper jaw with 25-30 teeth, posterior 1-2 (modally 1) on each side caniniform. 
Lower jaw with 20-27 teeth, posterior 1 (rarely 2) on each side caniniform. 

Both sexes show indication of fleshy crest development on head at 25 mm SL. 
Only a few males have a well developed crest; in most it remains very low. Highest 
development (about 2 mm) in 49.7 mm SL specimen. In females, only as sharp ridge. 

Coloration (Figs 1-2). Background pale yellowish brown. A mid-lateral row of 
7-8 dark brown blotches, more or less rectangular, horizontally elongated, irregularly 
shaped, alternating with 5-6 dark brown saddles along dorsal body contour and on base 
of dorsal fin. In some individuals, an intermediate row of horizontally elongated, 
irregular, dark brown spots or blotches; in a few individuals, blotches of three rows 
fused to form 3 stripes (Fig. lc). Numerous small, black spots on side, variably dis- 
tributed over whole body or only anteriorly or dorsally, almost missing in a few indi- 
viduals. A single individual plain pale yellowish brown, except for black dots (Fig. Id). 
Striped pattern present in both sexes, already distinct in specimens 28 mm SL. 

In individuals larger than about 30 mm SL, head yellowish brown, darker on 
top, with numerous small, black spots, some surrounded by a paler area (Fig. 4a). 2-3 
parallel rows of 2-6 black spots from eye to posterior corner of crest, sometimes fused 
to form short lines; 2 rows sometimes marking edges of a darker band. Usually 4 (3-5) 
oblique rows of 4-8 black spots on cheek, slanted posteroventrally, 3rd (counted from 
dorsalmost) originating at lower margin of eye; area above upper row dark brown, 
between rows 1 and 3 pale yellowish brown, between rows 3-4 dark brown, and below 
row 4 pale yellowish brown. Sometimes a short additional row starting at posterior 
extremity of upper lip. Throat uniformly yellowish brown. 

In specimens less than about 30 mm SL, bands on top of head more contrasty 
and a faint pale stripe from eye to tip of snout. Dark band on cheek distinct, but rows 
of spots absent. A darker patch at tip of lower lip, and a diffuse darker triangular patch 
on each side of throat. 

Dorsal fin with dark blotches or band along base; remainder of fin, in males 
more or less plain greyish brown, in females with 2-3 rows of dark brown spots on rays 
of segmented-ray portion. A black botch between anterior 2 spines, conspicuous in 
females and juveniles, faint to indistinct in males. Anal fin plain greyish brown, with a 
white distal and a dark brown subdistal bands. Caudal fin with a dark brown blotch 
continuing mid-lateral row on flank; 3-4 vertical rows of dark brown spots on rays, less 
contrasty in adult males. Pectoral-fin rays with several dark brown patches (tessellated) 
and a few black spots equivalent to those on head and flank. Pelvic hyaline. 

Distribution. Definitively known only from Trichords Lake (Fig. 5). The species 
possibly occurs in some or all of the other lakes of the Acheloos drainage (Lysimachia, 
Ozeros, Amvrakia), but this remains to be confirmed. 

Habitat and biology. Lake Trichords is 18.1 x 7.5 km large, up to 57 m deep, 
oligotrophic and drains to Acheloos River through lake Lysimachia. It has a late 
Pliocene origin (Economidis & Miller, 1990: 147, 166). The only other fish species 
endemic to Lake Trichonis is the miniature gobiid Economidichthys trichonis Econo- 



SALARIA ECONOMIDISI 



ìli 




Head pattern and cirrus of: a, Salaria economidisi, male, CMK 16968, 49.7 mm SL; b, S. flu- 
viatilis, female, CMK 17000, 63.4 mm SL. 

midis & Miller, 1990, but several other species are endemic to either the Acheloos 
drainage, or the lakes of the lower Acheloos drainage. 

On 6-7 October 2001, I observed and collected the species in Lake Trichonis 
east of Panetolio. The species was not seen in a small stream entering the lake, neither 
on sandy-muddy nor on gravely to stony substrate. It was observed only in the lake it- 
self, at depths of at least 0.4 m. The area was a shallow strand, mostly sandy, but with 
large patches of underwater vegetation (mostly clumps of Sagittaria (?) sp.). Salaria 
economidisi was observed only in these clumps; it was not seen over the sand or on or 
near stones even with the use of an electric fish-shocker. Other fish species collected in 
these clumps were the gobies Knipowitschia cf. caucasica (Berg, 1916), Economi- 
dichthys pygmaeus (Holly, 1929) and occasionally the loach Cobitis trichonica 
Stephanidis, 1974. However, P. S. Economidis informs me that he has observed the 
species on stony and gravel bottoms in other parts of the lake, especially at Pantanassa, 
about 10 km east of Panetolio. 

The reproduction and larvae are described in Economou et al. (1994). Eggs are 
laid in May- July. Early larvae are planktonic and settle at 14-15 mm. In October 2001, 
I observed large numbers of individuals 15-30 mm SL, probably young of the year. The 
largest collected male is 60.8 mm SL and the largest female 48.4. 

Etymology. Named for Panos Stavros Economidis, in appreciation for his help 
during several field trips in Greece, stimulating discussions, encouragement, and being 
a unique historical, gastronomic and oenological cicerone. 

DISCUSSION 

The blenny from Lake Trichonis has long been identified as S. fluviatilis, a 
species recorded from freshwaters all along the northern shore of the Mediterranean, 
from Israel to Portugal (Guadiana drainage) and Algeria and Morocco, including 
Sardinia, Corsica and Creta (Mouslih, 1984; Doadrio, 2001; Steinitz, 1950; Goren, 
1974). When I first encountered it in the field, I immediately suspected S. economidisi 
was distinct from S. fluviatilis because of habitat, behaviour, general coloration and 
patterning on the head. 



128 M. KOTTELAT 

I have observed S. fluviatilis in riverine habitats in Greece and France, and it 
was always observed among stones, usually preferring the deepest parts with fast cur- 
rents. This is also the habitat reported in the literature (e.g., Freeman et al., 1990; 
Hernandez et al., 2000). In lacustrine habitat, I observed it in Lake Maggiore, where it 
was found under stones, in at least 0.5 m depth. This is also the habitat described by 
Elmiger (2002). I have never observed the species over sandy bottom under vegetation, 
the habitat in which S. economidisi was collected in Lake Trichonis. The largest ex- 
amined individual of S. economidisi is 60.8 mm SL, while the largest S. fluviatilis is 
111.8 mm SL, and larger sizes are reported in the literature (up to 150 mm; Persat, in 
Keith & Allardi, 2001). 

The colour pattern of S. fluviatilis varies, depending on sex, size, sexual activi- 
ty and 'mood' of the individual fish (e.g., Wickler, 1957), but there is no published in- 
formation on geographic variation. I have seen too few specimens (or more precisely, 
most samples I have seen include too few specimens) to venture into a detailed inter- 
population comparison, and such a comparison would probably better be based on live 
specimens observed over an extended period of time, in order to account for seasonal 
variation. It is a sad reality that research on European fishes is made very difficult if 
not impossible for reasons which have nothing to do with sciences (Kottelat, 1997) and 
that access to fresh material badly needed for taxonomic revisions often is simply im- 
possible to competent taxonomists. But the following generalities can be made. 

I have not seen geographic variation in the colour pattern of S. fluviatilis. The 
following common features are shared by the examined populations. In juveniles, fe- 
males and small males, the body has a mid-lateral series of about 7-10 irregularly 
squarish dark blotches, more or less alternating or connected with similar blotches 
along the dorsum (e.g., Fig. 6b, f). Small black dots may be present on the body, usu- 
ally on the dorsal half. In larger females (over 60 mm SL), the blotches tend to become 
more vertically elongated, more irregular, often dissociated to form a marmorated pat- 
tern. In larger males, the blotches become less contrasty, and the body might vary from 
marmorated to plain dark brown to black or plain olive brown (e.g., Fig. 6c, e). In S. 
economidisi, 7-8 squarish blotches are present in most adults. In some individuals, they 
may fade out or fuse to form midlateral stripes. I have not observed plain dark brown 
males. The general background colour is yellowish brown in all individuals, paler than 
in any S. fluviatilis. 

Salaria fluviatilis exhibits a conspicuous and distinctive colour pattern on the 
head (Figs. 4b, 7), which typically includes: a) a dark brown band from eye to posterior 
extremity of the crest, margined anteriorly and posteriorly by pale grey to yellowish 
bands; b) a pale band from eye to tip of snout, margined posteriorly by a dark band; c) 
from the upper extremity of this last band (under the eye), a conspicuous dark band 
extends posteroventrally, margined above and below by pale bands, and below by a 
narrow dark line along the edge of the posterior part of the upper lip; d) ventral side of 
head with a blackish blotch at tip of lower jaw, an elongated blotch on each side of jaw, 
and an oblique band connected or not with band (c). These bands are usually filled with 
tiny black dots and are never margined by rows of larger black dots or black lines. 

This pattern is variously developed in all examined specimens. It is often partly 
or wholly obscured, especially in large dark males. I have seen it in material from 



SALARIA ECONOMIDISI 



129 




Fig. 5 
Greece and western Balban Peninsula; arrow indicates Lake Trichonis, type locality of Salaria 
economidisi. 



Spain (Ebro drainage), France (Tech, Rhône and Var) and Switzerland (Lake 
Maggiore). It is figured in material from France (Keith & Allardi, 2001), Sicily (Zava 
& Violani, 1991), Garda Lake (Wickler, 1957), Lake Maggiore (Elmiger, 2002), Spain 
(Doadrio, 2001) and Slovenia (Povz & Sket, 1990). This pattern is also obvious in pho- 
tographs of live specimens from Turkey (from Kirandere near Iznik Lake [40°25'N 
29°43'E], and from Aksu Köprüsu) provided by R. Wildekamp. From Greece, I have 
examined only 5 specimens from Pinios drainage (Péloponnèse), which includes a 
large 111.8 mm SL black male in which no cephalic pattern can be distinguished any- 



130 M. KOTTELAT 

more; in the four smaller specimens, only a faint indication of the dark cheek bar can 
be recognised. It is also present in material from Lake Kinneret (Israel) which is dis- 
cussed below. 

In S. economidisi over about 30 mm SL, elements (b) and (d) are missing. The 
throat is finely dotted by very regularly set melanophores. Elements (a) are very poorly 
contrasted; instead, there are 2-3 rows of black dots in positions corresponding to the 
limits between the pale and dark bands. Elements (c) are replaced by 3-5 (usually 4) 
rows of conspicuous black dots, with the space between the lower two rows usually 
slightly darker than between the other rows. In specimens less than about 30 mm SL, 
elements (b) are usually distinct, and the jaw markings of (d) are distinct; there are no 
rows of black spots in elements (a) and (c) which have an appearance similar to that of 
S.fluviatilis (suggesting that the S.fluviatilis pattern is probably the plesiomorphic con- 
dition). 

Besides colour pattern, S. economidisi is distinguished from S. fluviatilis in 
having a longer head (29.1-33.1 % SLin specimens 39.9-60.8 mm SL, vs. 25.3-30.6 in 
specimens 37.8-111.8; Fig. 8), more and smaller teeth (25-30 in the upper jaw, vs. 16- 
24; 20-27 in the lower jaw, vs. 16-20; I have not observed variation associated with size 
or sex). The orbital cirrus is simple, while it is branched in most examined specimens 
of S. fluviatilis larger than 38 mm SL. The lateral line is much less developed, with only 
5-9 pores (vs. usually 12-22, rarely 6-8, see below) in the continuous anterior canal, 
followed by 1-3 bi-pored tubes (vs. 4-12), the last one located between verticals 
through base of 4th to 9th dorsal-fin spine (vs. between vertical through base of 9th 
spinous to 13th segmented ray), on the anterior arched portion of the lateral line (vs. 
on the posterior horizontal portion). 

The larvae of S. economidisi are described by Economou et al. (1994) who 
compared their observations with those of Cipria (1936) on the larvae of S. fluviatilis 
from northern Italian lakes. They report that S. economidisi differs from S. fluviatilis in 
the absence (vs. presence) of a large melanophore between the end of the ventral row 
of melanophores and the end of the notochord. 

The synonymy of S. fluviatilis includes 12 nominal species (Kottelat, 1997) and 
the question arises whether some of these synonyms could be S. economidisi or could 
be valid species masquerading under the name S. fluviatilis. 

Blennius fluviatilis Asso, 1801 was described from the river Ebro in Zaragoza 
(Spain) and B. f rater Bloch, in Schneider, 1801 was based on an earlier description of 
Asso's Zaragoza material and thus represents the same species. Blennius inaequalis 
Valenciennes, in Cuvier & Valenciennes, 1836 was described on the basis of material 
from Séte (southern France), B. cagnota Valenciennes, in Cuvier & Valenciennes, 1836 
on the basis of material from Toulon (southeastern France; type locality restricted by 
Kottelat, 1997, by lectotype designation) and S. varus Risso, 1827 on the basis of 
material from river Var (southeasten France). I have examined material from Ebro and 
Var drainages as well as from Tech drainage (in intermediate geographic position); I 
have not seen differences among these populations and conclude that they represent a 
single species. 

I have not seen material from the Tyrrhenian coast of Italy, Sicilia, Sardinia and 
Corsica. Blennius fluviatilis Rafinesque Schmaltz, 1810 (an homonym of B. fluviatilis 



SALARIA ECONOMIDISI 



131 



Fig. 6 
Salaria fluviatilis; a- 
b, male 83.0 mm SL 
and female 71.2 mm 
SL, CMK 17568; 
France: River Tech; 
c-d, male 111.8 mm 
SL and female 53.1 
mm SL, CMK 17305; 
Greece: Péloponnèse: 
River Pinios; e-f, 
male 74.4 mm SL and 
female 37.8 mm SL, 
CMK 16844; France: 
stream Leysse. 





-.„.... 



132 M. KOTTELAT 

Asso, 1801) was described from Sicilia, B. lupulus Bonaparte, 1840 from Toscana and 
Romana, and B. anticolus Bonaparte, 1840 from the surroundings of Roma. There does 
not seem to be differences between the fishes on the published photographs of popula- 
tions from the Tyrrhenian basin (Zava & Violani, 1991) and the material I examined 
from Spain and southern France. 

Blennius peneri Heckel & Kner, 1858 is a nomen nudum based on material 
from Split, Croatia. Besides the account and photograph in Povz & Sket (1990) there 
is no published information specifically dealing with the populations of streams 
draining to the Adriatic basin. Little can be said on the sole basis of a photograph. 

The situation with populations of subalpine lakes is different. South of the Alps, 
S. fluviatilis has long been known from Lake Garda, from where it was described as B. 
vulgaris Pollini, 1816 (Ichthyocoris pollimi Bonaparte, 1846 is an unnecessary re- 
placement name); the species is also known at least from Lakes Maggiore and Lugano. 
North of the Alps, the species is known from Lake Bourget (from where it was de- 
scribed as B. alpestris Blanchard, 1866) and the Rhône drainage. As discussed below, 
there are slight morphological differences between the Bourget population and those 
from coastal drainages. Unfortunately, I have not been able to examine material from 
the Rhône mainriver and my sample from Lake Bourget is quite small. 

I have not seen material from Lake Garda, but have examined material from 
Lake Maggiore. Here again, there are slight morphological differences between the 
Lake Maggiore population and those from coastal drainages of southern France and 
Spain. It need to be clarified whether these differences are shared with populations 
from other Insubrian lakes and those from peri-Adriatic coastal drainages. Interesting 
is that the morphological traits that distinguish the Lake Maggiore and Lake Bourget 
populations from the riverine ones are identical. 

While all populations referred to S. fluviatilis apparently have the same head 
colour pattern, there is some variability in shape and development of the lateral line. 
The material from Lake Maggiore and Leysse have a longer head (27.7-30.6 and 28.6- 
29.6 % SL, respectively; Fig. 8) than those from the Ebro, Tech, Var and Pinios (25.3- 
28.6, with a single specimen 30.1) and the Leysse one has a deeper body (22.0-26.5 % 
SL, vs. 21.3-23.8 in Pinios, 21.4-24.4 in Tech, Var and Ebro, and 20.9-24.0 in Lake 
Maggiore). The sample sizes from the various localities are, however, too small to 
reach reliable conclusions on the value of these morphometric differences. The Lake 
Maggiore specimens have an unbranched orbital cirrus. The Leysse males have a much 
higher crest (Fig. 6e), even in a 37.8 mm SL male. Both Leysse and Lake Maggiore 
populations have about 1/3 of the specimens with 14 spinous dorsal rays, while speci- 
mens from all other samples have 13 (except one from Tech). In both Leysse (L) and 
Lake Maggiore (M), the lateral line is shorter, the anterior canal has 8-16 (L) and 6-8 
(M) pores and extends posteriorly below base of 9-1 lth (L) and 3-6th (M) dorsal spine, 
continued by a series of 4-12 (L) and 5-8 (M) bi-pored tubes, extending to below base 
of 13th spine to 6th segmented (L) and 9- 13th spine (M). Noteworthy is that the Lake 
Maggiore population is purely lacustrine and the Leysse population is from a tributary 
of Lake Bourget (the swiftest habitat where I collected blennies) about 2 km from the 
lake; these populations are possibly partly isolated from the riverine populations and 
occur in much colder habitats, at higher elevations. 



SALARIA ECONOMIDISI 



133 



Fig. 7 
Salario fluviatilis; head pattern; a-b, 
male 83.0 mm SL and female 76.6 mm 
SL, CMK 17568; France: River Tech; 
e, female 63.4 mm SL, CMK 17000; 
Spain: Ebro; d, female 53.1 mm SL, 
CMK 17305; Greece: River Pinios. 





134 



M. KOTTELAT 



35-1 



— ^ 30H 

E 
E 

S 25 - 

e 

0) 



tv 



20- 



15- 



10- 



• S. economidisi y=0.24x + 3.24, R - 0.91 

S. fluviatilis 
o Tech, Var, Pinios y=0.30x - 1 .47, R 2 = 0.98 

X Leysse T_ y =o.3-| x _ fj.98, R^ 0.99 

+ Lago Maggiore J 




40 



60 80 100 

standard length (mm) 



120 



Fig. 8 
Relationships between standard length and head length in Solaria economidisi and S. fluviatilis. 



A population of Salaria has been known from Lake Kinneret (= lake Galilee or 
Tiberias) in Israel since Günther (1864; see also Lortet, 1883; Steinitz, 1950; Krupp & 
Schneider, 1989). Lake Kinneret is part of the endorheic Jordan drainage and the local 
population must have been isolated from the peri-Mediterranean ones since lower 
Pleistocene (Krupp & Schneider, 1989: 350), which creates a suspicion that it could be 
specifically distinct. Externally the Kinneret population is only minimally distin- 
guished from the other populations I have examined. It has XIII-XIV dorsal spines 
(modally XIV, vs. XII-XIII, modally XIII) and usually a poorly contrasted cephalic 
colour pattern. But in individuals in which the pattern is contrasted, it is identical to 
what is observed in peri-Mediterranean coastal streams. The only difference possibly 
is the presence (vs. absence) of a blackish spot in juveniles at about 1/3 of length of 
pectoral fin and grayish lower fourth of pectoral. 

A tentative conclusion is that the coastal populations are apparently all conspe- 
cific, while some of the lacustrine ones might represent distinct species. 

Salaria economidisi is definititely recorded only from Lake Trichonis; it 
possibly also occurs in the other lakes of the lower Acheloos drainage. The presence of 
S. fluviatilis in riverine habitats of the Acheloos drainage should be expected; it might 
even be present in the lakes along rocky shores. 

Perdices et al. (2000) investigated allozyme systems in 5 populations of S. flu- 
viatilis from Spain and 3 from Greece. This limited sampling and the absence of geo- 



SALARIA ECONOMIDISI 135 

graphic spectrum significantly reduces the significance of the conclusions derived from 
the study. But of interest here is that the authors probably included S. economidisi in 
their analysis (their Trichonis sample). All their phenograms show S. economidisi as 
the sister group to a cluster including all other Greek and Spanish populations, a pattern 
congruent with my conclusion that it is a distinct species. 

Further, the two other Greek populations (Miras drainage and Doiran Lake) 
cluster together as sister group to all but one (Ruidera Lakes) Spanish populations. The 
Ruidera population comes out as sister group to all other S. fluviatilis populations. 
Perdices et al. do not mention any non-allozymic character. The branching sequence 
among their populations of S. fluviatilis is largely what is expected considering the vast 
geographic distance between the Spanish and Greek populations and the absence of 
material from intermediate areas. Inclusion of material from Mediterranean islands, 
various lacustrine populations, Turkey and North Africa would be a valuable addition 
to their study. 

Nevertheless, the puzzling position of the Ruidera population would justify that 
its identity be critically re-examined. Ruidera Lake is a system of small lakes in the 
upper Guadiana drainage, one of the few Atlantic drainages where S. fluviatilis is 
recorded. Speculations on the evolution of the European freshwater blennies is prema- 
ture before their diversity, variation and distribution are documented and a robust 
phylogeny has been hypothesised. 

COMPARISON MATERIAL 

Salaria fluviatilis: CMK 17305, 5, 44.7-111.8 mm SL; Greece: Peloponnisos: River Pinios at 
bridge 3 km west of Simopoulo, on road to Efira; 37°5048"N 21°32'07"E; M. Kottelat, 
M. Stoumboudi & R. Barbieri, 14 May 2002. - CMK 5432, 9, 18.2-60.5 mm SL; 
Switzerland: Ticino: Lago Maggiore at Locarno; 46°09'30"N 8°48'20"E; M. Kottelat, 25 
Aug 1985. - CMK 17648, 11, 58.8-85.0 mm SL; France: Alpes-Maritimes: River Var at 
Pont Napoléon III in Nice, about 1 km from sea; J.-M. Foissy & CSP team; 3 October 
2002. - CMK 16844, 6, 37.8-74.4 mm SL; France: Haute-Savoie: stream Leysse (tribu- 
tary of Lake Bourget) at bridge at Le Tremblay, west of Chambéry; M. Kottelat & H. 
Persat, 20 July 2001. - CMK 17568, 11, 38.6-83.0 mm SL; France: Pyrénées-Orientales: 
River Tech about 3 km upriver of bridge on road N114, south of Eine; 42°35'02"N 
2°58'13"E; M. Kottelat & CSP team, 6 September 2002. - CMK 17000, 2, 53.2-63.4 mm 
SL; Spain: Ebro drainage; B. Elvira, October 2001. 

Salaria pavo: CMK 16978, 2, 60.7-81.2 mm SL; Greece: Kleisova Lagoon at Mesologgi, east of 
port, at beginning of road to Turlida; 38°21'39.5"N 21°25'38.9"E; M. Kottelat & P. S. 
Economidis, 8 October 2001. 

Salaria sp.: CMK 17016, 35, 21.0-70.9 mm SL; Israel: Lake Kinneret; M. Goren, 13 May 1993. 
- CMK 17017, 25, 20.4-70.8 mm SL; same locality; M. Goren, 10 June 1993. 

ACKNOWLEDGEMENTS 

I am pleased to thank Panos Economidis, Maria Stoumboudi, Roberta Barbieri, 
Dominique Beaudou, Jean-Michel Foissy, Philippe Théate and Henri Persat for assis- 
tance in obtaining freshwater blennies, Menachem Goren and Benigno Elvira for the 
gift of material, Uli Schliewen for making available figure 2, Ruud Wildekamp for pro- 
viding photographs of live individuals, Victor G. Springer and Panos Economidis for 
commenting on the manuscript. 



136 M. KOTTELAT 



REFERENCES 



Bath, H. 1977. Revision der Blenniini (Pisces: Blenniidae). Senckenbergiana Biologica 57 

(1976 [1977]): 167-234. 
Bath, H. 1996. Beitrag zur Osteologie der Arten der Tribus Parablenniini. Die Beziehungen der 

Knochen des Schädeldaches zum Seitenorgan-System und zu den Weichteilbildungen 

der Kopfoberseite sowie die systematische Bedeutung der Befunde nebst Bemerkungen 

zu Lupinoblennius dispar Herre 1942 (Pisces: Blenniidae). Senckenbergiana Biologica 

76: 65-92. 
Bock, M. & Zander, C. D. 1986. Osteological characters as tool for blenniid taxonomy - a 

generic revision of European Blenniidae (Percomorphi; Pisces). Zeitschrift für 

Zoologische Systematik und Evolutionsforschimg 24: 138-143. 
Cipria, G. 1936. Uova, stadi embrionali e post-embrionali nei Blennidi. I. Blennius pavo Risso. 

IL Blennius inaequalis C. V. Memoria del R. Comitato Talassografico Italiano 231: 1-7, 

1 Pi- 

Doadrio, I. 2001. Atlas y libro rojo de los peces continentales de Espana. Direccion General de 

Conservación de la Naturaleza & Museo Nacional de Ciencias Naturales, Madrid, 

364 pp. 
Economidis, P S. & Miller, P. J. 1990. Systematics of freshwater gobies from Greece 

(Teleostei: Gobiidae). Journal of Zoology 221: 125-170. 
Economou, A. N., Daoulas, C, Psarras, T & Barbieri-Tseliki, R. 1994. Freshwater larval 

fish from Lake Trichonis (Greece). Journal of Fish Biology 45: 17-35. 
Elmiger, C. 2002. Benthische Kleinfischarten des Tessins: Verbreitung und Habitatsnutzung des 

Ghiozzo (Padogobius bottelli) und der Cagnetta (Salaria fluviatilis). Diplomarbeit, ETH 

Zürich, 71 pp. 
Freeman, M. C, Vinolas, D, Grossman, G. D. & de Sostoa, A. 1990. Microhabitat use by 

Blennius fluviatilis in the Rio Matarrana, Spain. Freshwater Biology 24: 335-345. 
Goren, M. 1974. The freshwater fishes of Israel. Israel Journal of Zoology 23: 67-118. 
Günther, A. 1864. Catalogue of the fishes in the British Museum. 5. British Museum, London, 

xxii+455 pp. 
Hernandez, R., Lacomba, R. T., Uvinas, Y. N. & Oltra, R. 2000. Distribution pattern of river 

blennies in the Jucar River basin (eastern Spain). Journal of Fish Biology 57: 250-254. 
Hubbs, C. L. & Lagler, K. F. 1947. Fishes of the Great Lakes region. Cranbrook Institute of 

Sciences, Bulletin, 26: 1-213. 
Keith, P. & Allardi, J. 2001. Atlas des poissons d'eau douce de France. Patrimoines Naturels 

47: 1-387. 
KOTTELAT, M. 1997. European freshwater fishes. An heuristic checklist of the freshwater fishes 

of Europe (exclusive of former USSR), with an introduction for non-systematists and 

comments on nomenclature and conservation. Biologia, Bratislava, Section Zoology 52 

(Suppl. 5): 1-271. 
Krupp, F. & Schneider, W. 1989. The fishes of the Jordan River drainage basin and Azraq Oasis. 

Fauna of Saudi Arabia 10: 347-416. 
Lortet, L. 1883. Poissons et reptiles du lac de Tibériade et de quelques autres parties de la Syrie. 

Archives du Muséum d'Histoire Naturelle de Lyon 3: 99-194, pis. 6-18. 
Mouslih, M. 1984. Présence de la blennie cagnotte: Blennius fluviatilis Asso 1801 (Pisces, 

Blennidae) et de la loche: Cobitis taenia Linné 1758 (Pisces, Cobitidae) sur le Causse 

Moyen- Atlasique. Bulletin de l'Institut Scientifique de Rabat 8: 171. 
Perdices, A., Doadrio, L, Côté, I. M., Machordom, A., Economidis, P. S. & Reynolds, J. D. 

2000. Genetic divergence and origin of Mediterranean populations of the river blenny 

Salaria fluviatilis (Teleostei: Blenniidae). Copeia 2000: 723-731. 
Povz, M. & Sket, B. 1990. Nase sladkovodne ribe. Mladinska Knjiga, Ljubljana, 374 pp. 
Sazima, I. & Carvalho-Filho, A. 2003. Natural history of the elusive blenny Lupinoblennius 

paivai (Perciformes: Blenniidae) in coastal streams of southeast Brazil. Ichthyologica! 

Exploration of Freshwaters 14: 175-184. 



SALARIA ECONOMIDISI 137 



Springer, V. G. & Gomon, M. F. 1975. Revision of the blenniid fish genus Omobranchus with 
descriptions of three new species and notes on other species of the tribe Omobranchini. 
Smithsonian Contributions to Zoology ill: 1-135. 

Springer, V. G. & Smith-Vaniz, W. F. 1972. A new tribe (Phenablenniini) and genus 
(Phenablennius) of blenniid fishes based on Petroscirtes heyligeri Bleeker. Copeia 1972: 
64-71. 

Springer, V. G. & Williams, J. T. 1994. The Indo-West Pacific blenniid fish genus Istiblennius 
reappraised: a resision of Istiblennius, Blenniella, and Paralticus, new genus. Smith- 
sonian Contributions to Zoology 565: 1-193. 

Steinitz, H. 1950. Contribution to the knowledge of the Blenniidae of the eastern Mediterranean 
III. Istanbul Université si fen Fakultesi Mecmuasi, Seri B 15: 60-87. 

Whitehead, P. J. P., Bauchot, M.-L., Hureau, J.-C., Nielsen, J. & Tortonese, E. 1984-1986. 
Fishes of the north-eastern Atlantic and the Mediterranean. UNESCO, Paris, 3 vols. 

Wickler, W. 1957. Vergleichende Verhaltensstudien an Grundfischen. I. Beiträge zur Biologie, 
besonders zur Ethologie von Blennius fluviatilis Asso im Vergleich zu einigen anderen 
Bodenfischen. Zeitschrift für Tierpsychologie 14: 393-428. 

Zava, B. & Violani, C. 1991. Contributi alla conoscenza dell'ittiofauna delle acque interne 
siciliane. I. Sulla presenza in Sicilia di Salaria fluviatilis (Asso, 1801) (Pisces, Blenn- 
niidae). Bollettino del Museo Regionale di Scienze Naturali, Torino 9: 313-324. 



Revue suisse de Zoologie 111 (1): 139-174; mars 2004 



A taxonomic revision of the family Oncopodidae IV. The genus 
Oncopus Thorell (Opiliones, Laniatores) 

Peter J. SCHWENDINGER 1 & Jochen MARTENS 2 

1 Muséum d'histoire naturelle, case postale 6434, CH-1211 Genève 6, Switzerland. 
E-mail: peter.schwendinger@mhn.ville-ge.ch 

2 Institut für Zoologie, Johannes Gutenberg-Universität Mainz, Saarstr. 21, 
D-55099 Mainz, Germany. E-mail: martens@mail.uni-mainz.de 



A taxonomic revision of the family Oncopodidae IV. The genus Oncopus 
Thorell (Opiliones, Laniatores). - The known Oncopus species are revised 
and their penis morphology is illustrated. Distinct sexual dimorphism in 
external characters of some species is pointed out. Oncopus alticeps Pocock 
is placed in the synonymy of O. feae Thorell and O. acanthochelis Roewer 
in the synonymy of O. truncatus Thorell. Four new species are described: 
O. malayanus sp. n. from Peninsular Malaysia, O. tiomanensis sp. n. from 
Tioman Island (Malaysia), O. lingga from two Indonesian islands in the 
Lingga Archipelago and O. expatriatus sp. n. seemingly from Thailand 
(dubious record; more likely occurring on Borneo). Nine valid species are 
recognized and placed in four species groups; relationships and zoogeo- 
graphy are discussed. 

Key- words: Opiliones - Oncopodidae - Oncopus - new species - taxonomy 
- zoogeography - Southeast Asia. 

INTRODUCTION 

This is our fourth contribution in the course of a revision of the family Onco- 
podidae, completing the treatment of known species (see Martens & Schwendinger, 
1998; Schwendinger & Martens, 1999, 2002). The present paper deals with the genus 
Oncopus, in which seven species were hitherto known [not including O. cuspidatus 
Schwendinger, which was only provisionally described in this genus (see 
Schwendinger, 1992: 197) and later transferred to Caenoncopus (see Martens & 
Schwendinger, 1998: 507)] and the taxonomy of which is in considerable confusion. 
As most previous authors were seemingly unaware of external sexual dimorphism in 
this genus (present also in some species of Gnomulus), did not take into consideration 
genital characters and described most species from single specimens (several of them 
$ 9 , one even a juvenile), species distinctions were by no means clear. No Oncopus 
species was hitherto known from both sexes, and only in O. megachelis was penis mor- 
phology described [the illustration of an Oncopus penis by Martens (1986: fig. 7) was 
not assigned to any species; it refers to O. truncatus]. Two species were described 



Manuscript accepted 25.07.2003 



140 P- J- SCHWENDINGER & J. MARTENS 

and named twice: O. feae Thorell (from Penang Island) originally from a $ and later 
again from a S (described as O. alticeps) by Pocock; O. truncatus Thorell (from 
Singapore Island) originally from a juvenile and subsequently again from a ? 
(described as O. acanthochelis) by Roewer. The latter author caused further confusion 
by identifying a S , which clearly belongs to O. doriae (occurring in Sarawak, Borneo), 
as "0. alticeps" and giving Thailand as its country of origin (see Roewer, 1923: 60). 
Another â in his collection (unpublished record; here described as O. expatriatus 
sp. n.) was identified as "Pelitnus alticeps", a species that was never formally described 
(probably a misunderstanding for Oncopus alticeps) and also referred to as originating 
from Thailand, but penis morphology suggests that this species probably occurs on 
Borneo as well. 



MATERIAL AND METHODS 

External structures were studied and drawn with a ZEISS SV11 stereomi ero- 
scope, the penes with a NIKON Optiphot compound microscope (each with a drawing 
tube). In contrast to other genera, the penes of Oncopus â â examined were often 
partly expanded, i.e. the glans raised from the proximad-directed resting position by 
about 30°. Drawings were made with the glans pressed back into its resting position by 
squeezing the apex of the penis between a microscopic slide with a well (filled with 
diluted glycerine) and a cover slide. 

Body measurements refer to the dorsal scutum. Leg articles were measured on 
their dorsal side, from midpoint of anterior margin to midpoint of posterior margin. All 
measurements are given in mm. Terminology of penis morphology follows that of 
Martens & Schwendinger (1998: fig. 1). 

Abbreviations used in the text. Fe - femur, Mt - metatarsus, Pa - patella, Ta - 
tarsus, Ti - tibia, Tr - trochanter. 

CCD - collection of Christa Deeleman-Reinhold, Ossendrecht; MAR - collec- 
tion of J. Martens, Mainz; MCZ - Museum of Comparative Zoology, Harvard 
University, Cambridge, Massachusetts; MHNG - Muséum d'histoire naturelle, 
Genève; MSNG - Museo Civico di Storia Naturale, Genova; NHML - Natural History 
Museum, London [formerly British Museum (Natural History)]; SMF - Naturmuseum 
und Forschungsinstitut Senckenberg, Frankfurt; ZMT - Zoological Museum, 
University of Turku; ZRC - Zoological Reference Collection of the Raffles Museum of 
Biodiversity Research, National University of Singapore. 



TAXONOMY 

Oncopus Thorell, 1876 

Oncopus Thorell (1876: 134-135). - Thorell (1891: 99-100). - Loman (1902: 182). - Roewer 
(1923: 57). - S0rensen (1932: 210-211). - Martens & Schwendinger (1998: 552). - 
Schwendinger & Martens (2002b: 425-426). 

Type species. Oncopus doriae Thorell, 1876 by original designation and by 
monotypy. 



THE GENUS ONCOPUS 141 

Diagnosis. Large to very large oncopodids (5.9-10.8 mm), characterized by 
only one tarsal article on all legs. Interocular area a low, forward-directed tubercle, a 
low rounded hump or not elevated; carapace region larger and more or less distinctly 
elevated in â â ; carapace-abdomen bridge indistinct, wide, undivided, never in the 
shape of two opposing teeth (as in Gnomulus and Biantoncopus); ventral scutal eleva- 
tions of S â keeled, with transversal bands of white enclosures embedded in the 
cuticle; ventral side of leg coxa II usually with scale- or knob-shaped apophysis at its 
anteroproximal corner [indistinctly developed (as in Gnomulus spp.) only in O. hosei 
and O. megachelis]; stigmatic pit narrow, its posterior margin without tubercle; 
chelicerae robust (stronger in S S than in 9 9), with anteriad-inclined prodorsal boss 
exte'nding into more or less strongly developed dorsomedian boss on proximal article 
and with ventral process on proximal, median and often also on distal article (= mobile 
finger); palps with more or less pronounced ventral tubercles on coxa, trochanter, 
proximal and subdistal part of femur, proximal and distal part of tibia; some species 
also with wide proventral process on patella. Penis with one or several pairs of termi- 
nal setae on distal margin of truncus; glans proximad-directed, more or less distinctly 
narrower than truncus at that point, with U-, V- or W-shaped median plate, outwards- 
bent lateral sclerites, long membranous tubes and slender, tubular stylus with 
moderately bulbous base and with pair of subterminal teeth. 

Remarks. Possible synapomorphies for the genus Oncopus are: 1) tarsalia 
reduced to a single article on all legs; 2) ventral process present on 3 rc * cheliceral article 
(mobile finger; Figs 77-85); 3) anteroproximal apophysis on ventral side of leg coxa II 
more strongly developed than in other genera, scale-like (Fig. 71) or (rarely) knob-like 
(Figs 24-26); 4) ventral scutal elevations in S S keeled, containing white enclosures 
(Fig. 27b); 5) distal margin of truncus penis carrying terminal setae (Figs 4-7); 6) 
sexual dimorphism in shape and size of carapace region and of chelicerae (see Figs 1 1 , 
12 and Figs 13, 14). The latter is also present in some Gnomulus species (e.g., G. suma- 
tranus Thorell) and indicates (together with its proximad-directed, symmetrical glans 
penis) a close relationship between Oncopus and Gnomulus (see Schwendinger & 
Martens, 2002b). 

Species account and distribution. Nine valid species are here recognized: O. 
expatriatus sp. n. allegedly from Thailand (dubious record; more likely occurring on 
Borneo); O.feae Thorell from Penang Island (W-Malaysia); O. malayanus sp. n. from 
central Peninsular Malaysia and from Pangkor Island off the west coast of the penin- 
sula; O. tiomanensis sp. n. from Tioman Island off the east coast of Peninsular 
Malaysia; O. truncatus Thorell from Singapore Island and from the southernmost part 
of Peninsular Malaysia; Oncopus lingga sp. n. from two islands in the Lingga Archi- 
pelago (Indonesia); Oncopus doriae Thorell from western Sarawak; Oncopus hosei 
Pocock from eastern Sarawak; Oncopus megachelis Schwendinger from Sabah. Two 
presumably undescribed species occur in northern Sumatra and in southeastern Kali- 
mantan, respectively (see "Unidentified material") (Fig. 1). 

The f£A£-group 

Diagnosis. Large species (5.9-9.7 mm body length) with more or less distinctly 
forward-inclined interocular area; paramedian processes on opisthosomal area VIII 



142 



P. J. SCHWENDINGER & J. MARTENS 




Fig. 1 
Known records of Oncopus species. - 1 Bangkok (O. expatriants sp. n.; dubious record), 2 
Penang Island (O. feae), 3 Padang Gerus (O. malayanus sp. n.), 4 Maxwell Hill (O. malayanus 
sp. n.), 5 Cameron Highlands (O. malayanus sp. n.), 6 Chenderiang (O. malayanus sp. n.), 7 
Pangkor Island (O. malayanus sp. n.), 8 Templer Park, Ulu Gombak, Kuala Lumpur (O. 
malayanus sp. n.), 9 Pasoh Forest (O. malayanus sp. n.?), 10 Bukit Charas (Oncopus sp.), 11 
Tioman Island (O. tiomanensis sp. n.), 12 Gunung Pulai (O. truncatus), 13 Singapore Island (O. 
truncatus), 14 Lingga Island (O. lingga sp. n.), 15 Singkep Island (O. lingga sp. n.), 16 forest 
near Prapat (Oncopus sp.), 17 Batu Lundu, Gunung Gading (O. doriae), 18 Gunung Santubong 
(O. doriae), 19 Bako National Park (O. doriae), 20 Gunung Serapi (O. doriae), 21 Balai Ringin 
(O. doriae), 22 Gunung Penrissen (O. doriae), 23 Kapit (Oncopus sp.), 24 Lambir Hills National 
Park (O. hosei), 25 Kuala Baram (O. hosei), 26 Kabili-Sepilok Forest Reserve (O. megachelis), 
27 Sangasanga Muara (Oncopus sp.), 28 Sepaku (Oncopus sp.). 



projecting beyond posterior margin of dorsal scuta; penes slender, more or less gradu- 
ally widening towards apex with almost straight distal margins carrying three to five 
terminal setae on each side. 

This species group is close to the truncatus-group and contains two species (O. 
feae and O. malayanus sp. n.) from the northern and central parts of Peninsular 
Malaysia. 



THE GENUS ONCOPUS 143 

Oncopus feae Thorell, 1 890 Figs 2-26 

Oncopusfeae Thorell (1890: 375-377); description of $ . - Roewer (1923: 58). - S0rensen (1932: 
211-213, figs 1-2). - Bristowe (1976: 7-11, figs 1-6, pi. 1-2). - Moritz (1993: fig. 299A); 
drawing. - Murphy & Murphy (2000: pi. 2.1); photo. 

Oncopus alticeps Pocock (1897: 287, fig. 2), syn. n.; description of S . - Roewer (1923: 59-60, 
fig. 62); see also treatment of O. doriae in present paper. 

Types. MALAYSIA (peninsula), Pulo Pinang (= Pulau Penang = Penang Island), 12,4 
juvenile syntypes (MSNG, examined), leg. L. Loria and L. Fea, 1889; 9 here designated lecto- 
type in order to preserve stability of nomenclature. 

Other material. From Penang Island: 1 S (holotype of O. alticeps), 1897.1.25.1 
(NHML); Batu Feringgi, 20 m, 2 juveniles, leg. G. Cuccodoro & I. Lobi, 10.XI.1999; Batu 
Feringgi, 100 m, 1 o\ 1 9, leg. G. Cuccodoro & I. Lobi, 13.XI.1999; forest above Botanical 
Garden, 250 m, 3 9 , leg. G. Cuccodoro & I. Lobi, 12.XI.1999; Penang Hill (= Bukit Benderà), 
650 m, 1 ó\ 5 9, 2 juveniles, leg. G. Cuccodoro & I. Lobi, 19.XI.1999; Penang Hill, 650-760 
m, 1 o\ leg. P.J. Schwendinger, 19.1.1995; Penang Hill, 710 m, 1 S , leg. P.J. Schwendinger, 
8./9.XII.1997; all latter specimens deposited in MHNG. 

Diagnosis. Distinguished by: interocular area more or less distinctly forward- 
inclined; pair of paramedian tubercles distinctly projecting beyond posterior margin of 
dorsal scutum; anteroproximal process on ventral side of coxa II developed as longitu- 
dinal bulge; second cheliceral article with ventroproximal hump; mobile cheliceral 
finger with ventroproximal boss; ventroproximal process on palpal tibia distinct; penis 
with three to five terminal setae on each side, lateral sclerites of glans distally wide, 
truncate. 

Redescription. Large species (8.7-9.7 mm long). Carapace region of dorsal 
scutum with more or less distinctly forward-inclined interocular area, low in 2 9 (Figs 
11, 18), arched and distinctly elevated above opisthosomal region in 6 6 (Figs 13, 
15-17); paramedian pairs of tubercles present on opisthosomal areas VI- VIII, the last 
one distinctly protruding beyond posterior margin of dorsal scutum (Figs 12, 14). 
Anteroproximal apophysis of coxa II knob-shaped in â â (Figs 24-26), scale-like in 
2 2 (Fig. 23); process posterior to apophysis of coxa II developed as longitudinal 
bulge, anteroproximal process on coxa III small (Figs 23-26). 

Chelicerae robust, stronger in â S (Figs 14, 20) than in 2 2 (Figs 12, 19), with 
pronounced ventral tubercle on first article; ventral side of second cheliceral article 
with distal process and proximal hump (indicated by arrow in Figs 19, 20); mobile 
finger of chelicera with distinct ventroproximal boss. 

Pedipalps (Figs 21, 22) with distinct ventral process on trochanter, small 
ventroproximal process on femur and distinct ventroproximal process on tibia; no 
proventral process on palpal patella. 

Legs: femur of leg II with small dorsoproximal tubercle, femur IV with indis- 
tinct ventroproximal tubercle. 

Penis (Figs 2-10) fairly slender, with slightly rounded distal margin carrying 
three to five terminal setae on each side; glans about as wide as truncus at that point 
(wider when lateral sclerites are pressed sideward during examination on a temporary 
slide), flanked by two setae on each side; lateral sclerites slender at base, distally 
widening to truncate tips with more or less distinctly serrate terminal edge, bent 
outwards and away from truncus; median plate broadly V-shaped, distally rounded; 
membranous tubes long, protruding far beyond median plate. 



144 



P. J. SCHWENDINGER & J. MARTENS 




Figs 2-10 
Oncopus feae Thorell: 3 holotype of O. alticeps (here placed in the synonymy of O. feae) (6, 
7); 3 6 from Penang Hill (2-5, 8, 9); 6 from Batu Feringgi, Penang (10). - Penis, dorsal (2) and 
lateral view (3); apex of penis, dorsal (6, 4) and lateral view (5, 7); glans penis, dorsal view (8- 
10). - Scale lines 1.0 mm (2, 3), 0.1 mm (4-10). 



Variation. Measurements (c?, in parentheses 9): body 8.67-9.67 (8.77-9.45) 
long, 4.80-5.98 (4.96-5.39) wide; carapace region 2.48-3.74 (2.08-2.39) long, 3.22- 
4.11 (2.91-3.16) wide; cT (n= 5), $ (n = 10). Variation in the shape of the interocular 
areas see Figs 11, 13, 15-18, of glans penis see Figs 4, 6, 8-10. In some â 6 the knob- 
shaped anteroproximal apophysis on coxa II is divided into two parts (Figs 24-26). 

Remark. In 2 ô the penis is subdistally collapsed and its distal margin therefore 
looks more strongly rounded [Fig. 6 (holotype of O. alticeps, here placed in the 
synonymy of O. feae)] than in the other 6 â examined (Fig. 4). 



THE GENUS ONCOPUS 



145 




Figs 11-26 
Oncopus feae Thorell: 9 lectotype (11, 12, 19, 21, 23); S holotype of O. alticeps (here placed 
in the synonymy of O. feae) (13, 14, 20, 22, 24); 3 other o* 6 (15-17, 25, 26); other 2 (18). - 
Body, lateral (11, 13) and dorsal view (12, 14); anterior part of body, lateral (15-18) and ventral 
view (23-26; 25, 26 showing only central portion); left chelicera (19, 20; arrows pointing to ven- 
troproximal hump of second article) and left palp (21, 22), retrolateral view. - All scale lines 
1.0 mm. 



146 P. J- SCHWENDINGER & J. MARTENS 

Relationships. Oncopus feae is externally similar to 0. doriae, but genital mor- 
phology shows that the former is most closely related to O. malayanus sp. n. 

Distribution. This species is known only from Penang Island [Fig. 1 (2)]. 

Oncopus malayanus sp. n. Figs 27-54 

Oncopus alticeps Pocock. - Silhavy (1974: 25); misidentification. 
Oncopus cfr. alticeps Pocock. - Giribet et al. (2002); molecular study. 

Oncopus feae Thorell. - Silhavy (1974: 25; sub O.feai, incorrect subsequent spelling); misiden- 
tification. 

Material. MALAYSIA (peninsula), Perak, Pangkor Island, forest near Kampung Sungai 
Cempedak, 30 m, 3 holotype, 1 o\ 7 2 paratypes (MHNG), leg. PJ. Schwendinger, 
15./16.XII.1997; Padang Gerus, 200 m, 1 9 paratype (MHNG), leg. P.J. Schwendinger, 
15.1.1995; Maxwell Hill (= Bukit Larut), 1100 m, 1 juvenile (MHNG), leg. G. Cuccodoro & I. 
Lobi, 21. XL 1999; Maxwell Hill, 1500 m, 1 9 paratype, 1 penis (MHNG; with label "Oncopus 
alticeps, det. Silhavy, 1972"), leg. T. Jaccoud, 6.VIIL1969; Chenderiang, 290-330 m, 1 9 
paratype (HMNG), leg. PJ. Schwendinger, 22. -3 1.1.1994; Perak, without exact locality and data, 
1 S paratype (SMF 299/2) with label "Oncopus alticeps Poe, 1 S , Roewer det. 1925", no name 
of collector, no date; Pahang, Cameron Highlands, Tanah Rata, Trail No. 9, 1400 m, 2 9 
paratypes (MHNG), leg. Lobi & Calarne , 27.III.1993; Templer Park (NW of Kuala Lumpur), 
370 m, 1 juvenile (MHNG), leg. P. J. Schwendinger, 13.VII.2001; Selangor, Ulu Gombak, 200 
m, 1 9 paratype, 1 juvenile (MHNG), leg. Agosti, 26.IX.1991; Ulu Gombak, 1 6 paratype 
(penis missing) (MAR), leg. Rohe & Maschwitz, XII. 1988; 30 km N of Kuala Lumpur, 1 6 
paratype (MAR), leg. A. Dorow, 11.1986; Penchala, Kuala Lumpur, 2 9 paratypes (MHNG; with 
label "O.feai, det. Silhavy, 1971"), leg. T Jaccoud, 8.IX.1969. 

Diagnosis. Similar to O. feae, distinguished by: paired paramedian tubercles on 
dorsal scutal area VIII smaller, only little protruding beyond scutal margin; ventral side 
of coxa II with scale-like anteroproximal apophysis in â S , process posterior to it low, 
widely rounded, posteroproximal process present; chelicerae without subbasal ventral 
hump on second article and without subbasai boss on mobile finger; penis with nar- 
rower glans carrying distally pointed lateral sclerites. 

Description. S (holotype). Coloration: body amber, with dark reticulation on 
carapace, chelicerae and palps; dark margin on dorsal scutum and dark pattern on its 
opisthosomal part (Fig. 27a, c); legs dark, except for lighter distal portion of metatar- 
si, lighter tarsi III and IV, and cream tarsi I and II. 

Carapace region of dorsal scutum large and moderately elevated, interocular 
area low, rounded, only slightly projecting beyond carapace front margin (Fig. 27c). 
Opisthosomal areas of dorsal scutum indistinctly elevated in posterior part, with pairs 
of small paramedian tubercles on areas VI- VIII, the last pair slightly protruding beyond 
posterior margin (Fig. 27a, c); ventral scutal areas keeled, with transversal bands of 
white enclosures (Fig. 27b, c). Palpal coxa with long, digitiform ventral process; ven- 
tral side of leg coxa I with conical anterolateral and knob-shaped central processes; 
coxa II with broad scale-like anteroproximal apophysis, with low, rounded process 
posterior to it, and with more pronounced, rounded posteroproximal process over- 
lapping short anteroproximal process on coxa III. Genital operculum rounded, slightly 
longer than wide (Fig. 27b). 

Chelicerae (Fig. 45) strong; proximal article with strong, rounded retroventral 
tubercle; second article with small ventrodistal process and with characteristic pro- 
dorsal subproximal hump (Fig. 45; see arrow); mobile finger without ventral process, 



THE GENUS ONCOPUS 



147 






Fig. 27 
Oncopus malayanus sp. n.: S holotype. - Body, dorsal (a), ventral (b) and lateral view (c). - Scale 
line 1 .0 mm. 



its inner side with sharp, keel-shaped cutting edge only in distal half (worn in proximal 
half?). 

Palps (Fig. 48): trochanter with small, slightly distad-inclined ventral process; 
femur unarmed; patella without proventral process; tibia with distinct retroventral 
proximal hump. 



148 P. J- SCHWENDINGER & J. MARTENS 

Legs 3124; femur II without dorsoproximal tubercle; only low rounded hump 
ventroproximally on femur IV. 

Penis (Figs 28-31): truncus relatively slender, more or less continually widen- 
ing from base to apex, slightly constricted at height of glans; distal margin slightly 
arched, with three and four lateral setae on each side. Glans penis much narrower than 
truncus at that point, flanked by two setae on each side; lateral sclerites slender, with 
pointed tips bent outwards and away from the truncus; median plate broadly rounded, 
almost semicircular; membranous tubes long, protruding far beyond median plate. 

9 (from the type locality). As the S but carapace region of dorsal scutum less 
elevated in posterior part (Fig. 43); ventral scutal areas only slightly elevated, round- 
ed; chelicerae much weaker but with more distinct dorsal hump on second article (Fig. 
46); palps weaker, with small ventroproximal tubercle on femur (Fig. 53); coxa II with 
smaller anteroproximal apophysis, with conical process posterior to it, and with less 
pronounced posteroproximal process. 

Measurements. S holotype (2 from the type locality in parentheses): body 7.50 
(6.79) long, 4.81 (4.26) wide; carapace region 2.41 (1.67) long, 3.21 (2.53) wide. - Palp 
and legs: 





Ti- 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


1.1 1 (0.80) 


1.94(1.27) 


1.11 (0.86) 


0.86 (0.65) 




1.91 (1.54) 


6.93 (5.12) 


Legi 


0.86 (0.74) 


2.65 (2.28) 


1.14(1.05) 


1.14(1.02) 


2.28 (2.04) 


0.43 (0.43) 


8.50 (7.56) 


Leg II 


1.05 (0.86) 


3.46 (2.96) 


1.45 (1.33) 


2.04(1.70) 


3.36 (2.90) 


0.43 (0.43) 


11.79(10.18) 


Leg III 


0.86 (0.74) 


2.25 (1.98) 


1.17(1.05) 


1.11 (0.99) 


2.47 (2.16) 


0.43 (0.43) 


8.29 (7.35) 


Leg IV 


1.11 (1.05) 


2.90 (2.78) 


1.60(1.42) 


1.98(1.79) 


3.95 (3.58) 


0.46 (0.43) 


12.00(11.05) 



Variation. Measurements (o*, in parentheses 9): body 6.88-8.55 (5.90-9.10) 
long, 4.26-5.52 (3.46-6.08) wide; carapace region 1.76-2.93 (1.56-2.35) long, 2.59- 
3.61 (2.16-3.46) wide; 6 (n = 5), 9 (n = 15). Three to five terminal setae present on 
each side of distal penis margin. Variation in external morphological characters 
between and within different populations of O. malayanus sp. n. is more pronounced 
than in other species: 

1 ô from the type locality has a low carapace region (Fig. 42), as otherwise on- 
ly seen in 9 9 (see Figs 43, 44). 

The interocular area is moderately pointed and bent forward in the S from Ulu 
Gombak (Fig. 40) and in the S (Fig. 41) and 9 from Kuala Lumpur, whereas it is 
rounded and not inclined in the other specimens examined (see Figs 27c, 39, 42-44). 

The specimens from Pangkor Island possess a very strong and prominent retro- 
ventral process on the first cheliceral article (Figs 45, 46), in others this process is dis- 
tinctly smaller (Fig. 47). 

The characteristic dorsal hump on the second cheliceral article is distinct in 
specimens from the northern populations (including Pangkor Island; Figs 45, 46, see 
arrows), less pronounced in the 9 9 from the Cameron Highlands and Ulu Gombak 
and in the 6 from "Perak", and absent in specimens from near Kuala Lumpur (Fig. 47). 

Some specimens possess a small ventroproximal process on palpal femur (Figs 
49, 50, 53), others, even within the same population, lack it (Fig. 48). 



THE GENUS ONCOPUS 



149 




Figs 28-38 
Oncopus malayanus sp. n.: S holotype from Pangkor Island (28-31); S paratypes from Ulu 
Gombak (32, 33), Maxwell Hill (34, 35), Kuala Lumpur (36), Pangkor Island (37) and Perak 
(38). - Penis, dorsal (28) and lateral view (29); apex of penis, dorsal (30, 32, 34) and lateral view 
(31); glans penis, lateral (33, 35) and dorsal view (36-38). - Scale lines 1.0 mm (28, 29), 0.1 mm 
(30-38). 



A retroventral proximal hump on the palpal tibia is distinct in most specimens 
(Figs 48, 50), less pronounced in the 2 from Chenderiang, and indistinct in S S and 
2 9 from Kuala Lumpur and Ulu Gombak (Figs 51, 52). 

The 2 2 from Padang Gerus, Maxwell Hill and the Cameron Highlands, as well 
as the S from "Perak" (without exact locality), are all more than 8 mm long and thus 
distinctly larger than the other specimens. Specimens of the northern populations 
appear to be generally larger in size than those of the southern populations. 



150 



P. J. SCHWENDINGER & J. MARTENS 




Figs 39-54 
Oncopus malayanus sp. n.: S holotype from Pangkor Island (45, 48); 6 paratypes from Perak 
(39), Ulu Gombak (40), Kuala Lumpur (41, 51), Pangkor Island (42); 2 paratypes from Pangkor 
Island (43, 46, 53), Cameron Highlands (44), Kuala Lumpur [1 st 2 (47, 52, 54), 2 nd 2 (49)], 
Maxwell Hill (50). - Anterior part of body, lateral view (39-44); left chelicera (45-47; arrows 
pointing to prodorsal subproximal hump of second article) and left palp (48, 49 deformed), retro- 
lateral view; trochanter to tibia (50), patella and tibia (51), and trochanter and proximal part of 
femur (53) of left palp, retrolateral view; right leg III (deformed), retrolateral view (54). - All 
scale lines 1 .0 mm. 



Remark. The 2 2 from Kuala Lumpur possess crippled limbs: in one of them 
the right leg III has an elongated tarsus attached to the femur (Fig. 54), in the other 2 
the shortened palpal tarsus is joined directly to the palpal femur (Fig. 49). 

Relationships. Oncopus malayanus sp. n. is most closely related to O. feae. 

Distribution. This species has a relatively wide distribution in the western cen- 
tral part of Peninsular Malaysia [Fig. 1 (3-8)]. The immature specimens (not examined) 
recorded by Suzuki (1983: 2) from the Templer Park [near Kuala Lumpur; Fig. 1 (8)] 
and from the Pasoh Forest [near Simpang Pertang; Fig. 1 (9); Suzuki, 1976: 9-12, figs 
1-2] presumably also belong to this species. 



the genus oncopus 151 

The truncatus-group 

Diagnosis. Large to very large species (6.2-10.8 mm body length) with inter- 
ocular areas not (or only slightly) forward-inclined and without paramedian tubercles 
on opisthosomal area VIII of dorsal scutum; palps characteristically modified: femora 
with more or less distinct subdistal ventral bulge, patellae with proventral process, 
tibiae very short; penes with widened apex (giving them the shape of a screwdriver) 
and one or two terminal setae on each side of straight distal margin. 

This species group is close to the feae-group and comprises three species (O. 
truncatus, O. tiomanensis sp. n., O. lingga sp. n.) from southern Peninsular Malaysia, 
Singapore and from two Indonesian islands in the Lingga Archipelago. 

Oncopus truncatus Thorell, 1891 Figs 55-91 

Oncopus truncatus Thorell (1891: 764-765); description of juvenile. - Pocock (1897: 286, 288). 

- Roewer (1923: 58-59). - Roewer (1927: 267). - S0rensen (1932: 209). - Bristowe 

(1976: 9). 
Oncopus acanthochelis Roewer (1915: 127, fig. 71), syn. n.; description of 9. - Roewer (1923: 

60, fig. 63). - Martens (1986: fig. 7); illustration of an oncopodid penis, without indication 

as to which species it belongs. - Schwendinger (1992: 194, figs 91-97, pi. 2). 

Type. SINGAPORE, juvenile holotype of O. truncatus (MSNG, examined), leg. L. M. 
D'Albertis, DC.1874. 

Other material. SINGAPORE: 1 9 , holotypus of Oncopus acanthochelis (SMF 1256) 
with label "Oncopus acanthochelis, Type, 1 <?", no name of collector, no date; 1 S, 1 9 (SMF 
298/1) with label "Oncopus truncatus, 2 Exempl., Roewer det. 1926", no name of collector, no 
date; 1 o\ 1 2 (SMF 2113/7) with label "Oncopus acanthochelis, 2 S, Roewer det. 1930", no 
name of collector, no date; 1 o* (MCZ 37715), 1 9 (MCZ 37716), leg. Y. M. Wang, Vm.1965; 
Bukit Timah Nature Reserve, 4 S, 4 9 (MAR 2140), leg. D. H. Murphy, III.-VIII.1967 (pitfall 
traps); 1 6,1 9 (MAR 2141), leg. D. H. Murphy, XI.1965, IIL-Vni.1967 (pitfall traps); 1 o\ 1 
9 (both fragmented, MAR), 1 9 (MHNG), leg. S. Huber, 1999; 1 9 (CCD), leg. C. Deeleman- 
Reinhold, 21. II. 1983; SE Bukit Timah N. R., Rifle Range Road, 50 m, 1 o\ 1 juvenile (MHNG 
SBH-96/3), 1 juvenile (MHNG SBH-96/2), 20.XI.1996, leg. B. Hauser; Labrador Hill, 40 m, 1 
9 , 1 juvenile (MHNG Sar-87/1), leg. B. Hauser, 21.XI.1987; NNW of MacRitchie Reservoir, 60 
m, 1 9 (MHNG SBH-96/5), 21.XI.1996, leg. B. Hauser; Singapore Island Country Club, W of 
MacRitchie Reservoir, 20 m, 1 9 , 2 juveniles (MHNG SL/SI-93/4), leg. B. Hauser, 23VII.1993. 
MALAYSIA (peninsula), Johor, Gunung Pulai (SW of Kulai), 3 o\ 1 2, 3 juveniles (RFS 
2001.2241; 1 S donated to the MHNG), no name of collector, 1935. 

Diagnosis. Distinguished by: genital operculum with pronounced distal 
tubercle; mobile finger of chelicera with small ventroproximal process, large tooth on 
its cutting edge (if present) situated more proximally than tooth on cutting edge of fixed 
finger; palpal femur with subdistal bulge, palpal patella with wide proventral process, 
palpal tibia very short; ventroproximal process on palpal tibia distinct; penis with one 
terminal seta on each side, lateral sclerites of glans tapering. 

Redescription. Very large species (7.5-10.8 mm long). Carapace region of dor- 
sal scutum in Ó* â (Figs 69, 70, 72-74) more elevated than in 9 9 (Figs 75, 76), 
interocular area low, not or only slightly forward-inclined; low median longitudinal 
keel present on anterior areas of opisthosomal region and small paramedian pairs of 
tubercles present on areas V-VII (Fig. 70). Anteroproximal apophysis of coxa II scale- 
like, basally constricted; small process posterior to it and small anteroproximal process 
on coxa III. Genital operculum longer than wide, with distinct ventrodistad tubercle 
(Figs 71, 91). 



152 P. J- SCHWENDINGER & J. MARTENS 

Chelicerae robust, in S S (Figs 77-81) stronger than in 9 9 (Figs 83, 84), with 
truncate or bifid retroventral tubercle and pointed, forward-inclined prolateral distal 
process on first article; ventral side of second cheliceral article with distal process; 
mobile finger of cheliceral hand with more or less pronounced subbasal process; cut- 
ting edge of each cheliceral finger in S S with strong tooth (worn in old S S , Figs 77, 
78; indistinct in 9 9, Figs 83, 84), the one on the mobile finger situated at the same 
position as (Figs 79, 80), or more proximally than (Fig. 81) the one on the fixed finger. 

Pedipalps (Figs 86-89) with short digitiform ventral process on trochanter, with 
more or less distinct ventroproximal process and distinct subdistal ventral transversal 
bulge on femur, and with pointed distad-inclined pro- and retrolateral distal processes 
and low rounded proximal retroventral hump on conspicuously short tibia (Fig. 86); 
palpal patella with wide proventral process in S 6 (Fig. 87; weaker in 9 5). 

Legs: femur of leg II with distinct dorsoproximal tubercle, femur IV with 
distinct ventroproximal one (see Schwendinger, 1992: figs 96, 97). 

Penis (Figs 55-68) fairly slender, markedly widened between glans and apex, 
with straight or slightly rounded distal margin carrying one terminal seta on each side; 
glans narrower than truncus at that point (slightly wider when lateral sclerites pressed 
sideward on temporary slide; see Figs 57, 61), flanked by two to three setae on each 
side; lateral sclerites with wrinkles on lower side, tapering, slightly bent outwards and 
away from the truncus; median plate broadly V-shaped, distally rounded; membranous 
tubes long, protruding far beyond median plate. 

Variation. Measurements (â , in parentheses 9): body 8.71-10.74 (7.42-9.11) 
long, 4.42-5.38 (4.39-5.32) wide; carapace region 2.29-3.12 (2.07-2.41) long, 2.60- 
3.06 (2.82-3.50) wide; S (n = 19), 9 (n = 21). 

In most 6 â the teeth on the cutting edges of the cheliceral fingers are small or 
completely absent (Figs 77, 78). As all of them also show short and blunt (sometimes 
broken) palpal claws, whereas all â 6 with pronounced cheliceral teeth also have 
sharply pointed palpal claws, this is presumably due to wear. 

Penes have a straight or slightly arched distal margin (Figs 59, 61, 63, 67) and 
more or less strongly curved lateral glans sclerites (Figs 66, 60, 58, 62, 64). In most 
S S examined the glans penis is flanked by two setae on each side, in a single 3 (from 
Singapore) by three setae (on both sides). 

Remarks. The juvenile holotype (from Singapore) corresponds with Oncopus 
juveniles from the same island in all relevant characters [subbasal process on mobile 
cheliceral finger (Fig. 85), short palpal tibia (Fig. 90), dorso- and ventroproximal 
tubercle on femur II and IV, respectively]. Therefore we have no doubt that all 
Oncopus specimens from Singapore belong to the same species. 

All 21 9 examined posses a small but distinct ventral tooth on the mobile 
cheliceral finger, which clearly distinguishes them from 9 9 of the closely related 
O. lingga sp. n. (2 9 examined) and of O. tioman sp. n. (6 9 examined). 

The larger tubercle on the genital operculum of O. truncatus (Fig. 91), in 
comparison with that of O. lingga sp. n., only becomes evident when specimens of both 
species are placed side by side. 

Relationships. In its external morphology O. truncatus is most similar to O. 
lingga sp. n., but its penis morphology corresponds more with that of O. tiomanensis 



THE GENUS ONCOPUS 



153 




Figs 55-68 
Oncopus truncatus Thorell: 5 6 from Singapore [1 st (55-58), 2 nd (59, 60), 3 rd (61, 62), 4 th (63, 
64), 5 th (65, 66)]; 2 6 from Gunung Pulai (67, 68). - Penis, dorsal (55) and lateral view (56); 
apex of penis, dorsal (59, 61, 63) and lateral view (62, 64); glans penis, dorsal (57, 65, 68) and 
lateral view (58, 60, 66). - Scale lines 1.0 mm (55, 56), 0.1 mm (57-68). 



sp. n. Which of them are most closely related and sister to the third species remains to 
be established by a character analysis later on. 

Distribution and bionomics. Known from several localities on Singapore Island 
[Fig. 1 (13)] and from a mountain in the south of Peninsular Malaysia [Fig. 1 (12)], 
about 25 km northwest of Singapore. This species appears to be fairly abundant in the 
leaf litter of moist forests; one specimen was even collected at the base of an isolated 
tree on a golf course (B. Hauser, pers. commun.). 

According to a casual observation during night-time by S. Huber, O. trunca- 
tus mates in an upright position with the ventral sides of both partners facing each 



154 



P. J. SCHWENDINGER & J. MARTENS 




Figs 69-91 
Oncopus truncatus Thorell: juvenile holotype (85, 90); S S from Singapore (69-74, 77-80, 
86-88, 91); 2 S from Gunung Pulai (81, 82); 9 2 from Singapore (75, 76, 83, 84, 89). - Body, 
lateral (69) and dorsal view (70); anterior part of body, ventral (71) and lateral view (72-76); left 
cheliceral hand, retrolateral view (77-85), right (86) and left palp (90), retrolateral view; patella 
and tibia of right palp, prolateral view (87); trochanter and femur of left palp, retrolateral view 
(88, 89); genital operculum, lateral view (91). - Scale unes 1.0 mm (69-90), 0.5 mm (91). 



THE GENUS ONCOPUS 155 

other, similar to theraphosid spiders. This is the only existing observation on the 
mating behaviour of oncopodids. These animals are very sensitive to light and air 
movement, and either freeze or run for cover upon the slightest disturbance. 

Oncopus truncatus (and other oncopodids as well) appears to be long-lived: a 
pair of mature specimens was fed with living termites and kept alive for more than two 
years (S. Huber, pers. commun.). 

Oncopus tiomanensis sp. n. Figs 92-112 

Material. MALAYSIA (peninsula), Pahang, Tioman Island, 2 km SE of Kampung 
Genting (2°47'N, 104°08'E), 50-100 m, S holotype (MHNG), 6 9 paratypes, 15 juveniles 
(MHNG, MAR), 23.-28.VI.2001, leg. A. Schulz & K. Vock; Tioman Island, at the foot of 
Gunung Kajang (2°47.181'N, 104°07.892'E), 160 m, 1 S (matured 10.VIII.2002), 1 ? para- 
types (MHNG), 2.X.2001, leg. L. Monod. 

Etymology. The specific epithet refers to the island where the new species was found. 

Diagnosis. Similar to O. truncatus, distinguished by: smaller in size; no process 
on mobile finger of chelicerae; cutting edges of cheliceral fingers in S S without 
prominent teeth; palps with subdistal-ventral bulge of femur and proventral process of 
patella less developed; carapace region of â â not more elevated than in $ $ , inter- 
ocular area not directed forward; paramedian tubercles on dorsal scutal areas V-VII 
less pronounced; genital operculum much wider, without anterior tubercle; dorso- 
proximal tubercle on femur II and ventroproximal tubercle on femur IV much shorter; 
penis with almost quadrangular apex, separated from rest of truncus by a more abrupt 
constriction at height of glans. 

Description. S (holotype). Coloration: body reddish amber, carapace region, 
chelicerae and palps mottled with dark; dorsal scutum with dark margin and dark 
opisthosomal areas; proximal article of chelicerae and legs dark brown, except for light 
amber leg tarsi and distal portion of metatarsi; tarsi I and II cream. Ventral side of body 
and distal articles of chelicerae yellowish amber. 

Carapace region of dorsal scutum fairly low, not arched, interocular area low, 
rounded, not projecting beyond carapace front margin (Fig. 98). Opisthosomal areas of 
dorsal scutum indistinctly elevated in posterior part, with pair of small paramedian 
tubercles on areas VI- VII (Figs 98, 99); ventral scutal areas moderately keeled, with 
transversal bands of white enclosures (Fig. 98). Palpal coxa with long, conical ventral 
process; ventral side of leg coxa I with small anterolateral process and knob-shaped 
central tubercle; coxa II with scale-like anteroproximal apophysis and with low, 
rounded process posterior to it; posteroproximal process of coxa II absent; antero- 
proximal process on coxa III indistinct. Genital operculum rounded, clearly wider than 
long, without distoventral tubercle (Fig. 100). 

Chelicerae (Fig. 106) strong; proximal article with conical retroventral tubercle 
and with anteriad-inclined prodorsal boss extending into rounded dorsomedian boss; 
second article with small ventrodistal process; mobile finger without ventral process, 
its inner side with sharp cutting edge only in distal half (presumably worn in proximal 
half); both cutting edges of cheliceral hand without prominent teeth. 

Palps (Figs 109, 110): trochanter with small, slightly distad-inclined ventral 
process; ventral side of femur with indistinct subdistal bulge and indistinct proximal 



156 P. J- SCHWENDINGER & J. MARTENS 

process; patella with widely rounded proventral process (Fig. 110); tibia with low 
retro ventral proximal hump. 

Legs 3142; low rounded hump present dorsoproximally on femur II and 
ventroproximally on femur IV. 

Penis (Figs 92-95): truncus fairly slender, more or less continually widening 
from base to height of glans, there abruptly widening to subquadrangular apex; distal 
margin almost straight, with single terminal seta on each side. Glans penis about as 
wide as truncus at that point (wider when lateral sclerites pressed sideward on tempo- 
rary slide), flanked by two setae on each side; lateral sclerites slender, with pointed tips 
bent outwards and upwards; median plate broadly rounded, widely V-shaped; mem- 
branous tubes long, protruding far beyond median plate. 

$ . As the S , but interocular area slightly more raised (Fig. 102), ventral scutal 
areas not elevated; chelicerae weaker (Fig. 108); palps weaker, subdistal bulge on 
ventral side of femur (Fig. Ill) and proventral process on patella (Fig. 112) less 
developed. 

Measurements. S holotype (2 in parentheses): body 6.25 (6.18) long, 3.56 
(3.63) wide; carapace region 1.75 (1.53) long, 2.32 (2.15) wide. - Palp and legs: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.74 (0.65) 


1.63(1.23) 


1.01 (0.79) 


0.64 (0.49) 


-- 


1.68(1.38) 


5.70 (4.54) 


Legi 


0.67 (0.62) 


2.03 (1.78) 


1.04(0.91) 


0.89 (0.84) 


1.80(1.53) 


0.49 (0.40) 


6.92 (6.08) 


Leg II 


0.86 (0.79) 


2.82 (2.47) 


1.33(1.19) 


1.73(1.51) 


2.96 (2.42) 


0.49 (0.40) 


10.19(8.78) 


Leg III 


0.69 (0.62) 


1.75(1.58) 


1.01 (0.91) 


0.89 (0.84) 


1.88(1.58) 


0.49 (0.40) 


6.71 (5.93) 


Leg IV 


0.89 (0.79) 


2.47 (2.22) 


1.43(1.31) 


1.63 (1.48) 


3.06 (2.67) 


0.59 (0.47) 


10.07 (8.94) 



Variation. Measurements of S (of 9 in parentheses): body 6.25-6.37 (5.88- 
6.67) long, 3.51-3.56 (3.41-3.98) wide; carapace region 1.75-1.78 (1.48-1.68) long, 
2.17-2.32 (2.07-2.27) wide; Ó* (n = 2), $ (n = 7). The 6 paratype has (like the 9 9) 
a slightly more raised interocular area (Fig. 101) than the holotype. The cutting edges 
of its cheliceral hand appear less worn than in the holotype but they also carry no 
prominent teeth (Fig. 107). 

Remark. The right leg IV of the holotype is crippled; only a stump-like 
trochanter is present. 

Relationships. External and genital characters show that O. tiomanensis sp. n. 
belongs to the truncatus-group and that it probably is more closely related to O. trun- 
catus (more congruence in penis details) than to O. lingga sp. n. 

Distribution. Known only from Tioman Island, off the southeastern coast of 
Peninsular Malaysia [Fig. 1 (11)]. 

Oncopus lingga sp. n. Figs 113-131 

Material. INDONESIA. Riau Province, Lingga Archipelago, Lingga Island, near water- 
fall ca. 7 km N of Daik (0°08'53.3"S, 104°36'13.2"E), 60 m, S holotype (MHNG), leg. P. J. 
Schwendinger, 15.VI.2001; Singkep Island, Batu Ampar Waterfall, ca. 10 km NW of Dabo 
(0°29'30.9"S, 104°28'31.4"E), 80 m, 2 9 paratypes, 2 juveniles (MHNG), leg. P. J. Schwen- 
dinger, 17./18.VI.2001. 



THE GENUS ONCOPUS 



157 



92 








Figs 92-97 
Oncopus tiomanensis sp. n.: 6 holotype (92-95); â paratype (96, 97). - Penis, dorsal (92) and 
lateral view (93); apex of penis, dorsal (94, 96) and lateral view (95, 97). - Scale lines 1.0 mm 
(92, 93), 0.1 mm (94-97). 

Etymology. The specific epithet (a noun in apposition, hence invariable) refers to the ar- 
chipelago in which the new species occurs. 

Diagnosis. Close to O. truncatus, distinguished by: mobile finger of cheliceral 
hand ventrally with low mound instead of prominent tooth, dorsal and ventral proxi- 
mal tubercles on leg II and IV smaller, tubercle near distal margin of genital operculum 
smaller. S with more elevated interocular area, without subdistal ventral bulge on 
palpal femur, with stronger proventral process on palpal patella, and with large 
rounded tooth on cutting edge of mobile cheliceral finger situated more distally than 
tooth on cutting edge of fixed finger. Penis distinguished by two terminal setae on each 
side of distal margin; glans with wide, W-shaped median plate. 

Description. S (holotype). Coloration: body amber, with dark margin around 
dorsal scutum and dark transversal bands on its opisthosomal elevations (medially 
connected by dark transversal stripe); dark reticulations in carapace region and on 
chelicerae (proximal article darker than hand); legs and palps dark amber, except for 
light amber tarsi and distal part of metatarsi. 



158 



P. J. SCHWENDINGER & J. MARTENS 




Figs 98-112 
Oncopus tiomanensis sp. n.: 6 holotype (98-100, 106, 109-110); S paratype (101, 104, 107); 2 
9 paratypes (102, 103, 105, 108, 111, 112). - Body, lateral (98) and dorsal view (99); anterior 
part of body, ventral (100) and lateral view (101-103); genital operculum (104, 105); left 
chelicera, retrolateral view (106-108); left palp, retrolateral view (109, 111); patella and tibia of 
left palp, prolateral view (110, 112). - All scale lines 1.0 mm. 



Carapace large and elevated, interocular area a low rounded hump (Fig. 117). 
Opisthosomal areas of dorsal scutum indistinctly keeled, with indistinct longitudinal 
median keel extending over anterior areas and with pair of paramedian tubercles on 
areas V-VII (increasing in size towards rear) (Fig. 118); areas of ventral scutum 
strongly keeled, with indistinct transversal bands of white enclosures (Fig. 117). Palpal 
coxa with large ventral process; ventral side of leg coxa I with anterolateral and central 
processes; coxa II with scale-like anteroproximal apophysis and rounded process 



THE GENUS ONCOPUS 159 

posterior to it; coxa III with triangular anteroproximal process. Genital operculum 
distinctly longer than wide, with small tubercle near anterior margin (Fig. 119). 

Chelicerae (Figs 124, 125) massive; proximal article with two-headed retro- 
ventral tubercle and anteriad-inclined prodorsal boss; second article with small 
ventrodistal tubercle, mobile finger with indistinct ventral hump on proventral face; 
large rounded tooth on cutting edge of mobile finger situated more distally than large 
conical tooth on cutting edge of fixed finger. 

Palps (Figs 128-130) robust; trochanter with small ventral process; femur 
unarmed; patella with large subquadrangular proventral process (Figs 129, 130); tibia 
short, with low retroventral process near its base; tarsus subdistally widened (Fig. 128). 

Legs 3142; femur II with dorsoproximal and femur IV with ventroproximal 
tubercle. 

Penis (Figs 113-116): truncus fairly slender, with subbasal constriction, 
continually widening in distal half, slightly constricted at height of glans, widened at 
apex; distal margin straight, with two setae on each side. Glans penis narrower than 
truncus at that point; lateral scleri tes pointed, bent outwards and away from the 
truncus; median plate wide, indistinctly W-shaped; membranous tubes long, distinctly 
protruding beyond median plate. 

9 . As the 6 , but smaller, carapace region less elevated and shorter, ventral 
scutal areas less elevated (Figs 120, 123); chelicerae weaker, teeth on their cutting 
edges smaller (Figs 126, 127); ventral side of palpal femur with small proximal process 
and indistinct subdistal bulge, patella with smaller proventral process (Fig. 131); legs 
3124. 

Measurements. 6 holotype (9 in parentheses): body 7.93 (7.00) long, 4.75 
(4.04) wide; carapace region 2.75 (1.98) long, 3.30 (2.47) wide. - Palp and legs: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


1.17 (0.80) 


2.72(1.54) 


1.48 (0.93) 


1.08 (0.68) 


-- 


2.28 (1.60) 


8.73 (5.55) 


Legi 


0.86 (0.68) 


3.02 (2.10) 


1.30 (1.64) 


1.33 (1.64) 


2.62(1.82) 


0.52 (0.40) 


9.65 (8.28) 


Legn 


1.17 (0.80) 


3.95 (2.90) 


1.73 (1.36) 


2.41 (1.79) 


4.01 (2.80) 


0.49 (0.43) 


13.76 (10.08) 


Leg in 


0.86 (0.74) 


2.41 (1.88) 


1.36 (1.08) 


1.23 (0.99) 


2.69 (1.94) 


0.56 (0.52) 


9.11 (7.15) 


Leg IV 


1.17 (0.93) 


3.27 (2.59) 


1.94 (1.48) 


2.31 (1.82) 


3.55 (3.15) 


0.49 (0.56) 


12.73 (10.53) 



Variation. The second 9 measures: body length 7.19, width 4.16; carapace 
length 1.98, width 2.59. 

Remarks. As the 9 9 examined are clearly smaller than the S holotype and 
were collected from a different island, it is possible that they are not conspecific. On 
the other hand, they appear to be sufficiently distinct from 9 9 of O. truncatus and 
therefore we regard them as belonging to O. lingga sp. n. 

The ô holotype was collected immediately after its final moult (when the body 
was still unpigmented) and then kept in captivity for a few weeks; its cheliceral cutting 
edges and palpal claws therefore show no signs of wear. 

Living mature specimens have a clearly visible white membrane dors ally 
between the proximal and second cheliceral article (Figs 118, 121), though this 
becomes less distinct in alcohol. 



160 



P. J. SCHWENDINGER & J. MARTENS 




Figs 113-116 
Oncopus lingga sp. n.: 3 holotype. - Penis, dorsal (113) and lateral view (114); apex of penis, 
dorsal (115) and lateral view (116). - Scale lines 1.0 mm (113, 114), 0.1 mm (115, 116). 

Relationships. The new species is most closely related to O. truncatus. Both 
species share a similar penis morphology and a very similar external morphology (e.g., 
short palpal tibia; pronounced pro ventral process on palpal patella of males; subdistal 
ventral bulge on palpal femur of females; dorsoproximal and ventroproximal tubercle 
on femur of leg II and IV, respectively; anterior tubercle on genital operculum). 
Oncopus tiomanensis sp. n. is phylogenetically more distant from O. lingga sp. n. 

Distribution. Known only from two islands (separated from each other by less 
than 10 km) south of the Malayan Peninsula [Fig. 1 (14, 15)]. This species and a 
presumably undescribed one from southern Kalimantan (see "Unidentified material") 
are the only Oncopus occurring south of the equator. 

The doriae-group 

Diagnosis. See diagnosis of O. doriae. 

At present this species group contains only a single species, O. doriae, from the 
western part of Sarawak. This species appears to be more closely related to the feae- 
group and the truncatus-gioup from the Malayan Peninsula and nearby islands than to 
congeners from northeastern Borneo. 



Oncopus doriae Thorell, 1876 



Figs 132-151 



Oncopus doriae Thorell (1876: 135-138); description of 2. - Thorell (1891: 764-765, figs 
31-36). - Pocock (1897: 288). - Roewer (1923: 57-58, fig. 60). - Bristowe (1976: 9). - 
Sorensen (1932: 209). - Schwendinger (1992: 197). 

Oncopus alticeps Pocock (= O.feae Thorell). - Roewer (1923: 60); misidentification of specimen 
SMF 1261. 



THE GENUS ONCOPUS 



161 




Figs 117-131 
Oncopus lingga sp. n.: 3 holotype (117-119, 124, 125, 128-130); 2 ? paratypes (120-123, 126, 
127, 131). - Body, lateral (117, 120) and dorsal view (118, 121); anterior part of body, ventral 
(119, 122) and lateral view (123); left chelicera (124), right chelicera (125) and left cheliceral 
hand (126, 127), retrolateral view; left palp (128) and trochanter to tibia of left palp (131), retro- 
lateral view; patella and tibia of left palp, prolateral view (129); patella and tibia of right palp, 
retroventral view (130). - All scale lines 1.0 mm. 



162 P. J- SCHWENDINGER & J. MARTENS 

Types. MALAYSIA (east), Sarawak, 1 ? , 1 juvenile syntypes (MSNG, examined), leg. 
G. Doria & O. Beccari, 1866; 9 here designated lectotype in order to preserve stability of 
nomenclature. 

Other material. MALAYSIA (east), Sarawak, Santubong, 32 km N of Kuching, 0-100 
m, 1 o\ 1 9, 2 juveniles (MHNG), leg. I. Lobi & D. Burckhardt, 11.-16.V.1994; Gunung 
Santubong (south end), 1 9 (California Academy of Science), leg. J. E. Tobler, 15. IV. 1970; Batu 
Lundu (presumably near Lundu), 1 9 (SMF 540/5) with label "Oncopus doriae Thorell, 1 
Exempl., Roewer det. 1927", no name of collector, no date; Gunung Gading National Park, 300 
m, NW of Lundu, 1 juvenile (MHNG), leg. S. Kurbatov, 18.X.2001; Gunung Serapi (near 
Matang), 320 m, 1 juvenile (MHNG), leg. B. Hauser, 9.XH1987; Bako National Park, 2 juve- 
niles (CCD), leg. C. Deeleman-Reinhold & P. Deeleman, 29./30.III.1985; Bako N. P., 1 juvenile 
(MAR), leg. A. Riedel, 27.m. 1990; Balai Ringin, 40 km E of Serian, 1 juvenile (CCD), leg. C. 
Deeleman-Reinhold, 7.IV.1985; Gunung Penrissen, 1000 m, 2 juveniles (MHNG), leg. I. Lobi & 
D. Burckhardt, 23.V.1994. - THAILAND, without exact locality (dubious record), 1 S (SMF 
1261), fragmented specimen, with label "Oncopus alticeps Pocock, Siam, 1 S, Roewer det. 
1914", no name of collector, no date. 

Diagnosis. Distinguished by: interocular area forward-inclined; pair of para- 
median tubercles projecting beyond posterior margin of dorsal scutum; palpal coxa 
with two ventral processes; mobile finger of chelicerae in S S with transversal ledge 
and wide ventroproximal boss; palpal femur with subdistal ventral process; penis with 
very large, paddle-shaped apex carrying two setae on each side of distal margin, lateral 
sclerites of glans distally widening, median plate long, V- or narrowly U-shaped. 

Redescription. Large species (7.1-8.4 mm long). Carapace region of dorsal 
scutum in S S (Fig. 138) arched and more elevated than in 9 9 (Fig. 141), with 
forward-inclined interocular area, rounded in S S (Figs 138, 139), more pointed in 
9 9 (Figs 141-143); opisthosomal region of dorsal scutum with indistinct longitudinal 
median keel extending over areas I-ILT (Fig. 142) and with small paramedian pairs of 
tubercles on areas VI- VIII, the last pair clearly projecting beyond the posterior scutal 
margin (Figs 138, 139, 141, 143); palpal coxa with two pointed ventral tubercles on 
moderately developed common socket; leg coxa II with scale-like anteroproximal 
apophysis, process posterior to it developed as a hump (strongly elevated in SS) 
gradually sloping down to the posteroproximal corner, the latter without process; 
anteroproximal process on coxa III indistinct; genital operculum longer than wide 
(Fig. 140). 

Chelicerae robust, in S S (Figs 144, 145) stronger than in 9 9 (Figs 146, 147), 
with rounded retroventral tubercle on first article; ventral side of second cheliceral 
article with strong distal process; mobile finger of chelicera in S S with broad ventral 
hump and with pronounced transversal ledge in proximal half of retrolateral side (Figs 
144, 145; both characters absent in 9 9); cutting edge of cheliceral fingers without 
marked teeth. 

Pedipalps (Figs 148-151) with quite long, pointed ventral process on trochanter, 
with small proximal and small subdistal process on ventral side of femur (weaker in 
SS, Figs 148, 149), and with small retroventral hump on tibia; no proventral process 
on palpal patella. 

Legs: femur of leg II with small prodorsal proximal tubercle, femur IV without 
ventral tubercle. 

Penis (Figs 132-137) fairly slender, gradually widening towards the apex, with 
constriction at height of glans; apex exceptionally long, widely paddle-shaped, with 



THE GENUS ONCOPUS 



163 



136 



132 133 





Figs 132-137 
Oncopus doriae Thorell: 6 from Gunung Santubong (132-135); 6 labelled as originating from 
Siam (136, 137). - Penis, dorsal (132) and lateral view (133); apex of penis, dorsal (134, 136) 
and lateral view (135, 137). - Scale lines 1.0 mm (132, 133), 0.1 mm (134-137). 



broadly rounded distal margin carrying two setae on each side (distal seta arising from 
distal edge of truncus, more proximal seta originating from distoventral side close to 
edge; Figs 134, 136); glans about as wide as truncus at that point, flanked by two to 
three setae on each side; lateral sclerites broad, bent outwards and away from the 
truncus, with broadly truncate, dentate tips; median plate long and narrow, V-shaped 
(Fig. 136) or tongue-shaped (Fig. 134); membranous tubes short, not or only slightly 
protruding below median plate. 

Variation. Measurements (<$, in parentheses 9): body 7.92-8.39 (7.08-8.17) 
long, 4.88-5.25 (4.35-4.91) wide; carapace region 2.83-3.11 (1.93-2.27) long, 3.32- 
3.66 (2.48-3.07) wide; S (n = 2), 9 (n = 4). In 2 9 (including the lectotype) and both 
S a examined the median longitudinal keel on opisthosomal areas I-III is discernible 
in dorsolateral view (Fig. 142), in the other specimens it is not. The genital operculum 
of the 9 lectotype and of one S (Fig. 140) is slightly longer than wide, in the other 
specimens it is as wide as long or slightly wider than long. The proventral process on 



164 



P. J. SCHWENDINGER & J. MARTENS 




Figs 138-151 
Oncopus doriae Thorell: 9 lectotype (141-143, 147, 150); 9 (146, 151) and S (138-140, 144, 
148) from Gunung Santubong; $ labelled as originating from Siam (145, 149). - Body, lateral 
(138, 141) and dorsal view (139, 143); anterior part of body, ventral (140) and dorsolateral view 
(142); left chelicera (144, 146, 147) and left cheliceral hand (145), retrolateral view; left palp 
(148), trochanter and femur (150, 151) of left palp, femur (149) of left palp, retrolateral view. - 
All scale lines 1.0 mm. 



palpal coxa is either pointed and as long as the retrolateral one, or shorter and more 
rounded. 

Relationships. O. doriae is very distinct from all other congeners and its 
relationships are not clear. Judging from penis morphology, it appears that this species 
is more closely related to the/eae-group (closest to O.feae) and to the truncatus- group 
than to O. hosei and O. megachelis. 



THE GENUS ONCOPUS 165 

Distribution. This species is quite widely distributed in the western part of 
Sarawak [Fig. 1 (17-22)]. The types presumably originate from near Kuching. Beccali 
and Doria collected together in this area until Doria had to return to Italy (because of 
deteriorating health) in March 1866 (Beccali, 1904). The â with the locality label 
"Siam" (written by Roewer) undoubtedly belongs to O. doriae, but it is very unlikely 
that this species occurs in western Sarawak and in Thailand and seemingly nowhere 
else in between. Therefore we assume that Roewer has added an incorrect label (see 
also the paragraph "Distribution" under O. expatriatus sp. n.). 

The hosei-grovv 

Diagnosis. Large species (7.0-9.5 mm body length) characterized by penis mor- 
phology: distal margin of truncus sinuate, with single terminal seta on elevated socket 
on each side; subdistal lateral setae arranged in one or two recurved rows; lateral scle- 
rites of glans basally with knee-like lateral projections (as seen in dorsal view); medi- 
an plate very wide. 

This species group contains three species (O. hosei, O. megachelis, O. expa- 
triatus sp. n.), presumably all from the northeastern part of Borneo (record of O. 
expatriatus sp. n. from Thailand dubious). Relationships remain unclear; in penis 
morphology these species are very distinct from all other congeners. 

Oncopus hosei Pocock, 1897 Figs 152-158 

Oncopus hosei Pocock (1897: 285-287, fig. 1); description of S. - Roewer (1923: 59, fig. 61). - 
Bristowe (1976: 9). - Schwendinger (1992: 194-196, figs 98-101). 

Type. MALAYSIA (east), Sarawak, Baram, 6 holotype (NHML 1895.10.15, examined), 
leg. C. Hose, collected before 1895. 

Other material. MALAYSIA (east), Sarawak, Lambir Hills National Park, south of Miri, 
1 juvenile (MHNG), leg. S. Kurbatov, 11.-13.X.2001. 

Diagnosis. Distinguished by: interocular area strongly forward-inclined; cara- 
pace region fairly low (clearly lower than in 6 S of O. doriae); paramedian tubercles 
on opisthosomal area VIII situated close to each other and projecting beyond posterior 
margin of dorsal scutum; anteroproximal process on coxa III long; mobile finger of 
chelicera without ventral tubercle; palpal femur without ventral process; truncus penis 
with one terminal seta on prominent socket on each side of distal margin, subterminal 
lateral setae arranged in two recurved rows; glans penis with widely rounded median 
plate and truncate lateral sclerites with bases bulged laterally. 

Redescription. Large species (dorsal scutum of a holotype 9.5 long, 5.2 wide; 
carapace region 2.8 long, 3.4 wide) with fairly slender body (Fig. 156). Carapace 
region only moderately raised, interocular area distinctly forward-inclined (Fig. 157); 
opisthosomal areas VI- VIII with paired paramedian tubercles, posterior ones largest, 
situated close to each other and projecting beyond posterior scutal margin (Fig. 156). 
Palpal coxa with low, rounded ventral process; leg coxa II with indistinct antero- 
proximal apophysis (as in Gnomulus species), process posterior to it developed as low 
bulge, posteroproximal process absent; anteroproximal process on coxa III long, 
pointed, triangular; genital operculum distinctly longer than wide (Fig. 158). 



166 



P. J. SCHWENDINGER & J. MARTENS 




Figs 152-158 
Oncopus hosei Pocock: S holotype. - Penis, dorsal (152) and lateral view (153); apex of penis, 
dorsal (154) and lateral view (155); body, dorsal view (156); anterior part of body, lateral view 
(157) and ventral view (158, showing only central portion). - Scale lines 1.0 mm (152, 153, 156- 
158), 0.1 mm (154, 155). 



Chelicerae (see Schweninger, 1992: fig. 100) robust, with low, domed retro- 
ventral process on proximal article, a small distoventral one on second article, none on 
mobile finger. 

Palp (see Schwendinger, 1992: fig. 101) with small pointed ventral process on 
trochanter and with low ventroproximal hump on tibia, no process on femur. 

Legs: femur of leg II with distinct dorsoproximal tubercle. 

Penis (Figs 152-155): truncus fairly slender, with sinuate distal margin carrying 
one lateral seta on each side, each seta raised on small mound; subterminal lateral setae 
arranged in two more or less clearly outlined recurved rows; glans penis narrower than 
truncus at that point, its membranous base rounded, flanked by three setae on each side; 
median plate widely rounded, not covering tips of long membranous tubes; lateral 
sclerites with wide, rounded bases bulged sideward like "knees" and with truncate tips 
bent away from truncus. 



THE GENU'S ONCOPUS 167 

Remarks. The holotype (with a pin running through its longitudinal axis) has a 
brittle cuticle and shows an unusually dark coloration (mentioned by Schwendinger, 
1992: 194). This is possibly the result of an unusual preservation method. 

The juvenile specimen is quite similar to the holotype (different only in inter- 
ocular area less forward-inclined and anteroproximal process on coxa III shorter), but 
shows a normal (amber) coloration. 

Relationships. Penis morphology shows a close relationship between O. hosei 
and O. megachelis. The penis of both species differ only in relatively minor details, i.e. 
O. hosei has a stouter truncus with a wider, laterally more rounded apex and narrower, 
distally more divergent, apically rounded lateral sclerites (Fig. 154, cf. Fig. 159). 

Distribution. Known from the type locality, somewhere along the Baram River 
{possibly near Kuala Baram [Fig. 1 (25)]}, and from the Lambir Hills N. P. [Fig. 1 
(24)], both in the northeastern part of Sarawak. 

Oncopus megachelis Schwendinger, 1992 Figs 159-161 

Oncopus megachelis Schwendinger (1992: 192-194, figs 81-90); description of S . 

Types. MALAYSIA (east), Sabah, Sandakan Residency, Sepilok, Kabili-Sepilok Forest 
Reserve, 40 m, 6 holotype, juvenile paratype (both in MHNG, examined), leg. B. Hauser, 
10.V.1982. 

Diagnosis. Large species (8.1 mm body length), similar to O. hosei, distin- 
guished by: interocular area more rounded (Fig. 160); no paramedian tubercles on 
opisthosomal area VIII of dorsal scutum; genital operculum wider than long (Fig. 161); 
palpal femur with small ventroproximal process; penis stouter, its subterminal lateral 
setae mostly arranged in one recurved row (a few setae in the middle in two rows); 
lateral sclerites of glans penis narrower, more strongly bent outwards, distally rounded, 
bases more distinctly knee-shaped (Fig. 159). 

Relationships. This species is most closely related to O. hosei. 

Distribution. Known only from the type locality, in the eastern part of Sabah 
[Fig. 1(26)]. 

Oncopus expatriates sp. n. Figs 162-171 

Material. THAILAND, near Bangkok (dubious record), S holotype (SMF 8658/13) 
with label reading "Pelitnus alticeps Pocock, 1 S , Siam: bei Bangkok, Roewer det. 1940", no 
name of collector, no date. 

Etymology. Latin participle: expatriatus - expatriate. The specific epithet refers to the 
questionable origin of the holotype. 

Diagnosis. Similar to O. megachelis, distinguished by: interocular area less pro- 
truding; anteroproximal process of leg coxa III distinctly smaller; truncus penis stouter, 
slightly constricted in its middle portion, its subterminal lateral setae arranged in two 
rows; glans penis shorter, its lateral sclerites more strongly bent upwards and with 
more strongly bent knee-shaped lateral bases, its median plate with straight distal 
margin and strong lateral teeth. 

Description. 6 (holotype). Coloration (faded): body light amber throughout, 
with slightly darker markings on dorsal scutum. 

Carapace region of dorsal scutum large and raised, interocular area a low 
rounded hump (Fig. 166). Opisthosomal areas of dorsal scutum indistinctly keeled, 



168 



P. J. SCHWENDINGER & J. MARTENS 




159 




Figs 159-161 
Oncopus megachelis Schwendinger: S holotype. - Apex of penis, dorsal view (159); anterior 
part of body, lateral (160) and ventral view (161, showing only central portion). - Scale lines 
0.1 mm (159), 1.0 mm (160, 161). 

with pairs of small paramedian tubercles on areas VII and VIII, posterior pair slightly 
protruding from posterior margin of scutum (Figs 166, 167); ventral scutal areas dis- 
tinctly keeled, with transversal bands of white enclosures (Fig. 166). Palpal coxa with 
long, digitiform ventral process; ventral side of leg coxa I with distinct anterolateral 
and indistinct central processes; coxa II with small scale-like trapezoidal antero- 
proximal apophysis and with small conical process posterior to it; coxa III with small 
anteroproximal process. Genital operculum rounded, longer than wide (Fig. 168). 

Chelicerae (Figs 169, 170) strong; proximal article with small, rounded retro- 
ventral tubercle and anteriad-inclined prodorsal boss (Fig. 170); second article with 
small ventrodistal tubercle, mobile finger without ventral process; cutting edge of mo- 
bile finger with conical tooth fitting between two teeth on cutting edge of fixed finger. 

Palps (Fig. 171): trochanter with strong, distad-inclined ventral process; femur 
with only an indistinct ventrobasal hump, otherwise unarmed; patella without proven- 
tral process; tibia with quite large retroventral proximal process. 

Legs 3124; femur II with dorsoproximal tubercle; only a low rounded hump 
ventroproximally on femur IV 

Penis (Figs 162-165): truncus remarkably stout, more or less continually 
widening from base to apex, slightly constricted after the proximal two-thirds, with 
subterminal lateral setae arranged in two irregular recurved rows; distal margin of trun- 
cus straight, each side with one terminal seta raised on low socket. Glans penis quite 
short, much narrower than truncus at that point, flanked by two setae on each side; 
lateral sclerites bent outwards and away from truncus, their wide bases strongly pro- 
truding laterally, their apices truncate; median plate with straight distal margin and 
with pair of strong lateral teeth; membranous tubes distinctly protruding beyond 
median plate. 

$ . Unknown. 

Measurements. S holotype: body 7.04 long, 4.28 wide; carapace region 2.26 
long, 2.83 wide. - Palp and legs: 



THE GENUS ONCOPUS 



169 







Figs 162-171 
Oncopus expatriatus sp. n.: <? holotype. - Penis, dorsal (162) and lateral view (163); apex of 
penis, dorsal (164) and lateral view (165); body, lateral (166) and dorsal view (167); anterior part 
of body, ventral view (168); left chelicera (169) and left palp (171), retrolateral view; proximal 
article of left chelicera, retrodorsal view (170). - Scale lines 1.0 mm (162, 163, 166-171), 0.1 mm 
(164, 165). 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.89 


1.77 


0.93 


0.79 


- 


1.77 


6.15 


Legi 


0.74 


2.41 


1.08 


1.13 


2.21 


0.49 


8.06 


Legn 


0.93 


3.20 


1.43 


1.87 


3.27 


0.34 


11.04 


Leg III 


0.79 


1.92 


1.18 


1.13 


2.34 


0.49 


7.85 


LeglV 


0.98 


2.56 


1.62 


1.87 


3.69 


0.54 


11.26 



170 P. J- SCHWENDINGER & J. MARTENS 

Remarks. The holotype has been pinned through its prosoma (not indicated in 
Fig. 167), but is otherwise in perfect condition. 

According to Roewer's substitute label of 1940 (Roewer replaced many original 
labels in his collection), the holotype was collected near Bangkok, but we strongly 
doubt if this is correct. If so, this would be the only Oncopus specimen ever found in 
Thailand (not considering the obviously incorrectly labelled S of O. doriae in 
Roewer's collection; see paragraph "Distribution" in the treatment of O. doriae), about 
900 km away from the closest locality of another Oncopus (i.e. O. feae on Penang 
Island). No indication is given as to when the type was collected and by whom, and 
Roewer was notorious for giving inaccurate localities (see Helversen & Martens, 
1972). On the other hand, strong similarities in the penes of O. expatriatus sp. n., O. 
hosei and O. megachelis indicate a close relationship between these three species, and 
therefore we assume that O. expatriatus sp. n. actually occurs in northern Borneo. We 
speculate that Roewer possibly has received the type from the collection of Stanley 
Flower, who was working at the Royal Museum of Bangkok and who also collected 
the type of O. alticeps (here placed in the synonymy of O. feae) (see Pocock, 1897: 
287). No confirmation of this was found among Roewer's documents in the SMF (P. 
Jäger, pers. commun.). There is, however, an entry for this specimen (serial number 
8658, tube number 13) in the collection files of Roewer, in which "Borneo" (in blue 
ink, as for other entries on the same page) was overwritten by "Siam" (in black ink). 

In the case that the holotype of O. expatriatus sp. n. really originates from near 
Bangkok, then there is little hope that the corresponding population of this species still 
exists there today. Leaf litter sifting in semi-evergreen rainforest in the lowland and 
lower montane rainforest (terminology according to Whitmore, 1991) of Khao Yai 
National Park (about 90 km NE of Bangkok; the most promising locality to find 
oncopodids in this area) by I. Lobi & D. Burckhardt and by P.J. Schwendinger has 
revealed the presence of Gnomulus, but not of Oncopus (see Schwendinger & Martens, 
2002: 107). 

Relationships. Congruencies in genital morphology (terminal setae of truncus 
penis resting on sockets; subdistal lateral setae arranged in one or two curved rows; 
glans penis with a very wide median plate and with laterad-directed "knees" at the base 
of the lateral sclerites) and in external morphology (i.e. indistinctly developed antero- 
proximal apophysis on ventral side of leg coxa II; no ventral process on mobile finger 
of chelicerae) indicate a close relationship between O. expatriatus sp. n. and the sister 
species O. megachelis and O. hosei. 

Distribution. According to its dubious label data, the holotype of O. expatriatus 
sp. n. was found in central Thailand [Fig. 1 (1)]. Morphological characters, however, 
indicate that this species actually occurs on Borneo (see remarks above). 

UNIDENTIFIED MATERIAL 

MALAYSIA (peninsula), Pahang, Bukit Charas [Fig. 1 (10)], 60 m, near Panching, 30 
km NW of Kuantan, 1 juvenile (MHNG), leg. P.J. Schwendinger, 8.VTL2001. This specimen 
possibly belongs to O. malayanus sp. n. 

MALAYSIA (east), Sarawak, confluent of Suan Oyan and Mujong rivers [Fig. 1 (23)], 
150m, east of Kapit, 1 juvenile (MHNG), leg. I. Lobi & D. Burckhardt, 19.V.1994. This quite 
small specimen does not show the characteristics of O. doriae and of O. hosei; it possibly 



THE GENUS ONCOPUS 171 

belongs to an undescribed species. 

INDONESIA, Kalimantan, East Kalimantan, from a degraded rainforest near Sepaku 
[Fig. 1 (28)], 40 km NNW of Balikpapan, 1 juvenile (MAR), leg. C. Deeleman, 2.-5.HI.1980. 
This specimen is similar to juveniles of O. doriae, but it possesses no subdistal ventral process 
on the palpal femur. 

INDONESIA, Kalimantan, East Kalimantan, Samarinda District, Sangasanga Muara 
[Fig. 1 (27)], 4 juveniles (ZMT), leg. P. Lehtinen. These specimens appear to be conspecific with 
the previous one. 

INDONESIA, Sumatra, West Sumatra Province, off the road Pematangsiantar - Prapat, 
18 km from Prapat [Fig. 1 (16)], 980 m, 1 juvenile (MHNG), leg. B. Hauser, 8.XI.1985. This is 
the only known record of Oncopus from Sumatra Island. In the course of that journey (Sum-85) 
B. Hauser also collected in Singapore, but a confusion of localities can be ruled out. This fairly 
large specimen from Sumatra is distinct from all other Oncopus juveniles examined in that it 
possesses a light median longitudinal stripe on the anterior part of the dorsal scutum, running 
from the carapace front margin to opisthosomal area III. No such pattern is present in the O. trun- 
catus juveniles examined from Singapore. Some small juveniles of O. doriae (not all of them 
and not the large juveniles) also possess a median longitudinal stripe over the carapace region, 
but it is less pronounced, narrower and not extending onto the anterior opisthosomal areas. 

DISCUSSION 

Taxonomic characters and relationships 

Bristowe (1975) was so impressed by the external morphology of the O. feae 9 
he found on Penang Island that he proposed a new suborder, Oncopodines, for the 
family Oncopodidae. This was obviously done in ignorance of the fact that the same 
one-sided interpretation (not taking genitalia into consideration) of oncopodid mor- 
phology has earlier led Silhavy (1961) to establish the suborder Oncopodomorphi. On 
the basis of plesiomorphic similarities in external morphology with the Cypho- 
phthalmi, both authors considered the Oncopodidae as a primitive group at the base of 
the opilionid system. It was not until genital morphology in opilionids became better 
known that this view changed considerably. The studies of Martens and collaborators 
(Martens, 1976, 1980, 1986; Martens et al, 1981) showed that oncopodids are actually 
a highly evolved branch within the Laniatores. This was corroborated by an analysis of 
morphological and molecular data by Giribet et al. (2002) and by our investigations on 
the penis morphology of all known oncopodid species (Schwendinger, 1992; Martens 
& Schwendinger, 1998; Schwendinger & Martens, 1999, 2002a, b and this paper). 

Oncopus species possess several characters which seem to be apomorphic 
within the family, i.e. large body size, massive chelicerae with a ventral process on mo- 
bile finger, reduced "carapace-abdomen bridge", reduced number of leg tarsalia (1- 
1-1-1), scale-like anteroproximal apophysis on ventral side of leg coxa II, white 
enclosures in keeled elevations of ventral scutum in â 6 , terminal setae on truncus 
penis, and proximad-directed glans penis. Therefore we consider Oncopus as a taxon 
that occupies an advanced position in oncopodid phylogeny, either as the sister group 
to Gnomulus or of Caenoncopus (see Schwendinger & Martens, 2002b: fig. 59). 

Four species groups are here distinguished. The feae-group (O. feae, O. ma- 
layanus sp. n.) and the truncatus-group (O. truncatus, O. tiomanensis sp. n., O. lingga 
sp. n.) from the Malay Peninsula and nearby islands are closely related to each other, 
and both probably have a more distant relationship to O. doriae (do ria e -group) from 



172 P. J- SCHWENDINGER & J. MARTENS 

northwestern Borneo. The hosei-group (O. hosei, O. megachelis, O. expatriatus sp. n.) 
appears to be further apart from the other three species groups and sister to them. 

Most species are clearly distinct in their penis morphology. Intraspecific 
variation in genital morphology of O. truncatus is considerable, and the penes of this 
species are only little different from those of O. tiomanensis sp. n. In external morpho- 
logy, however, these species are clearly distinguishable and therefore we treat them as 
separate species. 

We want to draw attention to the light enclosures embedded in the transversal 
keels of the ventral scutum in S S of some Oncopus species, which are not found in 
other oncopodids. These cuticular structures possibly have a similar function as the 
glandular setae of 6 6 in some Gnomulus species (see Schwendinger & Martens, 
1999: fig. 69a, b) and they deserve histological investigation. 

Cheliceral teeth and palpal claws are strongly worn in old specimens (mature 
Oncopus are presumably long-lived), which indicates that chelicerae and palps are 
used for digging and for handling obstacles. The shape of the cutting edges of the 
cheliceral fingers is therefore only useful as a taxonomic character in "young" mature 
specimens. 

Zoogeography 

The genus Oncopus occurs in Peninsular Malaysia, on Singapore Island, 
Sumatra and its islands to the east, and on Borneo. It obviously is restricted to the 
perhumid zone of the SE-Asian tropics and occurs roughly in between 6°N and 4°S. 
The two records from Thailand (one of them from near Bangkok and the other from an 
unspecified locality), based on 2 specimens from the collection of Roewer, are 
extremely dubious. No additional Oncopus specimens have since been collected in this 
country despite intensive leaf litter sifting in southern Thailand by various persons 
during recent years. Furthermore, the seasonally dry climate of central Thailand 
presumably does not meet the ecological requirements of this genus. We would, 
however, not be surprised if Oncopus is one day discovered in the most southern 
provinces of Thailand. 

In contrast to the species-rich genus Gnomulus, syntopic occurrences of two or 
more Oncopus species are not known and probably do not exist. Previous records of 
sympatric Oncopus species on relatively small islands (O. feae and "O. alticeps" on 
Penang and O. truncatus and "O. acantïiochelis" on Singapore) were long considered 
dubious, especially since the type of each "sympatric species" either belongs to the 
opposite sex or to a different postembryonic instar. 

Ranges of distribution in Oncopus species seem to be fairly small (with the 
exception of O. malayanus sp. n.) and therefore we expect that several unknown 
species remain to be discovered; most likely so on Sumatra and in the southern part of 
Borneo from where juvenile specimens are already known. 

ACKNOWLEDGEMENTS 

We gratefully acknowledge loans and donations of specimens from the follow- 
ing colleagues: Giulio Cuccodoro and Ivan Lobi (MHNG), Daniel Burckhardt (Natural 
History Museum, Basel), Christa Deeleman-Reinhold (Ossendrecht), Giuliano Doria 



THE GENUS ONCOPUS 173 

(MSNG), Gonzalo Giribet (MCZ), Manfred Grasshoff (SMF), Bernd Hauser (MHNG), 
Siegfried Huber (Oberuhldingen), Sergei Kurbatov (Moscow State University), Pekka 
Lehtinen (ZMT), Janet Beccaloni and Paul Hillyard (NHML), Lionel Monod 
(MHNG), Alexander Riedel (Staatliches Museum für Naturkunde, Karlsruhe), Andreas 
Schulz (Leichlingen), Chang Man Yang (ZRC). S. Huber also provided information on 
the behaviour of O. truncatus; Peter Jäger (SMF) searched the documents of Roewer 
for collecting data on oncopodids. Christine Lehmann-Graber (Geneva) skillfully pro- 
duced the whole-animal illustrations of O. malayanus sp. n., John Hollier (MHNG) 
checked the English text and Jürgen Gruber (Naturhistorisches Museum Wien) kindly 
reviewed the manuscript. The German Academic Exchange Service (Deutscher 
Akademischer Austauschdienst, Bonn) supported P.J. S. with a 1-month research grant 
at the beginning of this project. The Feldbausch Foundation of the University of Mainz 
provided travel funds to J.M. and the MHNG to P.J.S. 

REFERENCES 

Beccari, O. 1904. Wanderings in the great forests of Borneo. Archibald Constable & Co. Ltd., 
London, 456 pp. 

Bristowe, W. S. 1976. Rare arachnids from Malaysia and Sumatra. Journal of Zoology 178: 
7-14. 

Giribet, G., Edgecombe, G. D., Weeler, W. C. & Babbitt, C. 2002. Phylogeny and systematic 
position of Opiliones: a combined analysis of chelicerate relationships using morpho- 
logical and molecular data. Cladistics 18: 5-70. 

Helversen, O. von & Martens, J. 1972. Unrichtige Fundort-Angaben in der Arachniden- 
Sammlung Roewer. Senckenbergiana biologica 53 (1/2): 109-123. 

Loman, J. C. C. 1902. Neue aussereuropäische Opilioniden. Zoologische Jahrbücher, Abteilung 
für Systematik 16: 163-216. 

Martens, J. 1976. Genitalmorphologie, System und Phylogenie der Weberknechte (Arachnida: 
Opiliones). Entomologica Germanica 3: 51-68. 

Martens, J. 1980. Versuch eines phylogenetischen Systems der Opiliones (pp. 355-360). In: 
Gruber, J. (ed.). Verhandlungen, 8. Internationaler Arachnologen-Kongreß. Egermann, 
Wien, 506 pp. 

Martens, J. 1986. Die Großgliederung der Opiliones und die Evolution der Ordnung (Arach- 
nida) (pp. 289-310). In: Barrtentos, J. A. (ed.). Actas 10 Congreso Internacional de 
Aracnologia, Jaca/Espana, Septiembre 1986, vol. 1. Instituto Pirenaico de Ecologia & 
Grupo de Aracnologia, Barcelona, X+428 pp. 

Martens, J., Hoheisel, U. & Götze, M. 1981. Vergleichende Anatomie der Legeröhren der 
Opiliones als Beitrag zur Phylogenie der Ordnung (Arachnida). Zoologische Jahrbücher, 
Abteilung für Anatomie 105: 13-76. 

Martens, J. & Schwendinger, P. 1998. Ataxonomic revision of the family Oncopodidae I. New 
genera and new species of Gnomulus Thorell (Opiliones, Laniatores). Revue suisse de 
Zoologie 105 (3): 499-555. 

Moritz, M. 1993. 1. Unterstamm Arachnata (pp. 64-442). In: Grüner, H.-E. (ed.). Lehrbuch der 
speziellen Zoologie, Band I: Wirbellose Tiere. 4. Teil: Arthropoda (ohne Insecta) (4 th 
edition). Gustav Fischer Verlag, Jena, 1279 pp. 

Murphy, F. & Murphy, J. 2000. An introduction to the spiders of Southeast Asia. Malaysian 
Nature Society, Kuala Lumpur, 625 pp. + 32 colour plates. 

Roewer, C. F. 1915. 106 neue Opilioniden. Archiv für Naturgeschichte 81 (3): 1-152. 

Roewer, C. F. 1923. Die Weberknechte der Erde. Gustav Fischer Verlag, Jena, 1116 pp. 

Roewer, C. F. 1927. Weitere Weberknechte I. Abhandlungen herausgegeben vom naturwis- 
senschaftlichen Verein zu Bremen 26 (2): 261-402. 



174 P. J- SCHWENDINGER & J. MARTENS 



Pocock, R. I. 1897. Descriptions of some new Oriental Opiliones recently received by the 

British Museum. Annals and magazine of natural history (6) 19: 283-310. 
Schwendinger, P. J. 1992. New Oncopodidae (Opiliones, Laniatores) from Southeast Asia. 

Revue suisse de Zoologie 99 (1): 177-199. 
Schwendinger, P. J. & Martens, J. 1999. A taxonomic revision of the family Oncopodidae II. 

The genus Gnomulus Thorell (Opiliones, Laniatores). Revue suisse de Zoologie 106 (4): 

945-982. 
Schwendinger, P. J. & Martens, J. 2002a. A taxonomic revision of the family Oncopodidae III. 

Further new species of Gnomulus Thorell (Opiliones, Laniatores). Revue suisse de 

Zoologie 109(1): 47-113. 
Schwendinger, P. J. & Martens, J. 2002b. Penis morphology in Oncopodidae (Opiliones, 

Laniatores): evolutionary trends and relationships. Journal ofArachnology 30: 425-434. 
Silhavy, V. 1961. Die Grundsätze der modernen Weberknechttaxonomie und Revision des bis- 
herigen Systems der Opilioniden (pp. 262-267). In: Strouhal, H. & Beier, M. (eds). 

Verhandlungen des 11. Internationalen Kongresses für Entomologie in Wien (17.-25. 

August 1960), vol. 1. C. Reisser's Söhne, Wien, XLIV + 802 pp. 
Silhavy, V. 1974. Some phalangids from Ceylon and Malaysia. Revue suisse de Zoologie 81 (1): 

25-28. 
S0RENSEN, W. 1932. Descriptiones Laniatorum (Arachnidorum Opilionum subordinis) fecit 

William S0rensen opus posthumum recognovit et edidit Kai L. Hendriksen. Mémoires de 

l'Académie Royale des Sciences et des Lettres de Danemark, Copenhague (Section des 

Sciences) (9) 3 (4): 199-422. 
Suzuki, S. 1976. Report on a collection of opilionids from Pasoh Forest Reserve, West Malaysia. 

Nature and Life in Southeast Asia 7: 9-38. 
Suzuki, S. 1983. Additional notes on the Malaysian harvestmen (Arachnida: Opiliones). Acta 

arachnologica 32: 1-4. 
Thorell, T. 1876. Descrizione di alcune specie di Opilioni dell 'Archipelago Malese apparte- 
nenti al Museo Civico di Genova. Annali del Museo civico di Storia Naturale di Genova 

(1)9: 111-138. 
Thorell, T. 1890. Aracnidi di Pinang raccolti nel 1889 dai signori L. Loria e L. Fea. Annali del 

Museo civico di Storia Naturale di Genova (2) 10: 269-383. 
Thorell, T 1891. Opilioni nuovi o poco conosciuti dell'Arcipelago Malese. Annali del Museo 

civico di Storia Naturale di Genova (2) 10: 669-770, plates Vili + LX. 
Whitmore, T. C. 1991. An introduction to tropical rain forests. Clarendon Press, Oxford, 

226 pp. 



Revue suisse de Zoologie 111 (1): 175-181; mars 2004 



Colasidia wau, a new leleupidiine species from Papua New Guinea 
(Insecta, Coleoptera, Carabidae, Zuphiinae) 

Martin BAEHR 

Zoologische Staatssammlung, Münchhausenstr. 21, D-81247 München, Germany. 

E-mail: martin.baehr@zsm.mwn.de 



Colasidia wau, a new leleupidiine species from Papua New Guinea 
(Insecta, Coleoptera, Carabidae, Zuphiinae). - A new species of the 
leleupidiine genus Colasidia Basilewsky is described from Papua New 
Guinea: C. wau sp. n. The new species is most similar to C. papua Dar- 
lington, differing from it by even smaller eyes, narrower pronotum, and 
longer elytra. A key to all Australian-New Guinean species of Colasidia is 
provided. 

Key- words: Coleoptera - Carabidae - Zuphiinae - Leleupidiini - Colasidia 
- new species - Papua New Guinea. 

INTRODUCTION 

Within a sample of specimens collected in central Papua New Guinea in 1992 
by G. Cuccodoro of Geneva Museum, a single specimen of the genus Colasidia of the 
zuphiine tribe Leleupidiini was detected that is different from all described New 
Guinean species and, although it reminds C. papua Darlington in general shape and 
structure, it differs considerably from that species in body shape and size of eyes, and 
therefore is described as a new species. While working on the present paper, R. T Bell 
(Burlington) sent me another Colasidia specimen for determination, collected by him 
in the same area in 1982, that turned out to belong to the same new species. As both 
specimens are males, the species identity is doubtless, although even in external struc- 
tures the new species is different from all other known New Guinean Colasidia. 

This paper is a supplement to my monograph of the Oriental-Australian species 
of the genus Colasidia (Baehr, 1997). 

MATERIAL AND METHODS 

Two specimens of the new species are available that were collected by different 
collectors but approximately at the same locality and at same altitude. The holotype 
was kindly presented by the collector to Zoologische Staatssammlung, München, and 
is presently located in the working collection of the author (CBM-ZSM). The paratype 
is in Muséum d'histoire naturelle, Genève (MHNG). 



Manuscript accepted 25.05.2003 



176 M. BAEHR 

For dissection of the male genitalia the specimens were soaked in a wet jar for 
one night, the genitalia were then cleaned for a short while in hot 4% KOH. For the 
description normal taxonomic methods were used. The description follows the style of 
my synoptic paper of the Oriental-Australian Colasidia (Baehr, 1997). 

The habitus photograph was obtained using SPOT Advanced, version for 
Windows 3.5, and subsequently was worked using MS Corel Photo Paint 10. 

Measurements were taken using a stereo microscope with an ocular micrometer. 
Length has been measured from apex of labrum to apex of elytra. Length of pronotum 
was measured along midline, width of base of pronotum at the extreme tips of the basal 
angles. Length of head was taken from apex of labrum to anterior border of "neck", 
length of orbit was likewise measured to anterior margin of "neck". 

Colasidia Basilewsky, 1954 

For information about taxonomy and distribution of the genus Colasidia see 
Baehr (1997: 613). From New Guinea four species were recorded so far: C. madang 
Darlington, C. papua Darlington, C. kokodae Baehr, and C. garainae Baehr (Dar- 
lington, 1971; Baehr, 1991, 1997, 2000a). Surprisingly, no Colasidia has been recorded 
from Irian Jaya so far, and even the records from Papua New Guinea are few and from 
a rather restricted range in the central part. Probably, this is mainly due to absolutely 
inadequate sampling of these litter- or even soil inhabiting tiny flightless beetles that 
are probably best sampled by sieving or Berlese extraction of litter and soil. Therefore, 
it is not too surprising, that one specimen of the new species was collected by staff of 
the Geneva Museum who is well known for its very successful sampling of litter and 
soil inhabiting tiny insects. 

Almost all Colasidia from New Guinea (and Australia) are known from single 
specimens only which further demonstrates the rarity of these beetles and/or the little 
collecting efforts that were made. I suspect, however, that in New Guinea at least, dis- 
tribution of these beetles is rather scattered, and that it would need enormous efforts to 
get a better picture of species inventory and distribution. 

Colasidia wau sp. n. Figs 1, 2 

Type material: Holotype: 6, PAPUA NEW GUINEA Wau-Mt. Misim March 15, 1982 
Coll. R. t. Bell / 1400 m /under stone / Colasidia nr. madang (CBM-ZSM). 

Paratype: 1 6 , PAPUA NG: Morobe above Wau 1450m, 21. V. 1992 G. Cuccodoro #5b 

(MHNG) 

Diagnosis: Medium sized, fairly elongate, piceous species, characterized by 
elongate, posteriorly markedly rounded head with fairly small eyes, rather elongate 
elytra with convex apical margin, and rather coarse and sparse, regularly arranged 
puncturation and pilosity; further distinguished from related species by the short, com- 
pact aedeagus having a moderately elongate, slightly upturned apex, and by presence 
of an elongate, folded sclerite in the internal sac. 

Description 

Measurements: Length: 4.75-4.80 mm; width: 1.60-1.62 mm. Ratios. Length/ 
width of head: 1.64-1.65; length orbit/eye: 3.9-4.1; length/width of pronotum: 1.16- 



COLASIDIA WAU, A NEW LELEUPIDIINE SPECIES 



177 




D 




Fig. 1 



Colasidia wau sp. n. Male genitalia: A) aedeagus (left side), B) shape of apex (from below), C) 
left and D) right parameres, E) genital ring. Scales: 0.25 mm. 



1.22; width widest part/base of pronotum: 1.54-1.61; width pronotum/head: 1.21-1.30; 
length/width of elytra: 1.54-1.55; width elytra/pronotum: 1.80-1.88. 

Colour: More or less dark piceous, suture of elytra very narrowly reddish. 
Labrum, palpi, legs, and antennae somewhat lighter, dark yellowish to light brown. 

Head: Rather elongate, very slightly widened behind eyes, widest far away 
from base, orbits posteriorly very widely rounded. Clypeus and frons in middle slightly 
raised, frons not grooved. Eyes small, laterally barely projecting, length only about 1/4 
of orbit length. Surface above and behind eye with a narrow, deep groove. Clypeus 
anteriorly faintly concave, lateral angles (above base of antenna) barely projecting. 
Clypeal suture laterally with shallow grooves. Labrum anteriorly rather excised, 6- 
setose, inner 4 setae slightly shorter than outer ones, lateral margin densely pilose. 
Mandibles short. Mentum with triangular, at apex faintly excised tooth. Labium trun- 
cate. Maxillary palpus moderately elongate, apex obtusely rounded. Terminal segment 
of labial palpus large and very elongate. Antenna very short, barely attaining anterior 
third of pronotum. Median antennomeres >1.5 x as wide as long, 3 rd antennomere c. 
2/3 as long as 1 st , c. 1.5 x as long as 2 nd antennomere. Surface without microreticu- 
lation, highly glossy. Puncturation rather coarse, moderately dense, diameter of punc- 
tures slightly wider than distance between punctures. Pilosity moderately dense, rather 
elongate, moderately erect, inclined anteriorly. Both supraorbital setae elongate, fairly 
well distinguished from pilosity, posterior supraorbital setae situated far behind eye. 

Pronotum: Rather elongate, cordiform, anteriorly considerably widened, 
slightly longer than wide, considerably wider than head, widest in anterior third. Upper 
surface rather convex, faintly sulcate along median line. Lateral margin in anterior two 
thirds strongly convex, in front of posterior angles deeply but somewhat irregularly 
sinuate, basal third almost straight. Apex wide, faintly excised, anterior angles convex, 



178 M. BAEHR 




Fig. 2 
Colasidia wau sp. n. Paratype, habitus. Length: 4.8 mm. 



barely projecting. Base rather narrow, laterally angulately excised, basal angles pro- 
jecting as sharp, small denticles. Lateral margin slightly raised, with distinct border 
line, but without distinct marginal channel. Median line distinct, though but faintly 
impressed. Prebasal grooves deep. Both marginal setae very elongate, anterior seta 
situated slightly behind anterior fourth of pronotum, posterior seta situated right on 
basal angle. Surface without microreticulation, highly glossy, with moderately dense, 
very coarse puncturation. Diameter of punctures wider than distance between them. 
Pilosity fairly dense, elongate, hirsute, irregularly inclined, rather erect. 

Elytra: Rather elongate and but moderately wide, laterally evenly but gently 
curved, widest slightly behind middle, upper surface moderately convex. Intervals not 
raised. Humeri wide, rather projecting, rounded off. Apex wide, gently convex, slightly 
redressed to suture. Striae rather regularly marked by rows of punctures, puncturation 
moderately sparse, very coarse, rather regularly arranged to longitudinal rows. Dia- 
meter of punctures considerably wider than distance between them. Fixed setae in third 
interval very difficult to recognize within the coarse puncturation. Series of marginal 
pores extremely difficult to detect when setae broken, apparently consisting of 8 basal, 
3 postmedian, 6 apical pores, and 1 pore at apex of 3 rd stria. Setae very elongate. 
Surface without microreticulation, highly glossy. Pilosity rather sparse, elongate, 
hirsute, rather regular, inclined posteriorly, rather depressed. 



COLASIDIA WAU, A NEW LELEUPIDIINE SPECIES 179 

Male genitalia: Genital ring fairly narrow, irregularly triangular and rather 
asymmetric, basal part short, apical plate very wide, oblique. Aedeagus rather short, 
with moderately elongate, rather narrow, slightly upturned apex. Lower surface very 
gently bisinuate. Internal sac in middle with an elongate, folded sclerite. Both para- 
meres fairly short, of very different size and shape, left paramere rather parallel, with 
widely rounded apex. 

Female genitalia: Unknown. 

Variation: Apart from slightly wider pronotum in the paratype, very little 
variation noted. 

Etymology: The name refers to the locality where the new species was detected. 

Distribution: Central eastern Papua New Guinea. Known only from the imme- 
diate vicinity of Wau. 

Collecting circumstances: Holotype collected "under stone" at 1400 m, para- 
type sampled by sieving and Winkler extraction of rotting trunk and vegetational debris 
in mountain rain forest, at 1450 m. Like the other New Guinean species of the genus 
Colasidia, this is a ground living beetle of the rain forest litter that occurs at median 
altitude in the lower level of the montane rain forest zone. 

Key to species of Colasidia of New Guinea and Australia 

To facilitate identification of the new species the most recent key to the record- 
ed Australian-New Guinean species of Colasidia (Baehr, 2000a) is revised here. For 
the benefit of the user some figures from previous papers on the subject are mentioned 
in this key: B00: Baehr, 2000a; B87: Baehr, 1987; B91: Baehr, 1991. 

1 . Head about parallel, or wider across eyes than across orbits; base of head 
usually considerably rounded (doubtful specimens under both couplets) .... 2 
Head decidedly wider at posterior angles or across orbits than across 
eyes; base of head less rounded, more square 4 

2. Head short, eyes very large, ratio of distance from orbit to neck/eye 
length <1.5; basal angles of head very widely rounded (B91 fig. 6); 
puncturation of elytra irregular, rather confused; odd intervals raised in 
anterior half; aedeagus unknown. Kokoda, central Papua Peninsula, 

eastern Papua New Guinea kokodae Baehr 

Head longer, eyes smaller, ratio of distance from orbit to neck/eye length 

>3 ; basal angles of head less widely rounded; puncturation of elytra in 
regular rows; intervals not markedly raised 3 

3. Eyes larger, ratio of distance from orbit to neck/eye length c. 3; pro- 
thorax shorter, ratio length/width c. 1.10; prothorax less wide in 
comparison to head, ratio width of prothorax/width of head c. 1.16; 
elytra shorter and wider, ratio length/width c. 1.50; aedeagus unknown. 
Dobodura, central Papua Peninsula, eastern Papua New Guinea 

papua Darlington 

Eyes smaller, ratio of distance from orbit to neck/eye length c. 4; pro- 
thorax longer, ratio length/width >1.16; prothorax wider in comparison 



180 M. BAEHR 

to head, ratio width of prothorax/width of head >1.20; elytra longer and 
narrower, ratio length/width >1.54; aedeagus short, rather compact, with 
moderately elongate, faintly upturned apex (Fig. 1). Vicinity of Wau, 
eastern central Papua New Guinea wau sp. n. 

4. Eyes very small, ratio of distance from orbit to neck/eye length >5; head 
very elongate, markedly triangular (B87 fig. 1). Northeastern Queens- 
land, Australia monteithi Baehr 

Eyes larger, ratio of distance from orbit to neck/eye length <4.5; head 
shorter, usually less markedly triangular. New Guinea 5 

5. Pronotum shorter and wider, ratio length/width c. 1.05; elytra shorter 
and wider, less depressed, ratio width of elytra/width of prothorax >2.05, 
ratio length/width of elytra c. 1.40; aedeagus unknown. Damanti, Huon 

Peninsula, northern Papua New Guinea madang Darlington 

Pronotum longer and narrower, ratio length/width >1.12; elytra longer 

and narrower, more depressed, ratio width of elytra/width of prothorax 
<1.90, ratio length/width of elytra >1.48; aedeagus see fig. 1, BOO fig. 1. 
Distribution different 6 

6. Smaller species, length <4.35 mm; head shorter and wider, ratio 
length/width <1.5; elytra shorter and wider, ratio length/width <1.51; 
aedeagus elongate, with elongate, at tip very slightly upturned apex (BOO 
fig. 1). Garaina, northwestern Papua Peninsula, eastern Papua New 

Guinea garainae Baehr 

Larger species, length >4.75 mm; head longer and narrower, ratio 
length/width >1.64; elytra longer and narrower, ratio length/width 
>1.54; aedeagus short, compact, with moderately elongate apex (Fig. 1). 
Vicinity of Wau, eastern central Papua New Guinea wau sp. n. 

Remarks 

Even when sampling of these tiny, litter inhabiting beetles is difficult and prob- 
ably has been conducted quite inadequately in New Guinea so far, the absolute lack of 
any record from the western half of this island (Irian Jaya) is still enigmatic, because 
collectors like A. Riedel who conducted extensive sieving and Berlese extraction 
samplings in Irian Jaya, failed to find any specimens (see Baehr, 2000a). Therefore, it 
is uncertain, whether the western part of New Guinea represents one of the quite sur- 
prising distribution gaps within the extended range of Leleupidiini in the Oriental- 
Australian region, and whether Leleupidiini still escaped the attention of collectors in 
this region. 

When the leleupidiine fauna of this large island will be better explored, this 
question will be asked again, in combination with the questions, why not a single 
Colasidia has been ever collected in Java, the Lesser Sunda Islands, Sulawesi, and the 
Moluccas, whereas they seem to be quite speciose in Malaysia, Sumatra, (northern) 
Borneo, and Papua New Guinea, and one species even occurs in northeastern Australia. 
Then, probably the very complex paleogeographic history of New Guinea should be 
considered for explanation of the curious distribution pattern of the tribe on this large 
island. 



COLASIDIA WAU, A NEW LELEUPIDIINE SPECIES 181 

At present, we only can state that a rather limited area in central eastern Papua 
New Guinea is the single place in New Guinea where Leleupidiini have been recorded 
so far, and moreover, although being rare in terms of specimens, they are nevertheless 
quite speciose there. As in other regions where Leleupidiini occur, the species 
apparently occupy extremely restricted ranges, which is not surprising in view of their 
inability for flight and the rugged montane landscape of this island, where the ranges 
of neighbouring species of Leleupidiini usually are separated by deep valleys. As a 
consequence, the number of species still to be detected in New Guinea likely might be 
considerably augmented in future, in the same manner, as it has been demonstrated for 
other flightless, montane New Guinean carabid beetles (Baehr, 1995, 1999, 2000b, 
2001a, 2001b). 

ACKNOWLEDGEMENTS 

My sincere thanks are due to Drs I. Lobi and G. Cuccodoro of the Geneva 
Museum and Prof. Dr R. T. Bell of University of Vermont (Burlington) for the kind 
loan, respectively gift of the specimens. 

REFERENCES 

Baehr, M. 1987. Revision of the Australian Zuphiinae 2. Colasidia monteithi sp. nov. from 

North Queensland, first record of the tribe Leleupidiini in Australia (Insecta: Coleoptera: 

Carabidae). Memoirs of the Queensland Museum 25: 135-140. 
Baehr, M. 1991. On new and rare Leleupidiini from the Oriental and Australian Regions 

(Coleoptera, Carabidae, Zuphiinae). Mitteilungen der Münchner Entomologischen 

Gesellschaft 81: 193-202. 
Baehr, M. 1995. The genus Mecyclothorax Sharp in New Guinea (Insecta, Coleoptera, 

Carabidae, Psydrinae). Mitteilungen der Münchner Entomologischen Gesellschaft 85: 

3-19. 
Baehr, M. 1997. Leleupidiini from the Oriental Region. 1. New species of the genus Colasidia 

Basilewsky (Insecta, Coleoptera, Carabidae, Zuphiinae). Revue suisse de Zoologie 104: 

611-659. 
Baehr, M. 1999. The genus Dolichoctis Schmidt-Göbel in New Guinea (Insecta, Coleoptera, 

Carabidae, Lebiinae). Coleoptera 2: 121-169. 
Baehr, M. 2000a. A new species of the leleupidiine genus Colasidia Basilewsky from New 

Guinea (Insecta, Coleoptera, Carabidae, Zuphiinae). Spixiana 23: 41-45. 
Baehr, M. 2000b. Revision of the genus Idiagonum Darlington from New Guinea (Carabidae, 

Agoninae). Coleoptera 4: 55-92. 
Baehr, M. 2001a. Further new species of the genera Fortagonum Darlington and Collagonwn 

Baehr from New Guinea (Insecta, Coleoptera, Carabidae, Agoninae). Spixiana 24: 

53-72. 
Baehr, M. 2001b. The genus Rhytiferonia Darlington in New Guinea (Carabidae: Pterosti- 

chinae). Coleoptera 5: 41-55. 
Darlington, P. J. Jr 1971. The Carabid beetles of New Guinea. Part IV. General considerations, 

analysis and history of the fauna, taxonomic supplement. Bulletin of the Museum of 

Comparative Zoology 142: 129-337. 



Revue suisse de Zoologie 111 (1): 183-211; mars 2004 



Re vision of the Minettia fasciata species-group 
(Diptera, Lauxaniidae) 

Bernhard MERZ 

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

E-mail: bernhard.merz@mhn.ville-ge.ch 



Revision of the Minettia fasciata species-group (Diptera, Lauxaniidae). 

- The Minettia fasciata species-group is defined. It includes 4 Palaearctic 
species, namely Minettia czernyi Freidberg & Yarom, Minettia fasciata 
(Fallen) [= M. rivosa (Meigen), synonymy confirmed], M. subvittata 
(Loew) [= M. Cataracta (Pandellé), syn. conf.] and M. tabidiventris 
(Rondani) [= M. luteofrontata (Becker), syn. nov; = M. fasciata auctt. nee 
Fallen]. Sapromyza pallida Meigen is treated as a nomen dubium. All 
available types were studied, the 4 species are described, illustrated and 
compared with each other. A key to the species of Minettia with latero- 
ventral black spots on the scutellum is presented. 

Key- words: Diptera - Lauxaniidae - Minettia fasciata species-group - 
Palaearctic region. 

INTRODUCTION 

The Lauxaniidae are a large family of about 1800 described species worldwide. 
They are best represented in the tropics of the Old and New World (except for the 
Afrotropical region), and their diversity declines strongly towards the more temperate 
regions. Fewer than 180 species are recorded from Europe (Merz, 2003a). Most species 
are found in forests, on shrubs, trees, and leaves. They are less common in dry and wet 
grasslands. Despite their often remarkably high population densities and apparant 
importance in decomposing plant material, they are insufficiently known, even in the 
comparatively well studied Western Palaearctic region. 

Minettia Robineau-Desvoidy, with almost worldwide distribution, is one of the 
most speciose genera of Lauxaniidae with somewhat less than 100 species described 
so far. The Palaearctic fauna is the most diverse with 56 described species, although 
the other regions are less intensively studied and may be much richer than is known 
today. The genus was divided into 3 subgenera by Shatalkin (2000). Within Minettia s. 
str. a number of species-groups may be recognized, one of which is called the Minettia 
fasciata group which is restricted to the Palaearctic region. This group also includes 
Minettia nemorosa Robineau-Desvoidy, 1830, which is the type species of Minettia. 
The group is characterized by the grey microtrichose mesonotum, a pair of black spots 



Manuscript accepted 27.08.2003 



184 B. MERZ 

on the lateroventral side of the scutellum (Figs 2-5), 0+3 dorsocentral setae, 6 rows of 
acrostichal setulae, only one dorsal preapical seta on the midtibia, dark transverse 
stripes on the abdominal tergites (Figs 6-7) and characteristic terminalia. 

While studying a large number of specimens with this set of characters I 
realized that the limits of the described species differ from one author to another and 
that nomenclature is far from being resolved. The objective of this paper is to clarify 
the recognized synonymies by studying the types where still available, and to present 
diagnostic characteristics accompanied by illustrations, helping to define the limits of 
each species. 

HISTORICAL REVIEW 

The following account reviews the changing history of the names associated 
with the species of the Minettia fasciata group as briefly diagnosed above and 
extensively described below. This summary is not complete; only major revisions and 
catalogues are included which differ in their interpretations from other papers. 

The first two species which belong to this complex are Lauxania fasciata 
Fallen, 1 826, and Sapromyza rivosa Meigen, 1 826, with Fallen having priority. 

In 1830, Meigen described Lauxania pallida and Robineau-Desvoidy proposed 
in the same year the genus Minettia and described, among others, M. nemorosa which 
was designated as type species of Minettia by Westwood (1840). 

The various interpretations of the different species started with Macquart 
(1835), who synonymized nemorosa with rivosa. The following history is given in an 
abbreviated form. Papers with descriptions of new species relevant for this study are 
marked with an asterix (*): 

Westwood (1840): (l) fasciata (= rivosa, = nemorosa), (2) pallida; designation of 

nemorosa as type species of Minettia. 
*Loew (1847): (l) fasciata (= rivosa), (2) description of Sapromyza subvittata Loew 
Zetterstedt (1847): rivosa (= fasciata) 

Schiner (1863): (l) fasciata (= rivosa, = nemorosa), (2) pallida, (3) subvittata 
*Rondani (1877): (l) fasciata (= rivosa), (2) subvittata, (3) description of Sapromyza 

tabidiventris Rondani 
*Becker (1895): (1) fasciata (= rivosa, = nemorosa), (2) subvittata (= tabidiventris), 

(3) description of Sapromyza luteofrontata Becker (pallida Meigen not 

included) 
*Pandellé (1902): (1) fasciata (= rivosa, = nemorosa), (2) description of Sapromyza 

Cataracta Pandellé (= subvittata auctt, = fasciata var sensu Zetterstedt) 
Hendel (1908): (1) Cataracta, (2) dimidiata Loew (= pallida Meigen, nee pallida 

Fallen), (3) fasciata (= rivosa, = nemorosa), (4) luteofrontata, (5) subvittata (= 

tabidiventris) 
Czerny (1932): (l) fasciata (= rivosa, = nemorosa, = luteofrontata, = subvittata sensu 

Pandellé), (2) pallida Meigen (= dimidiata Loew), (3) subvittata (= Cataracta, 

= ? tabidiventris) 
*Czerny (1937): description of Minettia quadrisetosa Czerny 
Collin (1948): (1) fasciata, (2) rivosa (no synonymy) 



THE MINETTIA FASCIATA SPECIES-GROUP 1 85 

Papp (1984): (l) fasciata (= nemorosa, = subvittata, = luteofrontata, = Cataracta), (2) 

rivosa (= dimidiata). Nomina dubia: (3) pallida, (4) quadrisetosa, (5) tabidi- 

ventris 
Freidberg & Yarom (1990): (1) czernyi (= nom. no v. for quadrisetosa Czerny, nee 

Becker), (2) fasciata, (3) rivosa 
Shatalkin (2000): (1) czernyi, (2) fasciata, (3) pallida, (4) rivosa, (5) subvittata (no 

synonymy) 

It can be concluded from this non-exhaustive review of major papers about 
Lauxaniidae that the concept of each species and its synonymy varied from author to 
author. All papers prior to Czerny (1932) are unreliable as genitalic structures were not 
studied or illustrated, and types of the already described species were not examined. 
Even Czerny's monograph (1932), although illustrating for the first time the male 
terminalia of his Minettia subvittata, is not particularly helpful because a comparison 
with related species is missing. The first careful study of genitalic characters was 
presented by Collin (1948). Unfortunately, he named the two British species without 
the study of the types. The same deficiency characterizes all subsequent papers, 
including those of Papp (1979, 1984) and Shatalkin (1998, 2000). As a consequence of 
this unsatisfactory situation, faunistic papers, checklists and ecological studies 
including these species are difficult to use because it is impossible to know the identity 
of the species mentioned. 

One species which was often treated as member of the M. fasciata group is 
Sapromyza dimidiata (Loew, 1847) (Hendel, 1908; Papp, 1984; see above). A study of 
the male holotype (ZMHB) revealed that this species belongs to the M. lupulina 
species-group, and so it is not further discussed here. 

MATERIAL AND METHODS 

This study is based on about 1150 specimens which are deposited in the 
following collections: 
CAS California Academy of Sciences, San Francisco, USA (P. A. Arnaud, 

B. Fisher) 
CBM private collection B. Merz, Genève, Switzerland 
CCK private collection Christian Kassebeer, Kiel, Germany 
CGB private collection G. Bächli, Dietikon, Switzerland 
CMCT private collection Miguel Carles-Tolrâ, Barcelona, Spain 
CME private collection Martin Ebejer, Cardiff, Wales 
CPG private collection Paul Gatt, Rabat, Malta 

ETHZ Eidgenössische Technische Hochschule, Zürich, Switzerland (A. Müller) 
FNRA Institut National de la Recherche Agronomique, Montpellier, France (M. 

Martinez) 
MHNG Muséum d'histoire naturelle, Genève, Switzerland 
MHNN Muséum d'histoire naturelle, Neuchâtel, Switzerland (J.-P. Haenni) 
NML Natur-Museum, Luzern, Switzerland (L. Rezbanyai-Reser) 
MNHNP Muséum National d'Histoire Naturelle, Paris, France (J. Charbonnel) 



186 B. MERZ 

MZF Museo Zoologico "La Specola", Firenze, Italy (S. Whitman) 

NHMB Naturhistorisches Museum, Basel, Switzerland (D. Burckhardt) 

NHMW Naturhistorisches Museum, Wien, Austria (R. Contreras-Lichtenberg) 

NRS Naturhistorisk Rijksmuseum, Stockholm, Sweden (T. Pape) 

SIZK Schmalhausen Institute of Zoology, Kiev, Ukraine (V. A. Korneyev) 

SMNS Staatliches Museum für Naturkunde, Stuttgart, Germany (H. P. Tschorsnig) 

SMO Silesian Museum, Opava, Czech Republic (J. Rohâcek) 

TAU Tel Aviv University, Israel (A. Freidberg) 

USNM National Museum of Natural History, Washington D. C, USA (S. Gaimari) 

ZMB Zoological Museum Bergen, Norway (L. Greve- Jensen) 

ZMHB Zoologisches Museum, Humboldt Universität, Berlin, Germany (W. Mey, 

H. Wendt) 

ZML Zoological Museum, University Lund, Sweden (R. Danielsson) 

ZMUC Zoological Museum, University Copenhagen, Danmark (R. Meier) 

ZSM Zoologische Staats Sammlung München, Germany (W. Schacht) 

Morphological terms in the descriptions follow chapters 1.1-1.3 in Papp & Darvas 
(2000), but the interpretation of the antennal structures is according to Stuckenberg 
(1999). 

DESCRIPTION OF THE M1NETTIA FASCIATA SPECIES-GROUP 

Diagnosis 

Species of Minettia s. str. with the following combination of characters: 
scutellum lateroventrally with a pair of black spots (Figs 2-5); mesonotum uniformly 
grey or striped brownish-grey, 0+3 dorsocentral setae and 6 rows of acrostichal setulae 
at level of suture; midtibia with only one dorsal preapical seta; abdomen yellow to 
brown, in the middle of tergites 3-5 each with a darker transverse stripe which is 
interrupted medially (Figs 6-7); male pregonite composed of 2 subequal branches (Fig. 
20); female sternite 8 with 2 ear-like projections laterally (Figs 45, 48). 

Description of Male 

Head (Fig. 1) bicoloured: frons and parafacial yellow to brown; fronto-orbital 
plate, ocellar triangle, occiput and postgena grey microtrichose; face partly pale brown, 
partly grey; frons with yellow stripe dorsal of lunule and with dirty brown transverse 
stripe at level of anterior fronto-orbital seta; parafacial with brown spot lateral of 
antennal base; head in profile about 1.35 times as high as wide; gena one quarter as 
high as compound eye; the latter elongate ovoid; frons (including frontal plates) 
slightly wider than length from lunule to posterior margin of posterior ocelli; frons 
anteriorly with black setulae; face longer than antenna, flat with shallow antennal 
grooves and slightly raised at vibrissal corner; occiput straight, not conspicuously 
concave medially above occipital foramen; fronto-facial angle 120-130°; parafacial 
with single row of about 10 short setulae in ventral third; antenna yellow, sometimes 
postpedicel dorsally grey to black infuscated; postpedicel about twice as long as wide, 
apically rounded; arista plumose, longest rays about as long as width of postpedicel; 
proboscis with yellow to black palpus. The following paired, black setae are present: 2 



THE MINETT1A FASCIATA SPECIES-GROUP 187 

reclinate fronto-orbital setae of equal length; 1 strong ocellar seta and few smaller 
setulae posteriorly between ocelli; 1 very long inclinate medial vertical seta; 1 shorter 
lateroclinate lateral vertical seta; 1 slightly inclinate postocellar seta; one row of strong 
postocular setae and a slightly unordered row behind. 

Thorax with grey ground colour, but to a variable extent brown to yellowish- 
brown striped; mesonotum in M. czernyi yellow with lines of acrostichal setulae pale 
grey only; sometimes with blackish stripe medial of line of dorsocentral setae; post- 
pronotum usually mostly yellow; pleuron grey varying to orange with only traces of 
grey microtrichosity, or with a brown stripe on dorsal third of anepisternum only; 
scutellum dorsally grey, border grey or yellow, lateroventrally pale with a pair of 
conspicuous black spots laterally which are not connected with each other between 
apical scutellar setae (Figs 2-5); subscutellum either black or brown, microtrichose; all 
setae and setulae black; 0+3 dorsocentral setae of subequal length; at level of suture 
with 6 rows of acrostichal setulae, the setulae of the two median rows slightly longer; 
1 strong prescutellar acrostichal seta; 1 postpronotal seta; 2 notopleural setae; 1 pre- 
sutural and 1 postsutural intraalar seta; 1 supraalar seta; 2 postalar setae; basal and 
apical scutellar seta; 1 anepisternal seta; 2 katepisternal setae; 1 proepisternal seta. 

Wing entirely hyaline, sometimes slightly tinged with yellow, but without 
pattern; halter and calypters yellow to whitish yellow. 

Legs yellow, but femora on all legs may vary from greyish to black, in particular 
the forefemur; tibiae sometimes slightly darkened towards tip; anteroventral part of 
forefemur and ventral part of midfemur each with a brown to black subapical spot, 
which is sometimes weak; midfemur with a row of strong, outstanding setulae 
anteriorly in apical half; hindfemur with a strong anterodorsal preapical seta; all tibiae 
with 1 dorsal preapical seta; midtibia with 1 ventral apical seta. 

Abdomen brown to yellow, tergites 3-5 each with a brown to black transverse 
stripe in the middle which is broadly interrupted medially, or which may be reduced to 
more or less large lateral spots (Figs 6-7); sternites unmodified. 

Male terminalia (Figs 9-34). Epandrium semicircular; surstylus black setulose, 
well developed, distally with tooth or invagination, and medially without tooth in 
middle; subepandrial plate with 1 long and sometimes 1-3 smaller black setulae (Fig. 
19); hypandrium as in other species of genus; pregoni te 2-branched, both branches of 
almost same length (except for M. czernyi, Fig. 15), apically more or less pointed and 
directed to the right [when seen from inside, Fig. 20]; postgonite forming an aedeagal 
sheath, distal margin straight (Fig. 25) or with 3 tooth like projections (Fig. 14); phalla- 
podeme short to long; ejaculatory apodeme rather complicated, sclerotized medially 
(Fig. 33); aedeagus membranous, distally "spinulose" (Fig. 25). 

Description of Female 

As male, but tergites 3 and 4 of abdomen either unmodified (Fig. 6) or at pos- 
terior margin with 4-8 much thicker, enlarged, black setae medially of tergite 3 (Fig. 7) 
or tergite 4 (in M. czernyi); pleural membrane in some species with a conspicuous glan- 
dular area on level of segments 4-5 (Fig. 8). 

Female terminalia (Figs 35-48) forming retractible pseudoovipositor; tergite 8 
a well sclerotized plate (Fig. 40), or as two small "L-"shaped weak sclerites (Fig. 44); 



188 



B. MERZ 




Figs 1-5 
Minettia fasciata (Fallen) (1), M. czernyi Freidberg & Yarom (2, 4) and M. subvittata (Loew) 
(3, 5): 1, head; 2-3, scutellum, posterior view; 4-5, scutellum, lateral view. 



tergite 9 evenly curved distally, with (Fig. 40) or without (Fig. 44) soft, long setae 
apically; sternite 8 rather complex, with paired ear-like, heavily sclerotized, invaginat- 
ed projections laterally (Figs 37, 41); sternite 9 and cerei as usual in genus; 3 (2+1) 
spermathecae present, with smooth surfaces, paired spermathecae slightly smaller 
(Fig. 42). 



Variability 

Colouration may vary considerably. The lateroventral black spots on the scutel- 
lum may occasionally be minute or even absent. The transverse dark stripes on tergites 
3-5 may be indistinct or reduced to small dots laterally. The distribution of yellow and 
grey stripes on the mesonotum may vary within each species. Although the yellow pal- 
pus is a good characteristic for quick separation of M. subvittata from the other species 
of the group, it is sometimes almost entirely yellow also in M. fasciata or M. tabidi- 
ventris. In contrast, the terminarla are very characteristic (except for the distal margin 
of the surstylus in M. tabidiventris, see below), and its study is therefore recommend- 
ed for a safe identification. 



THE MINETTIA FASCIATA SPECIES-GROUP 



189 



I« 1 

»ft. 



ifwr 








Figs 6-8 
Abdominal structures of Minettia fasciata species-group. 6, dorsal view of abdomen of holotype 
female of Minettia fasciata (Fallen); 7, dorsal view of abdomen of lectotype female of M. sub- 
vittata (Loew); 8, lateral view of last abdominal segments of female of M. fasciata (Fallen). 



Remarks 

Palaearctic species of Minettia s. lat. have recently been divided into 3 genera: 
Frendelia Collin, Plesiominettia Shatalkin and Minettia s. str. (Shatalkin, 2000). No 
attempt has been made to classify European species of Minettia s. str. The only group 
which has been characterized is the M. biseriata species-group (Papp, 1981). This 
group shares with the M. fasciata species-group the shape of the head, the pectinate 
arista, 0+3 dorsocentral setae, and 6 rows of acrostichal setulae, but differs by the 
entirely yellow thorax, the mostly yellow abdomen (only last 1-4 tergites in some 
species with small black spots laterally), the lateroventrally uniform yellow scutellum 
(except for M. cypriota with a pair of black spots; the position of this species is still 
unresolved and needs further study) and details of the male terminarla with the 2 
branches of the pregonite usually of very different size. Other species with a latero- 
ventral pair of black spots on scutellum have either 1+3 dorsocentral setae and/or the 
abdomen entirely yellow without brown stripes contrasting strongly to the uniform 
grey mesonotum (M. desmometopa (de Meijere), M. dimidiata, M. flaviventris (Costa), 
M. longiseta (Loew), M. lupulina, M. tubifer (Meigen)), see also key below. Other 



190 B. MERZ 

species of Minettia s. str. have the scutellum lateroventrally uniformly yellow, grey or 
black and may therefore be easily separated from species of the M. fasciata species- 
group. 

The present diagnosis of the M. fasciata species-group includes 4 species: M. 
czernyi, M. fasciata, M. subvittata and M. tabidiventris. M. czernyi is placed in the 
group tentatively because it differs in the structure of terminalia of both sexes from the 
other 3 species. Pending a thorough study of the terminalia of all Minettia s. str. I prefer 
to keep it here because of the external similarities with some pale specimens of M. sub- 
vittata with which it may be confused. 

Biology and Distribution 

Nothing is known about immature stages of the 4 species of the M. fasciata 
species-group (Miller, 1977; Smith, 1989). Adults may be found in various biotopes. 
They may be observed abundantly on leaves in open deciduous forests, on shrubs and 
hedges and on isolated trees. As an exception for European Lauxaniidae they are some- 
times common in meadows, grasslands and even crop fields (Reddersen, 1994) where 
they feed on fungi (Reddersen, 1995). The species-group has its apparent center of 
distribution in the Mediterranean region where all 4 species may be found. The diver- 
sity declines north of the Alps and the Carpathian ridge with only 2 species penetrating 
as far north as Scandinavia. Surprisingly, one species, M. fasciata, occurs also in the 
Western parts of the Nearctic region. 

KEY TO THE PALAEARCTIC SPECIES OF MINETTIA S. STR. WITH A PAIR OF 
LATERO VENTRAL BLACK SPOTS ON SCUTELLUM 

1 Scutellum dorsally and lateroventrally of same colour: yellow, grey or 

black other species of Minettia 

1 * Scutellum lateroventrally either with a pair of black spots or entirely 

blackish on ventral side, contrasting to the grey or yellow dorsal surface 

(Figs 2-5) 2 

2 Scutellum lateroventrally entirely blackish, including the area between 

apical scutellar setae Minettia lupulina species-group 

(4 species; Merz, in prep.) 

2* Scutellum lateroventrally with a pair of black spots which are not fused between 
apical scutellar setae (Figs 2-5) 3 

3 1+3 dorsocentral setae present 4 

3* 0+3 dorsocentral setae present, presuturai dorsocentral seta absent 5 

4 Midtibia with 2 dorsal preapical setae; acrostichal setulae in 6 rows; 
palpus, postpronotum, dorsal surface of scutellum, and femora mainly 
yellow M. tubifer (Meigen, 1826) 

4* Midtibia with 1 dorsal preapical seta; acrostichal setulae in 4 rows; 
palpus, postpronotum, dorsal surface of scutellum, and femora mainly 
black M. flaviventris (Costa, 1844) part 



THE MINETT1A FASCIATA SPECIES-GROUP 191 

5 Acrostichal setulae in 4 rows; mesonotum entirely grey including post- 
pronotum; a pair of blackish longitudinal stripes near or along line of 
dorsocentral setae often present; abdomen yellow without transverse 
stripes on tergites; palpus black 6 

5* Acrostichal setulae in 6 rows; mesonotum yellow to grey, postpronotum 
at least partly yellowish; legs usually mostly yellow, rarely femora 
strongly black; other characters variable 7 

6 Postpedicel black; presuturai dorsocentral seta always absent; female: 
tergite 3 short, at posterior margin medially with 4-6 very long, black, 
thick setae M. longiseta (Loew, 1 847) 

6* Postpedicel orange-brown, blackish at most along dorsal margin; presu- 
tural dorsocentral seta usually present, sometimes absent on one, rarely 
on both sides; female: tergite 3 not modified, with normal short, black 
setulae at posterior margin M. flaviventris (Costa, 1844) part 

7 Mesonotum and abdomen more or less uniformly yellow, rarely pale 
grey microtrichose along acrostichal setulae; palpus yellow; black 
latero ventral spots on scutellum small; female: tergite 3 unmodified and 
without long setae at posterior margin M. cypriota Papp, 1981 

7* Mesonotum at least with conspicuous medial grey stripe along acro- 
stichal setulae; abdomen yellow to brown, but tergites 3-5 usually each 
with a transverse black stripe which is interrupted in middle (M. fasciata 
species-group, see also Tab. 1) 8 

8 Ground colour of subscutellum yellow; scutellum lateroventrally with 
two large black spots which are larger than separating area (Figs 2, 4); 
mesonotum mainly yellow to brown, with only a rather narrow grey 
stripe medially; palpus yellow to brown; male: surstylus connected with 
subepandrial plate by a conspicuous sclerite (Figs 10-11); postgonite 
with 3 dorsal hooks (Fig. 14); female: tergite 4 with 2 pairs of very long 
black setae at posterior margin in middle M. czernyi Freidberg & Yarom, 1990 

8* Ground colour of subscutellum black; black spots lateroventrally on 
scutellum smaller, and wider apart from each other than length of one 
black spot (Figs 3, 5); mesonotum usually more extensively greyish 
along lateral margin with usually a grey band on level of presuturai and 
postsutural intraalar seta; female: either all tergites with usual setae and 
setulae (Fig. 6), or tergite 3 with 4-6 pairs of very long setae leaving a 
gap between them (Fig. 7) 9 

9 Male 10 

9* Female 12 

10 Surstylus strongly concave, with undulated distal margin which is 
conspicuously invaginated (Figs 28-29), sometimes with an additional 
medial projection (Figs 30-31) (mesonotum usually uniformly grey 
without paler stripes over lines of dorsocentral setae, but sometimes with 
a narrow, black stripe medially of line of dorsocentral setae) 
M. tabidiventris (Rondani, 1877) 



192 B. MERZ 

10* Surstylus, indistinctly concave, with a sharp projection at distal posteri- 
or margin (Figs 17, 22-23) (mesonotum usually with a more or less de- 
veloped paler stripe over line of dorsocentral setae) 11 

11 Surstylus distinctly longer than wide (Fig. 18); subepandrial plate with 
1-3 weaker and shorter setulae in addition to the long seta (Fig. 19); 
palpus usually entirely black in distal half or only indistinctly brownish 
ventrally M. fasciata (Fallen, 1826) 

11* Surstylus only slightly longer than wide, almost square (Fig. 24); sub- 
epandrial plate usually with one strong seta only (Fig. 22); palpus 
usually yellow, sometimes dorsally bordered by black margin 
M. subvittata (Loew, 1847) 

12 Tergite 8 evenly sclerotized (Fig. 40); tergite 9 distally with 2 thick, 
long, black setae (Figs 39-40); sternite 8 with evenly convex surface; 
ear-like projection distally rounded (Fig. 39); abdominal pleura at most 
with a small glandular area on level of tergites 4-5 (mesonotum usually 
uniformly grey without paler stripes over lines of dorsocentral setae, but 
sometimes with a narrow, black stripe medially along dorsocentral setae) 
M. tabidiventris (Rondani, 1877) 

12* Tergite 8 with two faint, L-shaped sclerites (Figs 44, 47); tergite 9 
without long apical setae (Figs 43, 46); sternite 8 with a medial longi- 
tudinal rib (Figs 45, 48); ear-like projection weakly invaginated, 
posteriorly sharply pointed (Figs 43, 46); a large glandular area present 
on pleura on level of tergites 4-5 (Fig. 8) (mesonotum usually with a 
more or less developed paler stripe through dorsocentral setae) 13 

13 Tergite 3 without long black setae along posterior margin (Fig. 6); 
palpus usually black at least in apical half, sometimes with brown or 
yellow border ventrally, but tip almost always black or dark brown; 
sternite 8 with more elongated ear-like lateral projection (Fig. 43), its 
apex more pointed and its ventral plate in ventral view larger, half-moon 
shaped (Fig. 45) M. fasciata (Fallen 1826) 

13* Tergite 3 with a row of long black setae at posterior margin which is in- 
terrupted in the middle, the middle 2-4 setae are longest and distinctly 
longer than width of tergite 4 (Fig. 7); palpus usually yellow to brown 
throughout, dorsal margin sometimes bordered by black, rarely more ex- 
tensively darkened, but usually with at least a distinct brown stripe to 
tip; sternite 8 with shorter lateral ear-like projection (Fig. 46), in profile 
more triangular, its ventral plate in ventral view short, with a lateral 
point (Fig. 48) M. subvittata (Loew, 1847) 

DESCRIPTION OF THE SPECIES OF THE MINETTIA FASCIATA SPECIES-GROUP 

The following part lists for each species first the synonymy, then the type 
material, followed by additional material studied. Labels of primary types are cited 
verbatim. The diagnosis does not repeat the characteristics given in the description of 



THE MINETT1A FASCIATA SPECIES-GROUP 193 

the M. fasciata species-group, but only those characteristics which are needed to 
separate a species from the other members of this species-groups. 

Minettia czernyi Freidberg & Yarom, 1990 Figs 2, 4, 9-15, 35-38 

Minettia quadrisetosa Czerny, 1937: 89. Type locality: [Israel]: Rehoboth bei Jaffa. 
Minettia czernyi Freidberg & Yarom, 1990: 98 (replacement name for Minettia quadrisetosa 
Czerny, 1937, nee Becker, 1907). 

Type material 

Lectotype â (here designated, examined): "Rehoboth bei Jaffa, 27.VII.33, J. 
Aharoni coll." (printed), "Minettia quadrisetosa Cz. S , det. L. Czerny" (in Czerny's 
handwriting), "Type Czerny 1935" (in Lindner's handwriting), "Lectotypus S 
Minettia quadrisetosa Czerny, 1937, desig. B. Merz 2003", Minettia czernyi Freidberg 
& Yarom 1990, det. B. Merz' 2003" (upper side), "n. name for M. quadrisetosa Czerny 
1937 nee Becker, 1907" (lower side of same label) (SMNS). The specimen is double 
mounted with a minutien pin, and is in good condition with strong colours, all legs, 
wings and antennae present, body slightly covered with debris; left posterior fronto- 
orbital seta, left medial postalar seta and both left scutellar setae are absent; both 
anterior fronto-orbital setae partly broken. 

Paralectotypes (examined): 19, same labels as lectotype; lo\ same labels as 
lectotype, but 26.VII.33, and "Cotype Czerny 1935" (in Lindner's handwriting) 
(SMNS). 

Comment: The species was described based on 4ó* o* and 2 9 9 from Rehoboth, 
25-27.VII.1933. Only 2<S 3 and 19 are present in the SMNS (Tschorsnig, in litt.). 
Freidberg & Yarom (1990) studied a couple in the NHMW which apparently belongs 
to the type series as well. The fate of the last S is not known. In order to avoid further 
confusion about the identity of the species, a lectotype is designated here. Another o" 
with the label "Rehoboth bei Jaffa, 25.XI.1933" (SMNS) is not a syntype because it 
does not correspond with the data mentioned in the original description. 

Other material examined (35 specimens) 

CYPRUS: Lemasos. ISRAEL: Antipatris, Dan, Ein Bazan, Hammat Gader, Herzliyya, 
Panyas, Ramat-Aviv, Tel-Dan. LEBANON: Aakkar. (CBM, MHNG, USNM) 

Diagnosis 

Head almost entirely yellow, with faint brown crossband at level of anterior 
fronto-orbital seta; palpus yellow; mesonotum mat yellow with at least broad grey 
stripe over line of acrostichal setulae, often with grey pattern between dorsocentral and 
intraalar setae; pleuron mostly yellow; scutellum lateroventrally with pair of very large 
black spots; legs almost entirely yellow, only forefemur anteroventrally in apical 
quarter and midfemur ventrally subapically with rather conspicuous brown spots; 
abdomen yellow with rather indistinct brown, narrow, transverse bars on tergites 3-5. 
Male: surstylus in profile slightly recurved (Fig. 10), slightly wider than long when 
maximally extended (Fig. 12), posterior margin medially concave (Fig. 10) and slightly 
folded downwards (Fig. 11); subepandrial plate entirely separated from surstylus, with 
one seta (Figs 10-11); two branches of pregonite of unequal size present (Figs 15), with 



194 B. MERZ 

left gonite much larger, apically swollen and with an indistinct apical tooth; postgonite 
very large, deeply concave medially (Figs 13-14) and posteriorly with 2+1 strong, 
black teeth (Fig. 14). Female: tergite 4 medially at posterior margin with 4 very long, 
and 2 shorter black setae, the longest setae longer than the following tergite; tergites 8 
and 9 well developed, without longer setae (Figs 15-16); sternite 8 basally with two 
small, ear-like, roundish projections with a channel-like sclerotization (Figs 35, 37-38); 
cercus small, with some longer setae apically. Wing length: (4.00) 4.25-4.75 mm. 

Remarks 

This species differs from the other 3 species of the group by the slightly longer 
wings on average, the more yellowish ground colour, in particular of the head, pleuron 
(the latter mostly grey in the other species) and subscutellum; the black spots on the 
lateroventral side of the scutellum are much larger; the legs have only two limited 
brown spots, and the terminalia (S and 9) are characteristic. It may be argued that M. 
czernyi does not belong to the M. fasciata species-group, due to genitalic differences, 
but taking into account the structural similarity with pale specimens of M. subvittata 
and M. fasciata it is difficult to place the species elsewhere at the present moment. 
Further studies of all described species of Minettia are needed before placing the 
species more accurately. 

Distribution 

So far only known from three Eastern Mediterranean countries: Cyprus, Israel, 
Lebanon. 

Minettia fasciata (Fallen, 1826) Figs 1, 6, 8, 16-20, 43-45 

Lauxania fasciata Fallen, 1826: 15. Type locality: Sweden: Skâne, Esperöd, nr. Stenshufvud. 
Sapromyza rivosa Meigen, 1826: 265. Type locality: not given ("im Sommer ziemlich gemein 

an Gestraden") (according to Papp, 1984: "? Germany"), [synonymy confirmed] 
Minettia nemorosa Robineau-Desvoidy, 1830: 647. Type locality: not given (according to Papp, 

1984: "France"). 

Type material 

Type material of fasciata 

Holotype $ (examined): "Lauxania fasciata $, Fall. Suppl" (handwritten); 
"Holotype $, Lauxania fasciata Fallen, desig. B. Merz 2002" (red label, handwritten) 
(ZML). The type material of this species is extensively discussed by Merz (2003b) and 
the comments do not need to be repeated here. 

Type material of rivosa 

Lectotype 6 (here designated, examined): "Meigen" (upper side of round label, 
handwritten); "2402 40" (lower side of same round label, handwritten); "rivosa" 
(square label, handwritten); "Lectotype â , Sapromyza rivosa Meigen, 1826, desig. B. 
Merz' 2002" (red square label, handwritten); "Minettia fasciata (Fallen), det. B. 
Merz'2002" (white label, handwritten) (MNHNP). The specimen is directly pinned 
and in a good state of preservation (only right hindleg, last 4 tarsal segments of left 
midleg and few setae missing; both wings slightly damaged; tip of abdomen slightly 



THE MINETTIA FASCIATA SPECIES-GROUP 



195 




Figs 9-15 
Male terminalia of Minettia czernyi Freidberg & Yarom: 9, epandrium, surstylus and inner 
terminalia, lateral view; 10, epandrium and surstyli, posterior view; 11, surstyli, oblique view; 
12, surstylus in maximal lateral extension; 13, inner terminalia, anterior view; 14, postgonite, 
oblique view; 15, pregonites and phallapodeme in maximal extension. 



196 B. MERZ 

covered with debris). The genitalia are partly covered with debris, but the right sur- 
stylus is clearly visible (as in Figs 16-19). 

Paralectotype 9 (examined): "Meigen" (upper side of round label, hand- 
written); "2402 40" (lower side of same round label, handwritten); "Paralectotype 9, 
Sapromyza rivosa Meigen, 1826, desig. B. Merz' 2002" (red square label, hand- 
written); "Minettia fasciata (Fallen), det. B. Merz'2002" (white label, handwritten) 
(MNHNP). The condition of the specimen is worse than the lectotype, with the left eye 
slightly squashed and the lunule sunken into the head. 

Comment: Meigen (1826) described this species based on an unknown number 
of specimens. Two specimens remain in the collection of the MNHNP. They agree well 
with the original description (p. 265) and illustration (Morge, 1975-1976, plate CXLII, 
Figs 11-12) and are considered to be syntypes. Because of the difficult taxonomy in the 
species-group I am designating a lectotype herein in order to avoid further confusion 
about the identity of this species. 

The synonymy of S. rivosa with L. fasciata was already proposed by Westwood 
(1840), but they were treated as good species by Collin (1948) and subsequent authors. 
The comparison of all available type specimens confirms the opinion of Westwood 
(1840). 

Lauxania fasciata has priority over S. rivosa because the exact publication date 
is not known for the latter and is therefore by default 31 December (ICZN, 1999), 
whereas L. fasciata was published on 6 December. 

Type material of nemorosa 

This species was described from a large number of specimens, but the collection 
of Robineau-Desvoidy, now in the MNHNP, is mostly destroyed. No specimen of this 
species could be found in the MNHNP (Charbonnel, in litt.). Because of the insuf- 
ficient description presented by Robineau-Desvoidy it is not possible to unambi- 
guously establish the identity of the species. It seems most reasonable to follow 
Macquart (1835), Westwood (1840) and most subsequent authors who synonymized it 
with L. fasciata and/or S. rivosa. 

Other material examined (598 specimens) 

Palaearctic - AUSTRIA: Süd-Steiermark. BELGIUM: Flanders: Knokke, Oostkamp. 
BULGARIA: Achtopol. CROATIA: Brazza Island; Unie Island; Dalmatia: Savudrja. 
DANMARK: Jylland: Hansted reserve. ENGLAND: Cambs: Wicken Fen; Herts: Harpenden; 
Oxfordshire: Wychwood Forest; Norfolk: Wretham Heats; Suffolk: Bury St. Edmunds; 
Wiltshire: Marlborough. FRANCE: Alpes-de-Haute-Provence: Montagnes de Lure; Alpes 
Maritimes: Menton, Nice, Sospel; Ariège: Salege; Aude: Aunat, Belcaire, Marsa; Aveyron: Nant; 
Bouches-du-Rhône: Marseille, Peyrolles; Calvados: Luc-sur-Mer; Corsica: Ghisonaccia, Lare; 
Doubs: Liesle, Vallée de la Loue; Drôme: Chalancon, Dieulefit; Gard: Camprieux, Dourbies; 
Haute Savoie: Bossey, Le Pas de TEchelle, Monnetier, Salève; Manche: Pontorson; Seine-et- 
Oise: Forêt St. Germain; Var: Les Maures, Sollies Pont; Yvelines: Fontainbleu, Maisons Laffitte. 
GERMANY: Bayern: Dachau, Etterschlag, München, Schöngeising; Berlin: Berlin; 
Brandenburg: Eberswalde. GIBRALTAR: Gibraltar. GREECE: Corfu: Hricida; Epyrus: Ionina; 
Peloponnes: Akhaia, Lakonfas. HUNGARY: Kalocsa. ITALY: Cosenza: Cirella, Grisolia, 
Marcellina, Santa Maria del Cedro, Verbicaro; Liguria: Monterosso al Mare; Lucca: Orecchiella; 
Macerata; Mantova: Marninolo; Puglia: Mte. Gargano; Sardegna: Cuglieri, Dorgali, Iglesias, 
Oliena, Villacidro; Sicilia: Etna, Linguaglossa, Nebrodi, Randazzo; Verona: Grezzana, 
Malchesine. LIECHTENSTEIN: Balzers, Ruggell. MALTA: Buskett, Chadwick Lakes, Ghadira, 



THE MINETTIA FASCIATA SPECIES-GROUP 



197 




Figs 16-20 
Male terminalia of Minettia fasciata (Fallen): 16, epandrium, surstylus and inner terminalia, lat- 
eral view; 17, epandrium and surstyli, posterior view; 18, surstylus in maximal lateral extension; 
19, surstyli, anterior view; 20, pregonites and phallapodeme in maximal extension. 



Gozo-Mgarr-ix-Xini, Mgiebah, Salina Bay, Wied Qanotta. MOROCCO: Rabat. POLAND: 
Szczecin (= Stettin). PORTUGAL: Estonil, Valenca. SERBIA & MONTENEGRO: Radovici. 
NETHERLANDS: Noord-Holland: Zandvoort; Zeeland: Ouddorp. SPAIN: Cadiz: Hozgarganta 
valley; Catalagna: Barcelona; Galicia: Santiago de Compostela; Mallorca: Drach. SWEDEN: 
Skane. SWITZERLAND: Bern: Ostermundigen; Fribourg: Mt. Vuilly; Genève: Bernex, 
Cartigny, Champel, Chancy, Dardagny, Jussy, Russin, Sézegnin; Schaffhausen: Beringen, 
Merishausen, Rüdlingen; Ticino: Gordola, Minusio, Mt. S. Giorgio; Vaud: Cudrefin; Valais: La 
Rippe, Leuk; Zürich; Zürich. TUNISIA: Ain Sobah, Tabarka. WALES: Glamorgan, Oxwich 
Bay, Kenfig. Nearctic - CANADA: British Columbia: Vancouver area. USA: Oregon: Sand 
Lake, Tillamook Co; Washington: Pierce Co. (CAS, CBM, CCK, CGB, CMCT, CME, CPG, 
ETHZ, INRA, MHNG, MHNN, NHMB, NRS, SIZK, SMNS, SMO, USNM, ZMUC, ZSM) 



198 B. MERZ 

Diagnosis 

Palpus black or at least dark brown in distal half, the orangish-brown base some- 
times extending ventrally almost to the tip; mesonotum usually not uniformly grey, but 
with faint to strong orange to yellow stripes along dorsocentral setae; these stripes may 
be narrow or very wide and almost reach the line of intraalar setae; lateroventral black 
spots on scutellum small; legs yellow, forefemur sometimes with dark brown pattern or 
entire legs more extensively darkened, including the apical quarter of all tibiae. Male: 
surstylus elongated, with an apical tooth (Fig. 17) and usually with 1 strong and 1-3 
weak setae on the subepandrial plate (Fig. 19); medial branch of pregonite with a sub- 
apical tooth, and apical tooth usually not prominent (Fig. 20); postgonite without tooth- 
like projection distally. Female: tergite 3 with unmodified black setae along posterior 
margin (Fig. 6); pleurae on level of tergites 4-5 with a conspicuous black glandular area 
(Fig. 8); tergite 8 with a pair of "L-"shaped sclerites (Figs 43-44); tergite 9 distally 
without long, black setae, usually wider than long, almost rectangular, smoothly con- 
cave medially along posterior margin (Fig. 44); sternite 8 raised medially, with a pair of 
long ear-like projections (Fig. 45); opening in lateral view more slit-like, elongated, api- 
cally sharply pointed (Fig. 43); ventral plate of ear-like projection in ventral view large, 
semicircular (Fig. 45). Wing length: 3.7-4.25 mm. 

Remarks 

Externally, this species is somewhat intermediate between M. tabidiventris and 
M. subvittata. The colour of the mesonotum is quite variable. Specimens from southern 
Europe tend to be paler than those from northern Europe and the mountains, and may 
then be easily confused with M. subvittata. Specimens with an entirely grey meso- 
notum (as in M. tabidiventris) are rare. A faint paler stripe over the line of dorsocentral 
seta is usually visible under certain illumination. 

M. fasciata is the sister species of M. subvittata, both having the same type of 
genitalia in both sexes, and females of both species have a large glandular area in the 
abdominal pleurum. The differences in the colour of the palpus as presented in the key 
apply to about 90% of the specimens. Those with a black tip clearly belong to M. fas- 
ciata, but a pale palpus may be found in conjunction with a rather yellow thorax in 
specimens of both species from mediterranean countries. The males differ clearly in the 
shape of the surstylus which is distinctly longer than wide in profile in M. fasciata 
(Fig. 18) and more square in M. subvittata (Fig. 24). Usually, the subepandrial sclerite 
carries 1-3 weak setulae in addition to the strong seta in M. fasciata (Fig. 19), but only 
one strong seta is present in M. subvittata (Fig. 23). The females can be separated by 
the chaetotaxy of the abdomen and the shape of the ear-like projections of sternite 8, 
as explained in the key. 

Distribution 

Widespread and common all over Europe. Also known from North Africa, but 
apparently absent from Israel. According to Shatalkin (2000) also in the Caucasus and 
the "Near East". An isolated population is known in the Western part of the Nearctic 
region. These specimens are identical with specimens from Western Palaearctic region. 
At the present moment it is impossible to explain this remarkable bimodal distribution 
of this species. 



THE MINETTIA FASCIATA SPECIES-GROUP 



199 




Figs 21-26 
Male terminalia of Minettia subvittata (Loew): 21, epandrium, surstylus and inner terminalia, 
lateral view; 22, epandrium and surstyli, posterior view; 23, surstyli, oblique view (paralecto- 
type); 24, surstylus in maximal lateral extension; 25, inner terminalia, anterior view (paralecto- 
type); 26, pregonites and phallapodeme in maximal extension. 



Minettia subvittata (Loew, 1 847) 



Figs 3, 5, 21-26, 46-48 



Sapromyza subvittata Loew, 1847: 28. Type localities: Italy: Neapel; Greece, Turkey 
("Kleinasien") 

Sapromyza Cataracta Pandellé, 1902: 398. Type localities: France: Marseille; Italy: Trieste, [syn- 
onymy confirmed] 

Minettia rivosa auctt. nee Meigen, 1826: Freidberg & Yarom (1990). 



200 B. MERZ 



Type material 



Type material of subvittata 

Lectotype 9 (here designated, examined): (small, white label with 2 black 
triangles on top), "Italien, Neapel, 8.1845, Zeller" (handwritten, blue label); "Coll. H. 
Loew" (printed, white label); "Typus" (printed, red label); "11596" (printed, white 
label); "Sapromyza subvittata m. [handwriting Loew], Type 9 Loew [handwriting of 
unknown person]" (white label); "Zool. Mus. Berlin" (printed, yellow label"); "Lecto- 
typus 9, Sapromyza subvittata Loew, 1847, desig. B. Merz'2002" (handwritten, red 
label). (ZMHB). The specimen is directly pinned and is in good condition, with only 
the right posterior fronto-orbital seta and left medial vertical seta missing, some setae 
on head partly broken; left lateral postalar seta and right second postsutural dorso- 
central seta absent; left wing slightly broken. 

Paralectotype 6 (examined): (small square, pink label), "KleinAsien, coll. H. 
Loew" (handwritten, blue label); "Coll. H. Loew" (printed, white label); "Paratypus" 
(printed, red label); "Sapromyza subvittata Cotype 9 Loew, 1847" (handwritten [not 
by Loew], white label); "Zool. Mus. Berlin" (printed, yellow label"); "Paralectotypus 
8, Sapromyza subvittata Loew, 1847, desig. B. Merz'2002" (handwritten, red label). 
The abdomen has been removed and is stored in glycerol in a glass vial on the same 
pin as the remaining specimen. Paralectotype S (examined): "95" (handwritten, small, 
square white label); "Rhodus, coll. H. Loew, Erber 5." (handwritten, blue label); "Coll. 
H. Loew" (printed, white label); "Paratypus" (printed, red label); "Zool. Mus. Berlin" 
(printed, yellow label"); "Paralectotypus S , Sapromyza subvittata Loew, 1847, desig. 
B. Merz'2002" (handwritten, yellow label); "Minettia tubifer (Mg.), det. B. 
Merz'2002" (handwritten, white label) (both ZMHB). 

Comment: Another female from the collection H. Loew (ZMHB) labelled 
"Italien, Neapel, Zeller S., 29.7.1845" does not belong to the syntypic series, because 
the collecting date (July) does not correspond with the original description (August). 
This specimen belongs to Minettia tubifer (Meigen). The abdomen of the specimen has 
been removed, and is kept in glycerol in a glass vial on the same pin. 

The lectotype is labelled "Typus", and the other specimens "Paratypus", but 
Loew (1847) did not designate a holotype and the red "Typus"-label was apparently 
added later to the specimen. It was probably this specimen that Czerny (1932) studied 
for his re-description and which he appeared to consider as the holotype (only by 
interference, he never referred to one of the specimens as the "Type" or "Holotype", 
but he did refer to Paratypes). However, his action was ambiguous and does not fulfill 
the requirements of the ICZN (1999), articles 74.5 or 74.6. Therefore, a lectotype 
designation is necessary in order to fix unambiguously the identity of the species. The 
lectotype corresponds perfectly with the description of Loew and maintains the current 
concept of the species (Shatalkin, 1998, 2000). 

Type material of Cataracta 

Lectotype 9 (here designated, examined): "9, 6522" (handwritten), "Sapro- 
myza Cataracta (Pandellé's handwriting), "Lectotypus Sapromyza Cataracta Pandellé, 
1902, 9. desig. B. Merz'2003" (red label), "Minettia subvittata (Loew), det. B. 



THE MINETTIA FASCIATA SPECIES-GROUP 



201 




Figs 27-34 
Male terminalia of Minettia tabidiventris (Rondani): 27, epandrium, surstylus and inner termi- 
nalia, lateral view; 28, surstyli, posterior view (paralectotype); 29-30, two extreme forms of 
epandrium and surstyli, posterior view; 31, surstylus, medial view of extreme form with medio- 
apical projection; 32, inner terminalia, anterior view; 33, aedeagal apodeme; 34, pregonites and 
phallapodeme in maximal extension. 



202 B. MERZ 

Merz'2003" (white label) (MNHNP). The specimen is directly pinned and is in very 
good condition (colours strong, little debris on integument; mesonotum medially 
sunken, legs slightly shrivelled). 

Comment: Although the species was described at least from two specimens, 
only the above mentioned female could be found in the MNHNP. (Charbonnel, in litt.). 
According to the catalogue of Pandellé, the code "6522" means "Marseille, Abeille 
(abdomen 3ème arc. avec une série marginale d'aiguillons couchés robuste atteignant 
le 5ème arceau)" [= collected by Abeille in Marseille, 3rd abdominal segment with a 
series of long setae reaching 5th segment] (Charbonnel, in litt.). It cannot be excluded 
that further type specimens occur elsewhere. Therefore it is proposed here that a lecto- 
type is designated in order to fix the identity of this species. 

The specimen is characterized by the entirely yellow palpus, the strongly 
plumose arista, the pair of black spots lateroventrally on the scutellum, 0+3 dorso- 
central setae and 3 pairs of strong medial and 3 pairs of weaker lateral setae at the 
posterior margin of tergite 3. Although the genitalia are retracted into the preabdomen 
it is obvious that tergite 9 does not have long black apical setae, and sternite 8 is 
bulging medially. This combination of characters is the same as in S. subvittata. 
Therefore this synonymy which was first proposed by Czerny (1932) without studying 
the types is confirmed herewith. 

Other material examined (237 specimens) 

CROATIA: Verudela. CYPRUS: Akamas peninsula, Akrotiri peninsula, Diarizos valley, 
Lemasos, Lefkara, Kalvasos, Kissousa. FRANCE: Alpes-de-Haute-Provence: Montagnes de 
Lure; Alpes Maritimes: Menton; Aude: St. Pierre de la Mer; Bouches-du-Rhône: Arles, Etang de 
Berre, Les Baux; Drôme: Curnier; Gard: Pont du Gard, Remoulins. Hérault: Grables, 
Montpellier, St. Gily-du-Fesc; Pyrénées-Orientales: Port Vendres; Var: Cavalière; Vaucluse: 
Cucuron; Yvelines: Maisons Laffitte. GREECE: Crete: Agios Galini, Agia Pelagia; Epyrus: 
Ioanina; Kilkis: Polycastro; Madeconia: Florina, Lahanas hills, Struma valley; Thessaloniki; 
Peloponnes: Lakonfas, Akhaia. HUNGARY: Kalocsa. ISRAEL: Aniam, Antipatris, Herzliyya, 
Mezdat Nimrod, Meron, Montfort, Mt. Carmel, Panyas, Park HaYarden, Ramat Aviv, Ramot 
Naftali, Rehobot, Qusbiye, Snir, Tel Aviv, Tel Dan, Zomet. ITALY: Cosenza: Marcellina, Santa 
Maria del Cedro; Puglia: Mt. Gargano; Trieste: Trieste. SERBIA & MONTENEGRO: Radovici. 
SPAIN: Cataluna: Barcelona. TURKEY: Antalya: Antalya, Cavusköy, Kemer, Phaseiis, Side; 
Isparta: Kovada lake; Samsun: Samsun. (CAS, CBM, CGB, CMCT, CME, CPG, INRA, MH- 
NG, MHNN, SIZK, SMNS, SMO, TAU, USNM, ZSM) 

Diagnosis 

More brownish than grey, with very wide orange to brown stripes along the 
lines of dorsocentral setae, leaving comparatively narrow grey stripes along the acros- 
tichal and intraalar setae, but dark specimens may have a similar pattern to M. fascia- 
ta; palpus yellow, rarely with dorsal margin brown to black; black latero ventral spots 
on scutellum small (Figs 3, 5); legs yellow, but femora often partly grey to black; black 
transverse stripes on tergites 3-5 (Fig. 7) tend to be narrower than in fasciata and 
tabidiventris but this is not a reliable character. Male: surstylus rather square in profile 
(Fig. 24); apically with a tooth-like projection at posterior margin; subepandrial plate 
with 1 strong black seta (Figs 22-23); medial pregonite with a subapical tooth, apical 
tooth barely longer (Fig. 26); postgonites distally without tooth like projection. 
Female: 3-4 pairs of medially separated, very long, black setae at posterior margin of 



THE MINETTIA FASCIATA SPECIES-GROUP 



203 




Figs 35-42 
Female terminalia of Minettia czernyi Freidberg & Yarom (35-38) and M. tabidiventris 
(Rondani) (39-42): 35, 39, lateral view; 36, 40, dorsal view; 37, 41, ventral view; 38, ear-like 
projection, enlarged; 42, spermathecae. 



204 B. MERZ 

tergite 3 (Fig. 7); abdominal pleurae with a dark glandular area on level of segments 
4-5; tergite 8 with a pair of "L-"shaped sclerites (Fig. 47); tergite 9 with moderatly 
enlarged black setae at distal margin (Fig. 46), about as wide as long, almost rhomboid, 
at posterior margin medially smoothly concave (Fig. 47); sternite 8 raised medially, 
with a pair of rather short ear-like projections (Fig. 48); opening in lateral view trian- 
gular (Fig. 46); ventral plate of this ear-like projection in ventral view small, laterally 
pointed. Wing length: 3.65-4.55 mm. 

Remarks 

This species resembles very much pale specimens of M. fasciata and a safe 
separation is only possible by the study of the terminalia ( S or 9 ) as given in the key. 
As explained under M. fasciata the colour of palpus and mesonotum are not fully 
reliable characters. 

M. subvittata may also be confused with dark specimens of M. czernyi. The 
latter species, however, always has much larger black spots lateroventrally on the 
scutellum (Fig. 2, 4), and differs in the genitalia of both sexes. Tergite 4 instead of 
tergite 3 has conspicuously long apical setae in M. czernyi. 

Distribution 

A circummediterranean species with northernmost records from Hungary and 
the Paris area (Yvelines: Maisons Laffitte). 



Minettia tabidiventris (Rondani, 1877) Figs 27-34, 39-42 

Sapromyza tabidiventris Rondani, 1877: 61. Type locality: not given (according to Papp, 1984: 

"Italy"). 
Sapromyza luteo-frontata Becker, 1895: 218. Type locality: "Herkunft nicht angegeben, daher 

wahrscheinlich aus Posen oder Schlesien". Syn. n. 
Minettia fasciata auctt. nee Fallen, 1826: Collin (1948); Papp (1979, 1984); Remm & Eiberg 

(1979); Freidberg & Yarom (1990); Shatalkin (2000). 

Type material 

Type material of tabidiventris 

Lectotype S (here designated, examined): "1553" (printed in red), "Lectotypus 
S, Sapromyza tabidiventris Rondani, 1877, desig. B. Merz' 2003" (red label) (MZF). 
The condition of the specimen is good, with strong colours and antennae and legs fully 
present; surstylus mostly visible. The following setae are missing: left posterior fronto- 
orbital, left medial vertical, right posterior notopleural, right anterior dorsocentral, left 
anterior katepi sternal and a series of setulae on the dorsal surface of the abdominal 
tergites. 

Parai ectotypes (examined): IS, 3 9$, with same label "1553", "Paralecto- 
typus Sapomyza tabidiventris Rondani, 1877, desig. B. Merz'2003". The male para- 
lectotype was dissected and the abdomen with the genitalia are stored in gylcerol in a 
glass vial which is attached to the pin of the specimen. The condition of the paralecto- 
types varies from rather good to bad, each specimen with some damage. One additional 
S is glued on a cardpoint and has the same label "1553", but represents the species 



THE MINETTIA FASCIATA SPECIES-GROUP 



205 




Figs 43-48 
Female terminalia of Minettia fasciata (Fallen) (holotype) (43-45) and M. subvittata (Loew) 
(46-48): 43, 46, lateral view; 44, 47, dorsal view; 45, 48, ventral view. 



Minettia lupulina (Fabricius, 1787), and therefore differs in various external characters 
from the original description and is not considered part of this type series. 

Comment: This species was described from an unknown number of specimens 
("non infrequens, exemplaribus preaecedentis [= subvittata Lw.] permixta"). All five 
syntypic specimens belong clearly to the same species, fitting perfectly the original 
description. However, because of the difficult taxonomy of the group with the in- 
clusion of another species in the series with the same number ("1553"), a lectotype is 
designated in order to fix the identity of the species. 

Type material of luteofrontata 

Holotype 9 (examined): "14/8 46" (handwritten); "Coll. H. Loew" (printed); 
"11607" (printed); "Type" (red label, printed); "luteo-frontata Beck." (Becker's hand- 



206 B. MERZ 

writing); "Holotypus 5, Sapromyza luteo-frontata Becker 1895" (red bordered label, 
L. Papp's handwriting); "Minettia fasciata Fall. 9, det. L. Papp" (handwritten); "Zool. 
Mus. Berlin" (yellow label, printed); "Minettia tabidiventris (Rondani), $, det. B. 
Merz'2003" (handwritten) (ZMHB). The specimen is in a moderately good state of 
preservation (no debris, pin not damaged, colours strong, but right posterior half of 
pleura (meron, anatergite, katatergite, anepimeron) and right half of abdomen eaten by 
pest and some setae on head and thorax missing). 

Comment: Becker (1895) described this species from one specimen in the Loew 
collection. He alternatively considered this species to be a male (title) or a female 
(diagnosis). Although not very comprehensive, the description of Becker fits the 
present specimen well. It is thus concluded to be the holotype. 

The comparison of the types of tabidiventris and luteofrontata clearly shows 
that they are conspecific and their synonymy is proposed here. 

Other material examined (255 specimens) 

CROATIA: Dubrovnik (Ragusa). CYPRUS: Akrotiri, Diarizos valley, Kissousa, 
Kofinous, Kouklia, Lemasos. DANMARK: Jylland: Thy. Nors. FRANCE: Alpes-de-Haute- 
Provence: Montagnes de Lure; Alpes Maritimes: Le Bar-sur-Loup; Aude: Marsa; Doubs: Vallée 
de la Loue; Drôme: Chalancon; Gard: Claret; Haute Savoie: Bossey, Le Pas de l'Echelle, 
Monnetier, Salève; Hérault: Grables, Lattes; Hautes Alpes: Agnielles; Haute Savoie: Albertville; 
Pyrénées-Orientales: Argèles-s-Mer, Collioure; Vaucluse: Beaumont. GERMANY: Baden- 
Württemberg: Lottstetten; Bayern: Bamberg, Dachau, Erlau b. Passau, Etterschlag, München, 
Schöngeising; Rheinland-Pfalz: Mainz; Sachsen-Anhalt: Wernigerode. GREECE: Corfu: Corfu, 
Merlin Marah; Macedonia: Seres; Sterea Elada: Itea, Paranssos mountains. HUNGARY: 
Kalocsa. ISRAEL: Mt. Carmel, Mt. Meron, N. Amud, N. Bezet, Panyas, Park HaYarden. ITALY: 
Trento: Mt. Lessini; Sicily: Nebrodi, Randazzo; Verona: Erbezzo. LIECHTENSTEIN: Balzers, 
Ruggell. MALTA: Buskett. NORWAY: Oslo: Hovedoya; Tey: Bamble. PORTUGAL: Praia des 
Macas. SPAIN: Catalana: Barcelona; Granada: Sierra de Alhama. SWEDEN: Gotland: Ardre, 
Ljugarn. SWITZERLAND: Bern: Mörigen, Sonceboz; Genève: Bernex, Cartigny, Chancy, 
Chèvres, Collonge-Bellerive, Dardagny, Russin, Sézegnin; Fribourg: Mt. Vuilly; Graubünden: 
Brienz-Surava, Valbella; Neuchâtel: Neuchâtel; Sankt Gallen: Betlis-Quinten; Schaffhausen: 
Merishausen, Rüdlingen; Schwyz: Gersau; Valais: Baltschieder, Leuk; Zürich: Dietikon, 
Unterengstringen, Winterthur, Zürich. TURKEY: Antalya: Cavusköy, Seiale. WALES: Tenby: 
Manorbier Beach. (CBM, CGB, CPG, CMCT, INRA, ETHZ, MHNG, MHNN, NHMB, NML, 
NRS, SMNS, SMO, TAU, USNM, ZMB, ZMUC, ZSM) 

Diagnosis 

Palpus always black at least in apical half; mesonotum uniformly grey mi- 
crotrichose, without paler stripes along lines of dorsocentral setae, but sometimes with 
narrow black stripe just medially of line of dorsocentral setae; scutellum lateroven- 
trally with two small black spots (as in Figs 3, 5); legs yellow, but anterior femur often 
dark brown to black and apices of tibiae often also darkened; transverse dark bands on 
tergites 3-5 broadly interrupted, leaving sometimes only 1-2 pairs of dark spots (as in 
Fig. 7). Male: surstylus almost square in profile, distal margin characteristically 
concave and invaginated (Figs 28-31); left pregonite usually with a simple apical tooth 
(Fig. 34); postgonite distally without protuberances; subepandrial plate usually with 1 
rather short seta only (Figs 28, 31). Female: without long setae along posterior margin 
of tergite 3; pleurae usually with only weak glandular area on level of tergites 4-5; 
tergite 8 forming a large sclerotized plate (Fig. 40); tergite 9 with a pair of very long, 



THE MINETTIA FASCIATA SPECIES-GROUP 207 

black, soft setae in the middle of the posterior margin (Fig. 40); sternite 8 indistinctly 
raised medially, ear- like projections not separated by depression (Fig. 41); ear-like 
projection in profile more or less parallel-sided, with deep invagination (Fig. 39). Wing 
length: 3.75-4.45 mm. 

Variability 

Specimens from Southern Europe (France, Alpes Maritimes; Italy, Sicily; 
Greece, Macedonia; Malta) show considerable variation in the outline of the distal 
margin of the surstylus. It is evenly concave in typical specimens (Figs 28-29) but there 
is a tendency to develop an additional tooth-like projection near the middle. The most 
extreme forms exhibit a very strong medial projection giving the impression of the 
presence of 3 lobes (Figs 30-31). The shape and orientation of this projection is sub- 
ject to strong variability. Because of the continuous variation of this character and the 
presence of different types of projections in the same population it is concluded that 
one variable species only is involved. 

Remarks 

M. tabidiventris differs from the couplet fasciata and subvittata by the usually 
entirely grey mesonotum without paler stripes over the lines of the dorsocentral setae, 
in the male in the shape of the surstylus and the left pregonite and in the female in the 
entirely sclerotized tergite 8, the long pair of setae along the posterior margin of tergite 
9 and the structure of sternite 8 with a slightly concave surface and parallel- sided ear- 
like projection in profile. 

Distribution 

Known throughout Europe except the very north of Scandinavia, Turkey and 
Israel. Apparently absent from North Africa. According to Shatalkin (2000) also in the 
Caucasus. 

DOUBTFUL SPECIES 

Lauxania pallida Meigen, 1 830, nomen dubium 

Lauxania pallida Meigen, 1830: 381. Type locality: not given ("von Herrn von Winthem"). 

No syntypic specimens could be found in the collections of Meigen in Paris 
(MNHNP, Charbonnel, in litt.), nor in Vienna (NHMW, Contreras-Lichtenberg, in litt.). 
The description is short, and the species is compared by Meigen with Minettia lupu- 
lina (Fabricius, 1787), differing from the latter by the black antenna, entirely trans- 
parent wing and the apparently uniformly grey scutellum dorsally (see Morge, 1975- 
1976, plate CCX, Fig. 8). Although sometimes treated as a species of the Minettia 
fasciata group (Papp, 1984) it seems more reasonable to assume that L. pallida either 
is a senior synonym of Sapromyza longiseta Loew, 1847 (if Meigen's illustration of 
pallida is a male) or it is one of the species of the M. lupulina group, as it was 
suggested by various authors who synonymized it with Sapromyza dimidiata Loew 
(Becker, 1902; Hendel, 1908; Czerny, 1932). Because of the insufficient description 
and the lack of type specimens, L. pallida must be considered at the present moment 
as nomen dubium. 



208 



B. MERZ 



o 
e 

00 



ü_ 



u. 



o- 

3 

c 



«, 



M 

4-> CU 

3 ea 

^x 

"33 cu 
'S. c 

CS U 

tS >» 
03 Th 
_-■ 3 

CO 3 




X) 


3 


^8 


>i 


>i 


33 


32 


cd 


CO 


O 2 

Ih T3 


3 
co 


3 

CO 


3 


3 




& 


CU 




O 


'S 



.33 c 



£ 


£ 




u 


_o 


>-. 


|H 


C 

ce 


"S 


<u 


eu 




00 


> 
c 


S 

T3 


>> 

"3 


O 


ce 
o 
c 


'— 
C 



■3.8. 



00. 



«ê * 

. £^ 

eu 00 <u 

^? i CL 

x -C •- 

d c« i; 

'C "I » 

« O C 

> 3 3 



" -S 



Ses - ~ 
-r 4» "5! 



.11 

§■£ 

-"S eu 
ûû > eu 

ö V, c 

'3d o 2 

■° «"§ 

« u rt 

eu g O 

CA U Da 



se 



ÎS * p 

Cy CS 
.S -g 

il 

-£-Ox 



es u -g 

E 5 1 



^ o 

cS OC co 



O 

a. 



OC 


m — 








§i 


OC S 






c« 


G <- 
— T3 






CO 

(U 


».2 






CJ 

o 

■— 

eu 


4*8 








£ S 


■o '& 






33 


5 


eu es 






cd 


3-c 






s 


fi eu 


c« .33 

■è * 
11 

ai 






co p. 


•t. -M 

s- ~ 

AS 


-5 

o 

c 


00 

c 
o 


i-T ■" 
co X) 



e 
« 

oc 
c 
o 



^h co X) 



CU > 

c o 



.2 
■3 



-S J 



oc 

fi eu 
© « 



« .g 

a oc 



-o o 
.» 8, 3 



o 



ö ä 



es -j3 

-a ■— 
<u -a 



T3 

S="S 

cS <4 

Ih 

>-. O 
32 C 

« "3 
IS co 
^ -S 



£•8 

il 

«1 

S Ih 

5 00 
c« C 



>> O 31 



« «S 

O P 



E .S -S a 



es 



1) 2 



C .3 co ce 



■g 

I 

O 



eu 

IH 

M 

5 g 3 

S .S « 
— u 

-C 00^ 

I 8.8 



oc 5 
u — 



-3 
_ s 
5 E 



Th 

>> 3 
00 

(U 



eu co 
*5jd "«S 



»? 

■S *s oc 

111 if 

sfa -s 

CO CO S I-H 



c3 





C- 


-* 


'3 


çu 




le 


fi 


o 

X 


a 


en 


n 


O 


oc 


u 


s- 


c 



-3 -^ O 
3 "33 

oo 3 



"fi — 



eu 
-= 

OC 

'■S ü 

3 s C 

O _ 

.S 

=3 eu 

CÖ -Q 

si 



Xi 



c 




3 


eu 




c_ 




CU 


cS 




o 

— 




X 
3 
co 


co 






E 


c*- 


3 




X 


O 






•— 
z 


_3 


3 


C5Û 
— 

CS 


CO 


eu 


3 


O 


s 


'I 


(M 


eu 

CO 


U 


Ph' 


o 



oo eu 



33 00 



E -ru 
oo pq E 

..eu 

Dh PL, to 



THE MINETTIA FASCIATA SPECIES-GROUP 209 

TAXONOMIC SUMMARY 

The Minettia fasciata species-group contains at present four valid species: 

Minettia czernyi Freidberg & Yarom, 1990 

= M. quadrisetosa Czerny, 1932 (junior primary homonym of M. quadrisetosa 
Becker, 1907) 

Minettia fasciata (Fallen, 1826) 

= M. rivosa (Meigen, 1826) (synonymy first proposed by Westwood, 1840, 

confirmed herein) 
= M. nemorosa (Robineau-Desvoidy, 1830) [synonymy doubtful] (synonymy of 

rivosa with nemorosa proposed by Macquart, 1835: 403) 

Minettia subvittata (Loew, 1847) 

= M. Cataracta (Pandellé, 1902) (synonymy first proposed by Czerny, 1932, 

confirmed herein) 
= M. rivosa auctt. nee Fallen, 1826: Freidberg & Yarom (1990) 

Minettia tabidiventris (Rondani, 1877) 

= M. luteofrontata (Becker, 1895), syn. n. 

= M. fasciata auctt. nee Fallen, 1826: Collin (1948), Papp (1979, 1984), Freid- 
berg & Yarom (1990), Shatalkin (1998, 2000) 

ACKNOWLEDGEMENTS 

It is my pleasure to thank my colleagues and the curators of the institutions 
listed in the "Material and Methods" section who sent me precious specimens in their 
care and who supplied me with important information. In particular I would like to 
thank Mrs J. Charbonnel (Paris) for her patience with my frequent questions concern- 
ing types and label information. I extend my thanks to L. Papp (Budapest), A. Shatalkin 
(Moscow) and I. Yarom (Hazeva) for fruitful discussions and to W. Schacht (Munich) 
who put an unpublished version of the English translation of the keys of Shatalkin 
(2000) at my disposal. F Marteau (Geneva) helped me with the preparation of the 
plates, and S. Gaimari (Sacramento) reviewed a draft of the manuscript. 

REFERENCES 

Becker, Th. 1895. Dipterologische Studien II. Sapromyzidae. Berliner Entomologische Zeit- 
schrift 40: 171-264. 

Becker, Th. 1902. Die Meigen 'sehen Typen der sogen. Muscidae acalypterae (Muscaria 
holometopa) in Paris und Wien. Zeitschrift für systematische Hymenopterologie und 
Dipterologie 2: 209-256. 

Collin, J. E. 1948. A short synopsis of the British Sapromyzidae (Diptera). Transactions of the 
Royal entomological Society of London 99 (5): 225-242. 

Czerny, L. 1932. 50. Lauxaniidae (Sapromyzidae). In: Lindner, E. (ed.). Die Fliegen der 
Palaearktischen Region, Vol. 5. E. Schweizerhart' sehe Verlagsbuchhandlung, Stuttgart, 
76 pp. 

Czerny, L. 1937. Minettia quadrisetosa sp. n. (Dipt.). Konowia 16: 89. 



210 B. MERZ 



Fallen, C. F. 1826. Supplementum Dipterorum Sveciae. 16 pp. Lundae. 

Freedberg, A. & Yarom, I. 1990. The Lauxaniidae (Diptera) of Israel, with an emphasis on 
Minettia. Israel Journal of Entomology 24: 93-105. 

Hendel, F. 1908. Diptera. Farn. Muscaridae. Subfam. Lauxaninae [sie]. In: Wytsman, P. (ed.). 
Genera Insectorum, Fascicule 68, 66 pp. & 3 plates. 

ICZN, 1999. International Code of Zoological Nomenclature. Fourth Edition. The International 
Trust for Zoological Nomenclature, London, XXIX + 306 pp. 

Loew, H. 1847. Dipterologische Beiträge. Dritter Theil. Ueber die europäischen Arten der 
Gattung Sapromyza. Jahresbericht des naturwissenschaftlichen Vereins zu Posen für 
1846: 25-44. 

Macquart, M. 1835. Histoire Naturelle des Insectes. Diptères. Tome deuxième. Librairie Ency- 
clopédique de Roret, Paris, 703 pp. 

Meigen, J. W. 1826. Systematische Beschreibung der bekannten Europäischen zweiflügeligen 
Insekten. Fünfter Theil. Hamm, 412 pp. 

Meigen, J. W. 1830. Systematische Beschreibung der bekannten Europäischen zweiflügeligen 
Insekten. Sechster Theil. Hamm, 401 pp. 

Merz, B. 2003a. Einführung in die Familie Lauxaniidae (Diptera, Acalyptrata) mit Angaben zur 
Fauna der Schweiz. Mitteilungen der Entomologischen Gesellschaft Basel 52 (2-3) 
(2002): 29-128. 

Merz, B. 2003b. The Lauxaniidae (Diptera) described by C. F. Fallen with description of a 
misidentified species of Homoneura van der Wulp. Insect Svstematics and Evolution 34: 
345-360 . 

Miller, R. M. 1977. Ecology of Lauxaniidae (Diptera: Acalyptrata). I. Old and new rearing 
records with biological notes and discussion. Annals of the Natal Museum 23: 215-238. 

Morge, G. 1975-1976. Dipteren-Farbtafeln nach den bisher nicht veröffentlichten Original- 
Handzeichnungen Meigens: "Johann Wilhelm Meigen: Abbildung der europaeischen 
zweiflügeligen Insecten, nach der Natur". Beiträge zur Entomologie 25 (2) (1975): 383- 
500 & plates 1-80; 26 (1) (1976): plates 81-160; 26 (2) (1976): plates 161-305. 

PandellÉ, L. 1902. Etudes sur les Muscidaes de France. 3ème partie, suite. Revue d'Entomo- 
logie 20 (Supplément): 373-492. 

Papp, L. 1981. New species and taxonomical data of the Palaearctic Lauxaniidae and Carnidae 
(Diptera). Acta Zoologica Academiae Scientiarum Hungaricae 27: 159-177. 

Papp, L. 1979. 4. Füzet. Korhadeklegyek-Pajzstetülegyek. Lauxaniidae-Chamaemyiidae (57 
abraval). Fauna Hungariae 136: 1-59. 

Papp, L. 1984. Family Lauxaniidae (pp. 193-213). In: Soós, A. & Papp, L. (eds). Catalogue of 
Palaearctic Diptera. Vol. 9. Akadémiai Kiado, Budapest, 460 pp. 

Papp, L. & Darvas, B. (eds) 2000. Contributions to a Manual of Palaearctic Diptera. Volume 1. 
Science Herald, Budapest, 978 pp. 

Reddersen, J. 1994. Distribution and abundance of lauxaniid flies in Danish cereal fields in 
relation to pesticides, crop and field boundary (Diptera, Lauxaniidae). Entomologiske 
meddelelser 62: 117-128. 

Reddersen, J. 1995. Feeding biology of fungivorous insects from Danish cereal fields. Pedo- 
biologia 39 (4): 370-384. 

Remm, E. & Elberg, K. 1979. Terminalia of the Lauxaniidae (Diptera) found in Estonia, Latvia 
and Lithuania. Dipteroloogilisi Uurimusi: 66-117. 

Robineau-Desvoidy, J. B. 1830. Essai sur les Myodaires. Mémoires présentés par divers 
Savants à l'Académie royale des Sciences de l'Institut de France. Second Volume. Impri- 
merie Royale, 813 pp. 

Rondani, C. 1877. Species Italicae Ordinis Dipterorum, pars sexta. Stirps XLX, Sciomyzinae. 
Annuario della Società dei Naturalisti Modena 1 1 : 7-79. 

Schiner, J. R. 1863. Fauna Austriaca. Die Fliegen (Diptera) (fase. 9-10, pp. 81-288). Band 2. 
Carl Gerolds Sohn, Wien. XXXII + 658 pp. 



THE MINETTIA FASCIATA SPECIES-GROUP 21 1 



Shatalkin, A. I. 1998. Palaearctic species of the genus Minettia (Diptera, Lauxaniidae). 

Zoologiche skii Zhurnal 11 (7): 809-817. (In Russian). 
Shatalkin, A. I. 2000. Keys to the palaearctic flies of the family Lauxaniidae (Diptera). Zoo- 

logicheskie Issledovana 5: 1-102. (In Russian). 
Smith, K. G. V. 1989. An introduction to the immature stages of British Flies. Handbooks for the 

Identification of British Insects 10 (14): 1-280. 
Stuckenberg, B. R. 1999. Antennal evolution in the Brachycera (Diptera), with a reassessment 

of terminology relating to the flagellum. Studia Dipterologica 6 (1): 33-48. 
Westwood, J. O. 1840. Synopsis of the Genera of British Insects. In: An Introduction to the 

Modern Classification of Insects. Volume 2. London, 158 pp. 
Zetterstedt, J. W. 1847. Diptera Scandinaviae. Disposita et Descripta 6: 2163-2580. Lundae. 



Revue suisse de Zoologie 111(1): 213-223; mars 2004 



A new species of Astyanax (Characiformes, Characidae) from the 
upper rio Bermejo basin, Salta, Argentina 

Juan Marcos MIRANDE 1 , Gaston AGUILERA 1 & 
Maria de las Mercedes AZPELICUETA 2 

1 Fundación Miguel Lillo, Miguel Lillo 25 1 , 4000 Tucumân, Argentina. 
E-mail: mcpiranha@hotmail.com 

2 Division Zoologia Vertebrados, Facultad de Ciencias Naturales y Museo, Paseo del 
Bosque, 1900 La Plata, Argentina. 

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

A new species of Astyanax (Characiformes, Characidae) from the 
upper rio Bermejo basin, Salta, Argentina. - Astyanax latens sp. n., 
from the Bermejo river basin, is described in this paper. Astyanax latens 
is diagnosed by a combination of characters: somewhat rhomboidal 
body; presence of 24-29 branched anal fin rays; 1-4 maxillary teeth; long 
pectoral fins (23.2-26.6 % of SL) always reaching one third of pelvic fin; 
long pelvic fins (17.6-22.0 % of SL) always surpassing anal fin origin; 
anal fin base long (32.3-36.7 % of SL); 26-27 gill rakers on first 
branchial arch; lateral band extremely narrow; one large vertically elon- 
gated humeral spot and a second one very faint; relatively shallow cau- 
dal peduncle (10.0-11.4 % SL); fifth tooth of inner premaxillary series 
very small and scarcely posterior to main series; origin of anal fin below 
dorsal fin. 

Key- words: Characiformes - Characidae - Astyanax - new species - no 
Bermejo basin. 

INTRODUCTION 

The genus Astyanax Baird & Girard, 1854 includes more than 100 nominal 
species (Garutti & Britski, 2000), distributed from Southern United States to the rio 
Negro in Argentina (Almirón et al., 1997). Since the original description of the genus, 
the most complete revision of Astyanax has been done by Eigenmann (1921, 1927) 
who diagnosed the genus with characters that are present in many Characiformes. 
Subsequently, many authors followed that classification (among others, Ringuelet et 
ah, 1967; Géry, 1977; Bertaco & Malabarba, 2001). Traditionally, the genus is defined 
by the presence of two series of teeth in the premaxilla, the first series with several 
teeth, a second series with equal or graduated teeth, usually five in number, four in a 
few species; crowns of premaxillary and mandibulary teeth usually ridged and denti- 
culated; maxillary with few or none teeth; complete lateral line, gill rakers setiforms. 



Manuscript accepted 19.09.2003 



214 J- M. MIRANDE ET AL. 

and absence of predorsal spine and scales on the caudal peduncle base. The combina- 
tion of those characters still defines the species recognized within the genus which has 
never been considered in a phylogenetic revision. 

About 20 species of the genus Astyanax were previously cited in Argentina. 
Some of those records need to be revised because they probably correspond to misiden- 
tifications due to external morphological similarities with species from other basins. A 
detailed revision of the species of the Rio de la Plata basin is necessary, although, sys- 
tematic papers concerning species of the genus Astyanax were published in the last 
recent years. Those papers reveal the presence of seven new species in the northeastern 
Argentina (Azpelicueta & Garcia, 2000; Azpelicueta et al., 2002a, 2002b; Almirón et 
al., 2002; Casciotta et al., 2003a, 2003b). Also, new collecting trips in the northwestern 
Argentina show the presence of new species, fact that allows us to describe A. latens 
sp. n. from the upper rio Bermejo basin. The description of the new species is the 
objective of this paper, providing osteological information that may be useful in future 
phylogenetic studies. 

MATERIAL AND METHODS 

Specimens were cleared and counterstained (C&S) following Taylor & Van 
Dyke (1985). Measurements are straight distances taken with calliper to nearest 0.01 
mm. Standard length was measured from snout tip to hypural joint; head length 
includes opercular flap; length of caudal peduncle was the distance between last anal 
fin ray insertion and hypural joint. Counts of vertebrae include Weberian apparatus and 
the caudal complex centrum as one element. 

Material is deposited in Asociación Ictiológica, La Plata (AI, asociacionictio- 
logica@interlap.com.ar); Facultad de Ciencias Naturales y Museo, La Plata (MLP); 
Fundación Miguel Lillo, Tucumân (CI-FML); Museo Argentino de Ciencias naturales 
Bernardino Rivadavia, Buenos Aires (MACN); Museum d'histoire naturelle de 
Genève, Genève (MHNG); The Academy of Natural Sciences of Philadelphia, Phila- 
delphia (ANSP). 

Comparative material (SL in mm). Acrobrycon tarijae Fowler, 1940: CI-FML 3270, 1 
ex., 66.6 mm, Argentina, Salta, Orân, La Bambù, rio Bermejo basin, rio Bianco. Astyanax 
abramis (Jenyns, 1842): MLP 9427, 2 ex., 102.0-113.0 mm, Argentina, Misiones, rio Parana. 
Astyanax cf. abramis: CI-FML 3368, Argentina, Salta, Orân, La Bambù, rio Bermejo basin, rio 
Bianco. Astyanax alleni (Eigenmann & McAtee, 1907): MLP 6774, 5 ex., 50.0-64.2 mm, 
Argentina, Santa Fe, Laguna Setübal. Astyanax asuncionensis Géry, 1972: MLP 8660, 5 ex., 
43.6-61.4 mm, Argentina Santiago del Estero. Banado de Anatuya. MLP 8844, 7 ex., 25.0-44.9 
mm, Argentina, Formosa, rio Bermejo basin, arroyo Mbiguâ. Astxanax eigenmanniorum (Cope, 
1894): ANSP 21627, paratypes, 2 ex., 42.5-49.4 mm, Brasil, Rio Grande do Sul. Astyanax cf. 
eigenmanniorum: CI-FML 3262, 23 ex., 29.4-50.7 mm, Argentina, Salta, La Bambù, rio Ber- 
mejo basin, rio Anta Muerta. Astyanax cf. fasciatus: MLP 7115, 1 ex., 47.8 mm, Argentina, Santa 
Fe, rio Parana basin, San José del Rincón, AI 111, 2 ex., 57.8-62.6 mm, Argentina, Buenos Aires, 
rio de la Piata in Punta Lara. MLP 8647, 5 ex. 34.0-45.8 mm, Argentina, Santiago del Estero, rio 
Salado. Astyanax ita Almirón et al, 2002: MLP 9599, holotype, 64.0 mm, Argentina, Misiones, 
rio Iguazù basin, arroyo Tateto. Astyanax leonidas Azpelicueta et al., 2002: MLP 9580, holotype, 
45.6 mm, Argentina, Misiones, rio Parana basin, headwaters of arroyo Urugua-i. Astyanax 
lineatus (Perugia, 1891): CI-FML 3272, 3 ex., 35.3-72.1 mm, Argentina, Salta, Orân, La Bambù, 
rio Bermejo basin, rio Bianco. Astyanax ojiara Azpelicueta & Garcia: MLP 9470, holotype, 50.5 
mm, Argentina, Misiones, arroyo Benftez. Astyanax pynandi Casciotta et al., 2003a: MACN 



NEW ASTYANAX FROM RIO BERMEJO BASIN 215 



8543, holotype, 52.0 mm, Argentina, Corrientes, Laguna Iberâ. Astyanax saguazu Casciotta et 
al, 2003b: MLP 9603, holotype, 63.0 mm, Argentina, Misiones, rio Uruguay basin, arroyo Once 
Vueltas. Astyanax troya Azpelicueta et al., 2002, MACN 8310, holotype, 73.8 mm, Argentina, 
Misiones, arroyo Cunapiru Chico. Bryconamericus iheringii (Boulenger, 1887): MLP 9073, 110 
ex. (5 measured), 39.9-44.3 mm, Argentina, Buenos Aires, Sierra de la Ventana. MLP 9103, 15 
ex., 34.8-49.2 mm, Argentina, Buenos Aires, Berisso, Los Talas (Man-made ponds connected to 
Rio de la Plata). Bryconamericus thomasi Fowler, 1940: CI-FML 1969, 94 ex. (5 measured, 2 
males and 3 females), 40.3-55.4 mm, Argentina, Salta, rio Piedras. Moenkhausia intermedia 
Eigenmann, 1908: CI-FML 3257, 60 ex. (5 measured), 20.1-31.5 mm, Argentina, Salta, Orân, La 
Bambù, rio Bermejo basin, arroyo El Oculto. Odontostilbe microcephala Eigenmann, in 
Eigenmann & Ogle, 1907: CI-FML 3369, 2 ex., 48.2-51.6 mm, Argentina, Salta, La Bambù, rio 
Bermejo basin, rio Blanco. Odontostilbe pequira (Steindachner, 1882): CI-FML 3451, 1 ex., 
31.5 mm, Argentina, Salta, Rivadavia, rio Bermejo basin, Pozo de los Yacarés. Oligosarcus boli- 
viano (Fowler, 1940): CI-FML 3277, 4 ex., 89.8-113.8 mm, Argentina, Salta, Orân, La Bambù, 
rio Bermejo basin, rio Bianco. 

Cleared and stained material (Personal coallection). Astyanax abramis: 2 ex., 74.5-92.0 
mm, Argentina, Buenos Aires, rio de la Piata in Punta Lara. Astyanax asuncionensis: 2 ex., 80.4- 
92.7 mm, Argentina, Misiones, rio Uruguay in San Isidro. Astyanax eigenmanniorum: 1 ex., 45.0 
mm, Brasil, Rio Grande do Sul, Viamâo, açude Charolês; 2 ex., 17.7-33.0 mm, Argentina, 
Buenos Aires, desembocadura del rio Colorado; 1 ex., 60.3 mm, Argentina, Buenos Aires, Rio 
de la Plata. Astyanax cf. fasciatus: 2 ex., 91.0-106.5 mm, Argentina, Misiones, rio Uruguay in 
San Isidro. Astyanax latens sp. n.: 2 ex., 40.6-44.0 mm, Argentina, Salta, Orân, La Bambù, rio 
Bermejo basin, arroyo El Oculto. Astyanax leonidas: 6 ex., 33.0-45.6 mm, Argentina, Misiones, 
rio Parana basin, arroyo Urugua-i. Astyanax ojiara: 10 ex., 37.8-58.0 mm, Argentina, Misiones, 
arroyo Bem'tez. Astyanax paris: 3 ex., 66.2-68.5 mm, Argentina, Misiones, rio Uruguay, arroyo 
Yabotf Guazù. Astyanax saguazu: 3 ex., 41.6-54.2 mm, Argentina, Misiones, rio Uruguay basin, 
arroyo Once Vueltas. Astyanax troya: 2 ex., 76.0-81.5 mm, Argentina, Misiones, rio Parana, 
arroyo Cunapirü Chico. Astyanax sp. A: 1 ex., 73.5 mm, Argentina, Misiones, arroyo Cunapirù 
Chico. 



RESULTS 

Astyanax latens sp. n. Figs 1-5, Table 1 

Holotype. CI-FML 3400 male, 44.3 mm SL, Argentina, Salta, rio Bermejo basin, arroyo 
El Oculto (64° 30' W - 23° 07' S), coli. M. Mirande & G. Aguilera, October 2001. 

Paratypes. MHNG 2640.36, 5 ex., 44.0-52.0 mm SL, collected with the holotype. AI 
110, 4 ex., 40.3-45.0 mm SL, Argentina, Salta, rio Bermejo basin, arroyo El Oculto, coll. M. 
Mirande & G. Aguilera, February 2002. CI-FML 3401, 5 ex., 46.8-52.4 mm SL, Argentina, 
Salta, rio Bermejo basin, arroyo El Oculto, coll. M. Mirande & G. Aguilera, November 2002, 
CI-FML 3402, 10 ex., 42.8-47.8 mm, Argentina, Salta, rio Bermejo basin, arroyo El Oculto, coll. 
M. Mirande & G. Aguilera, February 2002. AI 112, 2 ex. C&S, 41.0-45.0 mm, Argentina, Salta, 
Orân, arroyo El Oculto, coll. M. Mirande & G. Aguilera, February 2002. 

Diagnosis. Astyanax latens is distinguished from all other Astyanax species by 
the possession of somewhat rhomboidal body (body depth 34.9-39.1 % SL); 24-29 
branched anal fin rays; 1-4 maxillary teeth; long pectoral fins (23.2-26.6 % of SL) 
always reaching one third of pelvic fin; long pelvic fins (17.6-22.0 % of SL) always 
surpassing anal fin origin; anal fin base long (32.3-36.7 % of SL); 26-27 gill rakers on 
first branchial arch; origin of anal fin placed below dorsal fin; fifth tooth of inner pre- 
maxillary series very small and scarcely posterior to main series; presence of one large 
vertically elongated humeral spot followed by a second one very faint; lateral band 
extremely narrow; 37-38 perforated scales in the lateral series, and relatively shallow 
caudal peduncle (10.0-11.4 % of SL). 



216 J- M. MIRANDE ETAL. 

Description. Morphometries of holotype, 20 paratypes and 2 C&S specimens 
are presented in table 1. Body somewhat rhomboidal (Fig. 1), laterally compressed, 
with maximum body depth anterior to dorsal fin origin. Dorsal profile of body straight 
from snout to supraoccipital area, slightly concave above supraoccipital spine, convex 
from this point to dorsal fin origin; slanted ventrally from dorsal fin origin to caudal 
peduncle; gently concave along caudal peduncle to base of caudal fin rays. Ventral pro- 
file of body convex from tip of lower jaw to pelvic fin origin, somewhat straight 
between pelvic and anal fin origins, posterodorsally slanted from anal fin origin to 
caudal peduncle, and slightly convex along caudal peduncle. Ventral portion of body 
between bases of pectoral and pelvic fins transversally rounded; ventral portion of 
body between origins of pelvic and anal fins laterally compressed. 

Dorsal fin origin usually nearer base of caudal fin rays than snout tip (see table 
1). Pelvic fin origin situated markedly anterior to vertical through dorsal fin origin. 
Anal fin origin located on vertical through base of posteriormost dorsal fin rays. Tip of 
pectoral fin always reaching one third of pelvic fin or more; tip of pelvic fin always sur- 
passing anal fin origin, in all specimens independently from size and sex. 

Dorsal profile of head convex on snout and over eyes and slightly concave on 
supraoccipital area. Mouth terminal, placed at level of middle eye. Premaxilla bearing 
two series of teeth; ascending process with broad base and acute tip. Outer row with 
3-5 (5 ex.= 3); (10 ex.= 4); (7 ex. including the holotype= 5) tricuspidate teeth, with 
circular base and distal tip slightly compressed; inner row with 5 teeth, anterior face of 
them slightly concave; symphysial tooth slender, with 4 cusps; remaining teeth with 
five, four or three cusps; central cusp large and lateral ones small; fifth tooth much 
smaller than other teeth in series, usually tricuspidate, inserted posteriorly to fourth 
tooth (Fig. 2). Ascending maxillary process slender, straight; lateral process of maxilla 
long, laminar, and narrow, with 1-4 (1 ex.= 1); (16 ex.= 2); (4 ex. including holotype= 
3); (1 ex.= 4) tricuspidate teeth (Fig. 3). Dentary with 11-13 teeth (2 ex.= 11); (1 ex.= 
12); (1 ex.= 13); first three or four pentacuspidate and notably larger than remaining 
tricuspidate to conic teeth (Fig. 4). Posterior face of large dentary teeth convex near its 
base and slightly concave near distal tip. Dentary and premaxillary teeth of inner row 
bearing a central cusp large, slightly curved posteriorly and lateral ones small, placed 
oblique to main axis of central cusp. 

Eye large, longer than snout, little more than two times in head; interorbital 
wide. Third infraorbital not contacting latero-sensory canal of preopercle either ven- 
trally or posteriorly. 

Dorsal fin with iii,8-9 rays (2 ex.= 8); (20 ex. including holotype= 9); distal 
margin of dorsal fin straight, with last unbranched and first branched dorsal fin rays 
longest. First unbranched dorsal fin ray mostly visible in cleared and stained speci- 
mens. Enlarged anal fin with iv-v, 24-29 rays (3 ex.= 24); (11 ex. including holotype= 
25); (7 ex.= 26); (1 ex.= 29). Males with distal margin straight, and females with last 
unbranched and first five or six branched rays lengthened to form a small lobe. Males 
with hooks on last unbranched anal fin ray and posterior branch of first 8-16 branched 
anal fin rays; one pair of hooks per segment. 

Caudal fin with principal rays i,17,i; ventral caudal lobe longer than upper one. 
Pectoral fin with i, 12-14 rays (3 ex.= 12); (13 ex.= 13); (6 ex. including holotype= 14), 



NEW ASTYANAX FROM RIO BERMEJO BASIN 



217 




Fig. 1 
Astyanax latens sp. n., holotype, CI-FML. 3400, male, 44.3 mm SL, Argentina, province of 
Salta, Orân, rio Bermejo basin, arroyo El Oculto. 



one non-type specimen with 11; distal margin straight. Pelvic fin with i,7 rays (one 
specimen with i,6); pelvic fin distal margin slightly rounded. Males bearing hooks in 
all branched pelvic fin rays, usually on posterior branch; sometimes, more than one 
pair of hooks per segment. 

Scales cycloid. Lateral series with 37-38 (8 ex.= 37); (14 ex. including holo- 
type= 38; one non-type specimen with 39) perforated scales. Scales between dorsal fin 
origin and lateral line 7; scales between lateral line and pelvic fin origin 6-7. Scales 
around caudal peduncle 14-15. Scales between tip of supraoccipital spine and base of 
dorsal fin 12-13, usually forming a regular row. One row of scales situated along anal 
fin base, covering base of all unbranched and first twelve to fifteen branched anal fin 
rays. Few scales on caudal fin base. 

Colour in life: Body uniformly silvery, dark on dorsal part of flanks. Lateral 
band inapparent. First humeral spot vertically elongated, narrow, and second one faint. 
Caudal spot visible, occupying complete depth of caudal peduncle. Anal and caudal 
fins orange, becoming redish in specimens recently collected. 

Colour in alcohol preserved specimens: Body uniformly yellowish. First black 
humeral spot vertically elongated, very narrow; second one faint although always 
marked. Chromatophores scattered over body, especially concentrated on dorsal 
portion of flanks, forming a reticulate pattern. In lower half of flanks, chromatophores 
following myosepta; also, chromatophores forming lines perpendicular to anal fin base. 
Lateral band very narrow, formed by small superficial chomatophores and deep 
chromatophores placed as small V, with vertex directed anteriorly. Caudal spot well 
developed, black, triangular or irregular in shape, extending on middle caudal fin rays. 



218 J- M. MIRANDE ET AL. 



Table 1. Astyanax latens sp. n. Morphometries of holotype and 20 paratypes. Minimum, 
maximum, and mean ± standard deviation in brackets. SL is measured in mm. SOC= supra- 
occipital 

males (n=ll) 
43.6-52.0 



51.7-55.3(53.7-1.0) 
42.6 _ 45.8 (46.0 ±1.2) 
57.6 -61.0 (59.5 ±1.2) 
34.9 -38.8 (37.5 ±1.2) 
33.6 - 36.7 (34.8 ± 0.9) 
23.2 - 25.9 (24.9 ± 0.8) 
17.6 -22.0 (20.0 ±1.2) 

18.2 -19.9 (19.0 ±0.5) 

15.6 -18.7 (17.2 ±1.0) 
25.8-28.1 (27.1 ±0.6) 
24.7 -28.3 (26.9 ±1.1) 
35.5 -40.5 (38.3 ±1.5) 
26.2 - 28.9 (27.7 ± 0.9) 
10.2- 11.3 (10.8 ±0.3) 
10.7- 13.2 (11.9 ±0.8) 



38.9 -51.2 (43.9 ±4.0) 
38.8 -41.3 (39.9 ±0.9) 
22.4 -26.4 (23.9 ±1.0) 
35.9 -41.2 (38.8 ±1.4) 
27.8 -32.2 (29.3 ±1.1) 
30.0 -35.2 (33.8 ±1.4) 
Premaxillary+maxillary length 41.8 37.4 -40.8 (39.5 ± 1.1) 36.7 - 41.8 (39.5 ± 1.6) 



Dorsal fin with scattered small chromatophores along fin rays. Adipose fin with few 
chromatophores on its base. Black chromatophores scattered on caudal and anal fins, 
especially on membranes and along ray margins. Pectoral and pelvic fins with few 
chromatophores along ray margins. Dorsum of head dark; chomatophores concentrated 
on premaxilla, many of them on maxilla and upper portion of dentary. A line of black 
chromatophores around eye. 

Osteological characters. In 6 cleared and stained specimens: first branchial arch 
bearing 26-27 gill rakers placed as follow: 9-10 on epibranchial, 1 on cartilage, 14 on 
ceratobranchial, and 2 on hypobranchial. 

Caudal fin with 8-9 dorsal and 7-8 ventral procurrent rays. Thirty five to 37 
vertebrae (percaudal vertebrae 15-16, caudal vertebrae 20-22). Ten or eleven dorsal fin 
pterygiophores between neural spines of vertebrae 10-11 and 18-19; 27-28 anal fin 
pterygiophores between hemal spines of vertebrae 17 and 28-29. First anal fin ptery- 
giophore bearing 4 unbranched anal fin rays. Five or six supraneurals; 10-11 pairs of 
ribs. 





holotype 


females (n=10) 




44,3 




42.3 - 52.5 


SL 








% of standard length 








Predorsal distance 


53.8 


51.9- 


-55.6 (54.4 ±1.1) 


Preventral distance 


42.8 


44.3- 


- 46.3 (45.5 ± 0.8) 


Preanal distance 


59.1 


59.9- 


-63.3 (61.5 ±1.2) 


Body depth 


38.5 


36.1- 


-39.1 (37.3 ±0.9) 


Anal-fin base 


35.7 


32.3- 


-36.6 (33.5 ±1.3) 


Pectoral-fin length 


25.9 


23.8- 


- 26.6 (25.0 ± 0.8) 


Pelvic-fin length 


20.3 


17.6- 


- 20.4 (19.4 ± 0.8) 


Distance between pectoral 








and pelvic fin origins 


18.5 


18.6- 


-21.8(20.1 ±1.0) 


Distance between pelvic 








and anal fin origins 


17.3 


16.6- 


-20.0 (18.4 ±0.9) 


Head length 


27.2 


25.8- 


- 28.5 (26.9 ± 0.7) 


Prepectoral distance 


26.6 


25.8- 


- 27.8 (27.0 ± 0.6) 


Postdorsal distance 


38.3 


36.4- 


-40.6 (38.3 ±1.3) 


Snout tip-SOC posterior tip 


28.6 


26.3- 


- 28.0 (27.2 ± 0.5) 


Caudal peduncle depth 


11.1 


10.0- 


-11.4 (10.6 ±0.5) 


Caudal peduncle length 


11.6 


10.7- 


-12.9 (11.5 ±0.8) 


% of head length 








Peduncle length 


42.6 


38.3- 


- 49.0 (43.0 ± 4.0) 


Peduncle depth 


40.6 


38.0- 


-41.8 (39.4 ±1.3) 


Snout length 


26.4 


22.4- 


- 25.0 (23.6 ± 0.8) 


Eye diameter 


40.3 


38.4- 


-42.1 (40.1 ± 1.3) 


Interorbital width 


29.7 


27.5- 


- 30.5 (29.2 ± 0.9) 


Maxillary length 


34.9 


31.6- 


-36.6 (34.0 ± 1.5) 



NEW ASTYANAX FROM RIO BERMEJO BASIN 



219 




Fig. 2 
Astyanax laiens sp. n., 44.0 mm SL, premaxilla in left internal view. Scale bar = 1 mm. 




Fig. 3 
Astyanax latens sp. n., 44.0 mm SL, maxilla in left internal view. Scale bar = 1 mm. 



Upper edge of dentary almost parallel to lower mandibular margin; dentary 
foramen placed anteriorly and scarcely above level of Meckel cartilage. Cusps of pre- 
maxillary teeth placed forming an almost straight line in ventral view. 

Ventral surface of mesethmoid with lateral and medial processes for vomer 
articulation forming an obtuse angle. Rhinosphenoides well ossified occasionally. 



220 



J. M. MIRANDE ET AL. 




FlG. 4 

Astyanax latens sp. n., 44.0 mm SL, lower jaw in medial view. Scale bar = 1 mm. 



Etymology. The specific epithet latens is a Latin word that means hidden; the 
Spanish name of the type locality also means hidden. 

Distribution. Astyanax latens is known from arroyo El Oculto and the rivers 
Anta Muerta and Blanco, which flow into upper rio Bermejo (Fig. 5). The type locality 
of A. latens, arroyo El Oculto, is a small stream of springs, with clear and slow water, 
muddy bottom and pools more than 1 m deep. The temperature and the caudal of the 
water are constant along the year, contrasting with those of streams and rivers of the 
area under marked seasonality. The place where A. latens was found is usually covered 
by shadow of the circumambient vegetation; many specimens were collected all 
around the year. A lower number of specimens of A. latens was occasionally found in 
the rio Anta Muerta, a tributary on rio Blanco in which other specimens were collected 
also. 



DISCUSSION 

Astyanax latens sp. n. is differentiated from the species of the genus living in 
southern South America by the presence of 28-34 and fin rays in combination with 
other characters. Astyanax pel e gr ini (41-45), A. erythropterus (45), A. correntinus (45), 
and A. allenì (39-41) have higher numer of anal fin rays. Low number of anal fin rays 
is present in A. brachypterygium (16-20), A. cremnobates (18-22), A. gymnogenys 
(21-22), A. scabripinnis paranae (17-23), A. ita (23-28), A. eigenmanniorum (22-26), 
A. ojiara (24-28), A troy a (22-26), A. leonidas (20-25), A. pynandi (21-26), and A. 
paris (24-27). The number of maxillary teeth of A. latens (1-4) is shared with A. paris. 

The pigmentation pattern of A. lineatus (with several bands along the flanks), A. 
laticeps (with horizontally oval humeral spot), and A. marionae (without caudal spot) 
distinguishes them from A. latens wich only has a very narrow lateral band, humeral 
spot vertically elongated, and triangular caudal spot. 



NEW ASTYANAX FROM RIO BERMEJO BASIN 



221 




Fig. 5 
Geographic distribution of Astyanax latens sp. n. in Argentina, Salta, rio Bermejo basin. 1, 
arroyo El Oculto (type locality); 2, rio Anta Muerta; 3, rio Blanco. Scale bar = 10 km. 



Astyanax asuncionensis (30-32) and A. abramis (30-34) have similar number of 
anal fin rays, but they also bear a horizontally elongated humeral spot, and 0- 1 maxil- 
lary tooth. A similar number of anal fin rays than that of A. latens is present in A. cf. 
fasciatus (24-31). However, A. cf.fasciatus has a wide lateral band, only one maxillary 
tooth (vs. 1-4), shorter pectoral fins never reaching pelvic fin origin (vs. pectoral fin tip 
always surpassing pelvic fin origin), and distal tips of premaxillary teeth placed form- 
ing an arch of 180°, with concavity directed anteriorly (vs. premaxillary teeth cusps 
placed forming a line or scarcely curved). Also, A. latens is a smaller species, it occa- 
sionally reaches more than 55 mm SL. 

Astyanax saguazu shares the number of anal fin rays, the number of scales in 
the lateral series, and the disposition of the teeth cusps with A. latens. Nonetheless, A. 
saguazu has a notable lateral band and 19-23 gill rakers on first branchial arch whereas 
A. latens has very narrow lateral band and 26-27 gill rakers on the same arch. Also, A. 
saguazu has a larger eye (41.1-45.5 % vs. 38.4-42.1 % HL), and a shorter predorsal dis- 
tance (46.3-52.9 % vs. 51.7-55-6 % SL), pectoral (21.6-24.7 % vs. 23.2-26.6 % SL) 
and pelvic (16.5-18.8 % vs. 17.6-22.0 % SL) fin lengths, and anal fin base (29.5- 
33.8 % vs. 32.3-36.7 % SL). 

OSTEOLOGICAL CONSIDERATIONS 

Few osteological characters of species included in the genus Astyanax are 
known; most of them concern teeth and jaws. With no doubt, a revision of the genus is 
needed and, as the diagnostic characters of the genus could be plesiomorphic charac- 
ters, a phylogenetic study will be welcomed. 



222 J- M. MIRANDE ET AL. 

In comparison, the teeth of different species of Astyanax bear several characters 
that could help to differentiate them. Astyanax latens shares with A. saguazu and 
Astyanax sp. A, a maximum of five cusps in the teeth, while most species of Astyanax 
examined have the larger teeth with seven cusps or more (e.g., A. cf. fasciatus, A. 
eigenmanniorum, A. troya, A. ojiara, A. asuncionensis, A. ita). Also, the base of the 
inner premaxillary teeth is broader than the distal portion in A. latens, A. saguazu, and 
Astyanax sp. A. whereas some species have premaxillary teeth with broad distal por- 
tions (e.g., A. troya, A. ojiara, A. pynandi, A. giton, A. hastatus following Melo, 2001). 

Most of the Astyanax species examined have cusps of the inner premaxillary 
teeth forming an arch of 180° in ventral view, and leaving an anterior concavity 
(among others, A. lineatus, A. asuncionensis, A. abramis, A. eigenmanniorum, A. troya, 

A. ojiara, A. leonidas, A. cf. fasciatus, A. alleni). Astyanax latens, A. saguazu, and 
Astyanax sp. A have premaxillary cusps arranged in a straight line. 

The crest for insertion of the adductor operculi, in the internal surface of 
opercle, is very small in A. latens whereas the crest is longer, occupying more than half 
of the distance between anterior and posterior opercular margins in other species such 
as A. ojiara, A. ita, A. paris, A. lineatus, or A. asuncionensis. 

Astyanax latens shares some characters with species of different genera. For 
example, the ventral mesethmoides processes for articulation of the vomer are usually 
T-shaped as in Brycon (Weitzman, 1964), Oligosarcus (per. obs.), and other species of 
the genus Astyanax (e.g., A. abramis, A. asuncionensis, A. lineatus, pers. obs.); in 
contrast, the ventrolateral and ventromedial processes of mesethmoides form a slight 
angle (almost Y-shaped) in A. latens, Moenkhausia intermedia, and Acrobrycon 
tarifae. This angle was observed to be more pronunciated in Bryconamericus iheringii, 

B. thomasi and Odontostilbe pequira. 

The postero ventral process of pterotic is present in Brycon (Weitzman, 1964; 
per. obs.), A. abramis, and A. asuncionensis, whereas it is absent in A. latens, most 
species of Astyanax, Moenkhausia intermedia, Acrobrycon tarijae, Bryconamericus 
thomasi, and Odontostilbe pequira. 

All these observations show that detailed revisions of the genus Astyanax and 
other genera of tetragonopterine characiforms of the family Characidae, probably 
closely related as Hyphessobrycon, Moenkhausia, or Hemigrammus, are necessary to 
establish polarity of characters and the intrageneric relationships among them. 

ACKNOWLEDGEMENTS 

Authors thank L. Malabarba and J. Lundberg for loan, exchange or information 
of paratyes of A. eigenmanniorum, Fundación Miguel Lillo and Laboratorio de 
Investigaciones de las Yungas for help in various stages of this paper, Fundación 
Proyungas and Fundación Vida Silvestre Argentina for financial support, and Conicet 
for permanent support. 



NEW ASTYANAX FROM RIO BERMEJO BASIN 223 



REFERENCES 

Almirón, A. E., Azpelicueta, M. de las M., Casciotta, J. R. & Lopez Cazorla, A. 1997. 

Icthyogeographic boundary between the Brazilian and Austral subregions in South 

America, Argentina. Biogeographica 73: 23-30. 
Almirón, A. E., Azpelicueta, M. de las M. & Casciotta, J. R. 2002. Astyanax ita sp. n. - a new 

species from the Rio Iguazü basin, in Argentina (Teleostei, Characiformes, Characidae). 

Zoologische Abhandlungen 52: 3-10. 
Azpelicueta, M. de las M. & Garcia, J. O. 2000. A new species of Astyanax (Characiformes, 

Characidae) from Uruguay river basin in Argentina, with remarks on hook presence in 

Characidae. Revue suisse de Zoologie 107: 245-257. 
Azpelicueta, M. de las M., Casciotta, J. R. & Almirón, A. E. 2002a. Two new species of the 

genus Astyanax (Characiformes, Characidae) from the Parana river basin in Argentina. 

Revue suisse de Zoologie 109: 243-259. 
Azpelicueta, M. de las M., Casciotta, J. R. & Almirón, A. E. 2002b. Astyanax paris: A new 

species from the rio Uruguay basin of Argentina (Characiformes, Characidae). Copeia 

2002 (4): 1052-1056. 
Bertaco, V. & Malabarba, L. R. 2001. Description of two new species of Astyanax (Teleostei: 

Characidae) from headwater streams of Southern Brazil, with comments on the "A. 

scabripinnis species complex". Ichthyological Exploration of Freshwaters 12: 221-234. 
Casciotta, J. R., Almirón, A. E., Bechara, J. A., Roux, J. P. & Ruiz Diaz, F. 2003a. Astyanax 

pynandi sp. n. (Characiformes, Characidae) from Esteros del Ibera wetland, Argentina. 

Revue suisse de Zoologie 110(4): 807-816. 
Casciotta, J. R., Almirón, A. E. & Azpelicueta, M. de las M. 2003b. A new species of 

Astyanax from rio Uruguay basin, Argentina (Characiformes: Characidae). Ichthyo- 
logical Exploration of Freshwaters 14: 329-334. 
Eigenmann, C. H. 1921. The American Characidae. Memoirs of the Museum of Comparative 

Zoology 43: 209-310. 
Eigenmann, C. H. 1927. The American Characidae. Memoirs of the Museum of Comparative 

Zoology 43: 311-428. 
Garutti, V. & Britski, H. A. 2000. Descriçào de urna espécie nova de Astyanax (Teleostei: 

Characidae) da bacia do alto rio Parana e consideraçôes sobre as demais espécies do 

gènero na bacia. Comunicacòes do Museu de Ciências e Tecnologia da PUCRS, série 

Zoologia, Porto Alegre 13: 65-88. 
Géry, J. 1977. Characoids of the world. TFH Publications, Neptune City, 672 pp. 
Melo, F. A. G. 2001. Revisào taxonômica das espécies do gènero Astyanax Baird & Girard, 

1854, (Teleostei: Characiformes: Characidae) da regiào da Serra dos Orgàos. Archivos 

do Museu Nacional, Rio de Janeiro, 59: 1-46. 
Ringuelet, R. A., Aramburu, R. H. & Arämburu, A. A. de. 1967. Los peces argentinos de agua 

dulce. Comisión de Investigaciones Cientificas de la provincia de Buenos Aires, 602 pp. 
Taylor, W. R. & Van Dyke, G. C. 1985. Revised procedures for staining and clearing small fish- 
es and other vertebrates for bone and cartilage study. Cybium 9: 107-119. 
Weitzman, S. H. 1964. The osteology of Brycon meeki, a generalized fish with an osteological 

definition of the family. Stanford Ichthyological Bulletin 8: 1-77. 



Revue suisse de Zoologie 111 (1): 225-238; mars 2004 



Morphology and phylogenetic relationships of the Cyprus racer, 
Hierophis cypriensis, and the systematic status of Coluber gemo- 
nensis gyarosensis Mertens (Reptilia: Squamata: Colubrinae) 

Urs UTIGER 1 & Beat SCHÄTTI 2 

1 Eichstrasse 26, CH-8045 Zürich, Switzerland. E-mail: cypriensis@herpetology.ch 

2 Apartado postal 383, San Miguel de Allende, Gto. 37700, Repüblica Mexicana. 



Morphology and phylogenetic relationships of the Cyprus racer, 
Hierophis cypriensis, and the systematic status of Coluber gemonensis 
gyarosensis Mertens (Reptilia: Squamata: Colubrinae). - External mor- 
phology, skull bones, and hemipenis features of Hierophis cypriensis 
(Schätti) were examined and new lowland findings are reported. Morpho- 
logical and molecular data confirm sister species status of the Cyprus racer 
vis-à-vis the western Mediterranean racers H. gemonensis (Laurenti) and H. 
viridiflavus (Lacépède). Apart from the Cyclades viper, Macrovipera 
schweizeri (Werner), H. cypriensis is the only endemic Mediterranean in- 
sular snake species. Coluber gemonensis gyarosensis Mertens from Gyaros 
Island (Cyclades) is a junior synonym of H. viridiflavus. This Aegean 
population, highly isolated from the continuous range of the species, was 
most probably introduced by human activity in historical times. Molecular 
data suggest validity of H. viridiflavus carbonarius (Bonaparte) for mela- 
notic populations of the European whip snake including the Gyaros racer. 

Key-words: Hierophis cypriensis - morphology - zoogeography - Cyprus - 
Coluber gemonensis gyarosensis - Hierophis viridiflavus - phylogeny - 
mtDNA (COI, 12S rRNA). 

INTRODUCTION 

Since the description of Coluber cypriensis Schätti, 1985, the systematically 
difficult racer taxon Coluber auct. underwent drastic taxonomic changes. Coluber 
Linnaeus, 1758 is a purely Nearctic genus. Old World racers and whip snakes formerly 
parading under Coluber auct. (e.g., Schätti & Wilson, 1986) belong to at least four 
different genera, viz., Hemerophis Schätti & Utiger, Hemorrhois Boie, Hierophis 
Fitzinger, and Platyceps Blyth (Schätti, 1986, 1987; Schätti & Utiger, 2001). 

In an unpublished thesis, Schätti (1988) briefly discussed presumed phyloge- 
netic affinities of the Cyprus racer (or whip snake), stating that the in situ hemipenis 
shows strong resemblance to Hierophis spp. Schätti & Sigg (1989b) formally referred 
Coluber cypriensis to the predominantly southern Palaearctic genus Hierophis 
Fitzinger in Bonaparte, 1834 with the type species H. viridiflavus (Lacépède). Böhme 
& Wiedl (1994) figured an everted hemipenis of//, cypriensis and considered "[...] the 



Manuscript accepted 28.07.2003 



226 U. UTIGER & B. SCHÄTTI 

overall phenetic similarity being greatest with C.[oluber] (H.) gemonensis" (Laurenti). 
However, these authors continued to refer the Cyprus racer to Coluber auct., and not 
to "the Hierophis section of Coluber". 

Memphis Fitzinger is made up of two European species (viz., H. gemonensis 
and H. viridiflavus) as well as H. caspius (Gmelin) from the Balkans, the eastern 
Mediterranean H. jugularis (L.) and H. schmidti (Nikolskij), the endemic H. 
cypriensis, and H. spinalis (Peters) from Central Asia to Korea (Schätti 1987, 1993; 
Schätti & Utiger, 2001). 

The systematic status of Coluber gemonensis gyarosensis Mertens, 1968 from 
Gyaros Island (Aegean Sea, Cyclades) is controversial (see Böhme, 1993). Schätti 
(1988) stated that this population cannot be separated from the western Mediterranean 
Hierophis viridiflavus as far as external morphology is concerned. 

This paper describes external morphological features and osteological charac- 
ters of Hierophis cypriensis, compiles the known distribution of this insular endemic, 
and provides sequence data for two mitochondrial genes, COI and 12S rRNA. The 
objective of the present study is to clarify the phylogenetic relationships of the Cyprus 
racer as well as the validity and systematic status of the racer population from Gyaros 
Island. 

MATERIAL AND METHODS 

The material examined for the purpose of this study is deposited in The Natural 
History Museum (formerly British Museum, Natural History), London (BMNH), 
Department of Zoology, The Hebrew University, Jerusalem (HUJ), Muséum d'histoire 
naturelle, Geneva (MHNG), Naturhistorisches Museum, Vienna (NMW), and Zoo- 
logisches Museum und Forschungsinstitut Alexander Koenig, Bonn (ZFMK). The 
specimens are: BMNH 87.9.27.25 (<?), "Cyprus, Lord Lilford" (leg. Guiilemard, 
paratype). HUJ 3331 (9), Kykko Monastery (Panagia tou Kykko), approx. 34°58'N 
32°44'E, c. l'IOO m a.s.l., Nicosia District, leg. Mavromoustakis; HUJ 3786 (S), 
"Polemedia Hills" (Polemidia), c. 200 m (600 ft.), Limassol, leg. Mavromoustakis. 
MHNG 2206.30 (9), between Arakapâs and Kalokhorio, approx. 34°51'N 33°06'E, 
470 m, Limassol, B. Schätti, 14 April 1983 (holotype); MHNG 2453.60 (Ó*), between 
Kannaviou and Stavros tis Psokas, c. 750 m, Paphos, W. Teschner, 11 May 1989; MH- 
NG 2618.99 (cT), between Germasógeia (Yermasoyia) and Akroünta, approx. 34°45'N 
33°06'E, c. 75 m, Limassol, B. Schätti, 27 May 2001 (roadkill). NMW 15185.1 (9), 
"Platraes, Troodos", Limassol, 1'400 m, F Werner, May 1935 (paratype). ZFMK 
48877 (c?), 1 km northwest of Pano Platraes, Limassol, K. Osenegg & S. 
Demetropoulos, 26 June 1988; ZFMK 51283 (S), Pano Panagia (Panayia), l'OOO m, 
Paphos, H. Wiedl, October 1989; ZFMK 51881 (<?), Mt. Triphylos, approx. 34°59'N 
32°40'E, Paphos, H. Wiedl, September 1990. Furthermore, MHNG 2618.100, consist- 
ing of a portion of a shed skin from the upper Livadi Priver area at approx. 35°04'N 
32°36'E (c. 600 m) in Paphos District, was used for verification of dorsal scale 
characters (apical pits, transverse position of reduction levels) and molecular studies 
(12S rDNA, see footnote 3). 

External morphological data (pholidosis, length, dorsal colour pattern) are 
based on this material (ten complete specimens) and two male roadkills from the 



THE CYPRUS RACER 227 

vicinity of Pano Platraes (Troodos road, "ca. 5 km südlich Troodos") and Kakopetria 
(leg. S. Demetropoulos 1988) reported by Osenegg (1989: Tb. 24). Osteological 
characters were ascertained in MHNG 2453.60, MHNG 2618.99, and ZFMK 51881; 
additional teeth counts (right dentigerous bones) are based on HUJ 3331 (dentary, 
maxilla, palatinum), HUJ 3786 (dentary, maxilla), MHNG 2206.30 (maxilla), and 
NMW 15185.1 (maxilla). 

Methods and definitions of terms used in the descriptive part and comparative 
section of this text are explained in Schätti (1987, 1988). The position of bilateral 
dorsal scale row reductions along the trunk is given in terms of absolute numbers of 
ventrals and as a percentage thereof (%ven), and calculated on the basis of the average 
of right and left side counts. The length of the hemipenis and the M. retractor penis 
magnus are expressed in absolute numbers of subcaudals and as a percentage thereof 
(%sub). Vertebra measurements used in this study are the length of the centrum (lc), 
length of neural crest (nc), least width of neural arch (wn), and the width across 
prezygapophyses (wp). 

DNA sequences of two mitochondrial genes, cytochrome oxidase I (COI), and 
the small ribosomal subunit (12S rRNA), of 13 Palearctic and Afrotropical racers as 
well as the Oriental racer Coelognathus flavolineatus (Schlegel) (see Utiger, 2002) are 
from an earlier study (Schätti & Utiger, 2001) and listed in the Appendix. Eight tissue 
samples belonging to four different taxa and frozen at -80°C are new: Hierophis 
cypriensis (MHNG 2618.99, liver), H. gyarosensis (MHNG 2402.91 and 2401.92, 
Gyaros Island, Cyclades; muscle), H. spinalis (MHNG 2443.5, Wonju, Kangwon, S 
Korea; muscle; MVZ 211019, N shore Huang He, Shapotou, Yinnan Prefecture, China; 
liver), and H. viridiflavus (MHNG 2402.11, Chizé, Deux-Sèvres, France; muscle; MH- 
NG 2402.66, Finaita, Sicily, Italy; muscle; MHNG 2402.83, Krk Island, Croatia; 
muscle). The corresponding Genbank accession numbers are AY 541502-541512. 

DNA isolation, purification, polymerase chain reaction (PCR) and sequencing 
procedures were performed as described in Schätti & Utiger (2001). 

Because of considerable constraints in laboratory facilities, only one gene 
region, 12S rDNA, was analysed for the new specimens of Hierophis viridiflavus. 
These sequences were added to an alignment file consisting of two partially sequenced 
genes (COI and 12S rRNA) and a selection of 14 species from an earlier study (Schätti 
& Utiger, 2001). Where sequence information on the COI gene region was not 
available, characters were coded as "missing". 

All phylogenetic analyses were performed with PAUP* version 4.0b 10 for Mac 
(Swofford, 1998). Neighbour joining (NJ, Saitou & Nei, 1987) calculates gene distances 
based on the number of mutation events between all pairs of taxa. Under the present cir- 
cumstances, the calculation of genetic distances between specimens with complete and 
incomplete sequence data is inappropriate because of different mutation properties 
within each gene region. In this case, maximum parsimony (MP) is a good alternative 
to NJ as it reconstructs a tree based on both gene regions. In addition, MP counts the 
mutation steps that occurred in the 12S gene region between specimens with complete 
sequence data and those lacking information on COL This gives a qualitatively satis- 
fying result but deals with an underestimate of the intraspecific sequence variability of 
Hierophis viridiflavus compared to the branch lengths of the overall topology. 



228 



U. UTIGER & B. SCHÄTTI 



Gaps in the 12S sequence were treated as fifth character state. After a first run 
with heuristic search and tree-bisection reconnection (TBR) branch swapping, charac- 
ters were weighted with the rescaled consistency index (RC, Farris, 1989) and a second 
heuristic search was performed. The weighting procedure was repeated once; sub- 
sequent weighting procedures did not alter the parameters of the resulting tree. 
Nonparametric bootstrap calculations (Felsenstein, 1985) with 1000 replicates were 
performed for unweighted as well as weighted characters. 

The justification for including gaps and weighting characters is given in Utiger 
et al. (2002). Essentially, gaps supply additional phylogenetic information whereas 
weighting procedures favour characters which bear phylogenetic content. Using these 
measures, tree dependent (RC) and independent phylogenetic signals (tRASA, Lyons- 
Weiler, 2001) intensify (Tb. 1). 



Table 1. Sequence properties and tree reconstruction parameters. 



gaps: missing gaps: 5th character state 



unweighted MP unweighted MP weighted MP 



Length of sequence alignment (COI/12S) 1016 (407/609) 1016 (407/609) 1016 (407/609) 

Total of variable characters (COI/12S) 322(142/180) 333(142/191) 333(142/191) 

- parsimony-informative (COI/12S) 225(116/109) 232(116/116) 232(116/116) 

Number of most parsimonious trees 12 1 

Tree length 918 956 267.38 

Rescaled consistency index (RC) 0.232 0.242 0.548 



RESULTS 



Hierophis cypriensis (Schätti, 1985) 

Morphology 

Loreal and preocular single, two postoculars. Eight supraoculars (nine on left 
side of MHNG 2453.60), fourth and fifth (fifth and sixth) in contact with eye, sixth and 
seventh (seventh and eighth) largest. One anterior subocular situated between the third 
and fourth supralabial. One (holotype) or two anterior temporals, two or three scales in 
second row; lower anterior temporal vertically divided in NMW 15185.1. Ten subla- 
bials, anterior four in contact with first inframaxillary, sixth largest. Anterior infra- 
maxillaries slightly shorter than and more or less the same width as posterior pair, 
which is cranially separated by one row of small scales and caudally by two to four 
rows. 

Ventrals in males 194-202 (n=8), females 204-206 (3), subcaudals 123-133 (6) 
and 121-130 (3), respectively; anal plate divided. Dorsal scales smooth, with two 
apical pits; arranged in 17 longitudinal rows at the level of the tenth ventral and at 
midbody, 13 immediately prior to the vent. First reduction involving third and fourth 
row between ventrals 121 and 133 (62-68%ven) in males and 126-132 (61-64%ven) in 



THE CYPRUS RACER 229 

females; second reduction (rows 6+7 or 7+8) situated between ventrals 136-159 (70- 
79%ven, 6 S) and 133-156 (66-73%ven, 9 9) D. 

Total length in males 93,7 (62,0 + 31,7) cm (MHNG 2618.99), females approx. 
92 (62 + 30) cm (NMW 15185.1); tail/body ratio 0.48-0.52 (S <?) and 0.45-0.48 ( 9 9 ). 
Presumed maximum length probably not exceeding 120 cm. 

Dorsal colouration in adults dark olive to anthracitic. Supraocular with a light 
streak running from the posterior border of the eye toward the frontal, and often also 
present along the lateral edge of the supraocular. Irregular light markings (dots and 
streaks) on the frontal and parietals. Supralabials mostly light except upper borders. 
Preoculars and postoculars yellowish white. Posterior temporal region with light dots. 
Throat pale yellow or off-white, irregularly mottled with fine greyish or sometimes 
black dots. Holotype with caudally converging dorsolateral nuchal stripes (PI. 1). 
Anterior portion of neck usually somewhat lightened laterally. Forebody dorsally with 
fine white or creamish transverse bands. Posterior third of trunk and tail uniformly 
greyish olive (scales with lighter centres). Median part of venter pigmented with dark 
spots (less so anteriorly), with a fine whitish line along the ventral edges; lateral region 
invaded by dorsal colouration. ZFMK 48877 virtually black throughout the posterior 
three quarters of the underside. An irregular black line running along the median edges 
of the subcaudals. 

The dorsal colour pattern is subject to ontogenetic change, i.e., juveniles (beige 
above) and subadult specimens (dorsum olive) have dark transverse blotches on the an- 
terior two thirds of the body and an orange iridescent venter (H. Wiedl, pers. comm.). 
Illustrations of this species are found, for instance, in Schätti (1985), Gruber (1989), 
Osenegg (1989), and Schätti & Sigg (1989b). 

Maxillary teeth 14-16 + 2 (n=6), with a distinct diastema and the posterior teeth 
enlarged (last offset laterad). Counts for other dentigerous bones 10-11 (palatinum, 
n=3), 17-20 (pterygoid, 2), and 19-20 (dentary, 4). Palatine processus of maxilla 
comparatively long, lateral border more or less straight and ending in a distinct tip; 
posterior processus poorly developed. Lateral processus of palatinum stout, its 
posterior tip reaching insertion of choanal processus. Anterior portion of pterygoid 
narrow and slightly constricted behind the simple lateral processus. Parasphenoid not 
constricted at the basis, middle posterior border of basisphenoid somewhat convex 
(Fig. 1). Midbody vertebra ratios (n=2) are 1.38-1.58 (lc/wn), 0.74-0.80 (lc/wp), and 
1.11-1.28 (nc/wn). 

Hemipenis with a basal hook (Böhme & Wiedl, 1994: Fig. 6), distally followed 
by a series of spines (less numerous on sulcate side), apex distinctly calyculate, bor- 
ders of depressions spinose on asulcate side. Tip of apex in situ reaching to subcaudal 
9-10 (7-8%sub, n=4), M. retractor penis magnus inserting at subcaudal 30-32 (24- 
25%sub, n=6). 



*) Osenegg (1989: Table 24) gave 121 subcaudals for a male specimen (no. III). This figure as 
well as the reduction level to 15 dorsal scale rows are not included in the data because, strangely 
enough, subcaudal and reduction data for specimens III and V-VI are identical. Definitely, the 
reduction pattern of two specimens of the type series, i.e., MHNG 2206.30 and NMW 15185.1 
("Osenegg, 1989: nos. V-VI) is erroneous. 



230 



U. UTIGER & B. SCHÄTTI 




Plate 1 
Female holotype of Hierophis cypriensis (MHNG 2206.30). 

Distribution and Ecology 

The Cyprus racer is reported from Limassol District, southwestern Nicosia 
(Lefkosa), and the Stavros woods in eastern Paphos, i.e., the Troodos massif and its 
foothills in western Cyprus (Fig. 3) 2 \ 

Most records are from pine forest with abundant coppice between 400 to 1'700 
m a.s.l. near Pano Platraes (H. Wiedl, pers. coram.). The Cyprus racer also occurs at 
lower elevations, viz. below 250 m as exemplified by specimens collected in the im- 
mediate hinterland of Limassol (HUJ 3331 and 3786, MHNG 2618.99). 

Based on the limited number of preserved specimens and field observations, 
Hierophis cypriensis might appear to be either an uncommon, secretive, or locally re- 
stricted species. However, the Cyprus racer seems to be fairly common at least in the 
Xyliatos dam area (c. 550 m) and the Lagoudhera valley situated on the northern 
Troodos foothills at approx. 35°00'N 33°02'E in Nicosia District. There, from a total 
of 58 field observations during three years, 47 individuals found in the immediate sur- 
roundings of the dam were usually observed between 10 a.m. and 3 p.m., mostly from 
July to September, with a single record 50 m from the water in March (Blosat, 1998). 

Virtually all specimens housed in scientific collections and many field observa- 
tions are from habitats with dense bushy vegetation close to creeks and rivers. In many 



2 ' Apart from records shown on Böhme & Wiedl's (1994: Fig. 5) map, Wiedl found Hierophis 
cypriensis "together with Telescopus fallax" at Kâmpos and "Chalkistra" (Tsakistra, 
35°01'25"N 32°43'27"E) in western Nicosia District (Böhme & Wiedl, 1994: 40), a few kilo- 
metres roughly north of Kykko Monastery on the Agios Mamas road. 



THE CYPRUS RACER 



231 







Fig. 1 
Skull bones of Hierophis cypriensis: Left maxilla (A), right palatinum (B), right pterygoid (C), 
and basisphenoid (D, parasphenoid broken, reconstructed). Scale equals 2 mm. Drawings by 
Corinne Charvet (A: MHNG 2453.60) and Beatrice Naef (B-D: ZFMK 51881). 



of these places, Hierophis cypriensis is sympatric with Macrovipera (or Daboia) 
lebetina (L.). In the Xyliatos dam area, the Cyprus racer is found in the same habitat 
as Natrix natrix cypriaca (Hecht). 

Hierophis cypriensis preferably feeds on smaller species of lizards, in particular 
Ablepharus kitaibelii (Bibron & Bory) and Ophisops elegans (Ménétries), and green 



232 



U. UTIGER & B. SCHÄTTI 




Fig. 2 
Hierophis cypriensis: Lateral view of right hemipenis of MHNG 2618.99 (roadkill). Photograph 
by Claude Ratton. 

frogs, i.e., Rana cf. bedriagae Camerario (see Plötner et al., 2001). Blosat (1998) 
observed one specimen that captured a "Lacertide", probably Lacerta laevis troodica 
Werner, and an incidence of ophiophagy (Natrix natrix cypriaca). The holotype regur- 
gitated a centipede. In captivity, H. cypriensis also preys on mammals, i.e., small mice 
(personal observation). 

Contrary to most congeneric species, Hierophis cypriensis is a placid snake that 
does not bite upon collecting. In one instance, the Cyprus racer was found feigning 
death (W. Teschner, in litt.). Since this was observed on a road, it cannot be ruled out 
that a car had driven over the specimen before being encountered by the correspondent. 



THE CYPRUS RACER 



233 




Fig. 3 
Known distribution of Hierophis cypriensis. Solid circles denote examined specimens with pre- 
cise locality data and MHNG 2618.100 from the upper Livadi River, Paphos (see Material and 
Methods and footnote 3). Open symbols correspond to four localities in the central Troodos 
(Kakopetria, Platania, and "ca. 5 km südlich Troodos") and eastern Paphos District (Stavros tis 
Psokas) cited in Osenegg (1989: Fig. 44) as well as the Xyliatos dam area, an unspecified sta- 
tion near the type locality (Böhme & Wiedl, 1994: Fig. 5), and Kampos and Tsakistra (see foot- 
note 2). Elevations from 400 m to 1'200 m hatched, areas over 1'200 m in the central Troodos 
blank. Map partly redrawn from Böhme & Wiedl (1994). 



Zoogeography and relationships 

Böhme & Wiedl (1994) presumed that Hierophis cypriensis had been "existing 
much longer on Cyprus than the rest of the herpetofauna." Arguing that "snakes are 
generally primarily absent from oceanic islands", these authors though that "the case 
of Coluber cypriensis in particular would corroborate" the assumption of a land bridge. 
This alludes to Haas (1952) who postulated a terrestrial connection across the Golf of 
Iskenderun in the Pliocene or early Quaternary (see, e.g., Schmidt, 1960; Spitzen- 
berger, 1979). However, there is no geological evidence for a land bridge during that 
period. Rather, Cyprus is isolated from the surrounding mainland since the Upper 
Miocene salinity crisis of the Mediterranean (Hsii, 1972) as assumed by Schätti & Sigg 
(1989a). 

Hierophis cypriensis is most similar to H. gemonensis and H. viridiflavus with 
respect to certain osteological features (Fig. 1) as, for instance, the comparatively long 
palatine processus of the maxilla (also found in H. jugularis but not in H. caspius). H. 
cypriensis and H. gemonensis have more maxillary teeth (16-19 vs. 12-16) than the 
remaining species of this genus. These two species as well as H. spinalis (in prep.) 
possess an enlarged spine (hook) at the basis of the hemipenis. 



234 



U. UTIGER & B. SCHÄTTI 



Based on morphological evidence and preliminary molecular data (mtDNA 
sequences), Schätti & Utiger (2001: 934, 939) considered Hierophis cypriensis to be 
"most closely related to congeneric European species" 3 ). The present results (Fig. 4) 
confirm sister species status of H. cypriensis vis-à-vis the western Mediterranean H. 
viridiflavus and H. gemonensis, viz. the Balkans and European whip snake. 

Hierophis caspius 
Hierophis schmidti 
Hierophis jugularis 
Eirenis modestus 

Hierophis gemonensis 




32 



76 



100 



57 



59 



Hierophis viridiflavus carbonarius 

(Gyaros) 

Hierophis viridiflavus carbonarius* 

(Gyaros) 

Hierophis viridiflavus carbonarius * 

(Dalmatia) 

Hierophis viridiflavus carbonarius* 

(Sicily) 



Hierophis viridiflavus viridiflavus 

(Sardinia) 

Hierophis viridiflavus viridiflavus ' 

(France) 

— Hierophis cypriensis 



Hierophis spinalis 



93 



- Platyceps ventromaculatus 

Platyceps collaris 

Platyceps najadum 



■ Spalerosophis diadema 
Hemorrhois hippocrepis 



-Hemorrhois ravergieri 
■ Hemerophis socotrae 



Coelognathus flavolineatus 
5 changes 



Fig. 4 
Most parsimonious tree from a weighted MP analysis of two partial gene sequences, COI and 
12S rDNA. Numbers above branches indicate bootstrap values from an unweighted, those below 
from a weighted MP analysis. Asterisks denote specimens lacking COI sequence data (see also 
footnote 3). 



3) Schätti & Utiger (2001) examined a fragment of the 12S rDNA subunit of MHNG 2618.100 
(epidermal skin, see Material and Methods). The sequence shows only small difference (2 out of 
592 base pairs) vis-à-vis MHNG 2618.99. The specimen was not used for the phylogenetic 
reconduction because the lack of the COI partition significantly lowered the resolution of the 
tree topology. 



THE CYPRUS RACER 235 

Molecular data and considerable morphological divergence of Hierophis 
cypriensis (e.g., dorsal colour pattern, paravertebral reductions, processi of dentigerous 
bones, general shape of basisphenoid) from congeneric, and in particular the most 
closely related western Mediterranean, species support the theory of the isolation of 
Cyprus for the last approx. 5,2 my. 

Apart from Macrovipera schweizeri (Werner) from the Cyclades, Hierophis 
cypriensis is the only endemic Mediterranean insular snake species. 

Hierophis viridiflavus (Lacépède, 1789) 

Coluber gemonensis gyarosensis Mertens, 1968 syn. nov. 

The systematic status and specific allocation of Coluber gemonensis gyaro- 
sensis Mertens, 1968 have been a matter of controversy (see Introduction). Specific 
status (Schätti, 1988) was accorded based on morphological differences vis-à-vis two 
whip snake species {Hierophis caspius, H. gemonensis) distributed in the Aegean 
region. Böhme (1993) considered the melanotic taxon from Gyaros to be the rarest 
European snake species. 

Racers from Gyaros most strikingly resemble the European whip snake, 
Hierophis viridiflavus, in general appearance, habitus, and morphology (Schätti, 1988). 
In fact, the insular taxon cannot be distinguished from H. viridiflavus and molecular 
data (Fig. 4) clearly demonstrate that Coluber gemonensis gyarosensis Mertens, 1968 
is a junior synonym of//, viridiflavus (Lacépède, 1789). 

The western Mediterranean Hierophis viridiflavus has a continuous range from 
the Pyrenees region (NE Spain) to northern France, Luxembourg, and western 
Switzerland as well as throughout the Appenine Peninsula, Ticino, and eastward to 
northern Dalmatia. The species is widespread in the Tyrrhenian Sea, found on Sicily 
and many surrounding islands and islets, Malta, Gozo, and in the Adriatic Sea (Krk, 
Tremiti Islands). The isolated Aegean occurrence on Gyaros is most probably due to 
human introduction in historical times. To conclude from our molecular data (Fig. 4), 
H. viridiflavus was most likely transported to Gyaros from the northern Adriatic region. 

Gyaros (Gioura, Yioura), nowadays known as Yiaros or Nisos, is a mountainous 
island (up to 490 m a.s.l. in the central part) of ca. 20 km 2 at 37°37'N 24°43'E in the 
Cyclades, roughly 15 km northwest of Syros about halfway between Kea and Tinos. 
From the beginning of our era until the Middle Ages, the island was a haven for 
marauding corsairs who attacked merchant ships throughout the Aegean. During the 
Roman Imperial epoch, Gyaros served as a place of exile. 

All surrounding larger islands (viz., Andros, Kea, Kythnos, Syros, and Tinos) 
are inhabited by Hierophis caspius, an eastern Mediterranean species widespread in the 
Aegean Sea and found on all larger Cyclade islands surrounding Gyaros (e.g., Schätti, 
1988; Szczerbak & Böhme, 1993). The absence of the Caspian whip snake from 
Gyaros may have favoured the colonization of this island by H. viridiflavus. 

External morphological characters of Hierophis viridiflavus analysed by Schätti 
& Vanni (1986) hardly allow a distinction between typical and melanotic populations 
commonly referred to H. v. carbonarius (Bonaparte). These authors considered this 
phenotype as a colour morph of the European whip snake, H. viridiflavus. Based on 



236 U. UTIGER & B. SCHÄTTI 



molecular data, our limited sample shows genetic separation of typically coloured 
specimens (France, Sardinia) and melanotic individuals from Dalmatia (Krk), Sicily, 
and Gyaros Island. Pending studies using specimens from further regions, this result 
may justify the revalidation of H. v. carbonarius for populations from northeastern 
Italy and Dalmatia, the southern Appenine Peninsula including islands such as Sicily 
and Malta, and the isolated Gyaros population. 

ACKNOWLEDGEMENTS 

The junior author expresses his gratitude to Nicolas E. Arnold (London), 
Wolfgang Böhme (Bonn), Franz Tiedemann (Vienna), and Yehudah L. Werner (Jeru- 
salem) for approving the loans of specimens in their respective care. Theodore J. 
Papenfuss (Berkeley, MIZ) and J. Robert Macey (Walnut Creek) supplied tissue 
samples. Lara Modolo, Wolfgang Scheffrahn, Claudia Zebib-Brunner (all Zurich), and 
Jennifer Pastorini (New York) supported the laboratory work, and Vincent Ziswiler and 
Paul Ward (Zurich) provided the facilities. A most special thank you to Corinne 
Charvet (Geneva) for technical assistance. We feel obliged to Walter Teschner 
(Wuppertal) and "Snake George" Hansjörg Wiedl (Paphos) for field observations on 
the Cyprus racer. Regula Böniger, Claudia Fischer, Barbara Straub (all Zurich) helped 
with bibliographical research, and Beatrice Naef (Geneva) prepared some of the 
drawings. 

REFERENCES 

Blosat, B. 1998. Morphologie, Aut- und Populationsökologie einer Reliktpopulation der zypri- 
otischen Ringelnatter, Natrix natrix cypriaca (Hecht, 1930). Bonn, Inaugural-Disser- 
ation, 174 pp. 

Böhme, W. 1993. Coluber gyarosensis Mertens, 1968 - Gyaros-Pfeilnatter (pp. 111-114). In: 
Böhme, W. (ed.). Handbuch der Reptilien und Amphibien Europas, Band 3/1. Wiesbaden, 
Aula-Verlag, 479 pp. 

Böhme, W. & Wœdl, H. 1994. Status and zoogeography of the herpetofauna of Cyprus, with 
taxonomic and natural history notes on selected species (genera Rana, Coluber, Natrix, 
Vipera). Zoology in the Middle East 10: 31-52. 

Bonaparte, C. L. 1834. Coluber leopardinus [4 unnumbered pages]. In: Iconografia della Fauna 
Italica per le quattro classi degli animali vertebrati [1832-1841]. IL Amfibi [Fase. VII, 
punt. 38]. Roma, Salvinoci. 

Farris, J. S. 1989. The retention index and the rescaled consistency index. Cladistics 5: 417-419. 

Felsenstein, J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. 
Evolution 39: 783-791. 

Gruber, U. 1989. Die Schlangen Europas und rund ums Mittelmeer. Kosmos-Namrführer. 
Stuttgart, Franckh 'sehe Verlagsbuchhandlung, 248 pp. 

Haas, G. 1952. Remarks on the origin of the herpetofauna of Palestine. Revue de la Faculté des 
Sciences de l'Université d'Istanbul (B) 17 (2): 95-105. 

Hsü, K. 1972. When the Mediterranean dried up. Scientific American 227: 27-36. 

Lacépède, B. G. E., Comte de 1789. Histoire Naturelle des Quadrupèdes ovipares et des 
Serpens. Vol. II. Paris, Hôtel de Thou, 9 + 20 +144 + 474 pp. 

Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, 
species, cum characteribus, differentiis, synonymis, locis. Vol. I (editio decima, refor- 
mata). Holmiae [Stockholm], L. Salvii, TV + 823 [+ 1] pp. 

Lyons-Weiler, J. 2001. RASA 3.0 Turbo for the Macintosh. http://bio.uml.edu/LW/RASA.html 



THE CYPRUS RACER 237 



Mertens, R. 1968. Eine schwarze Zornnatter von den Cycladen: Coluber gemonensis gyarosen- 

sis n. subsp. Senckenbergiana biologica 49 (3/4): 181-189. 
Osenegg, K. 1989. Die Amphibien und Reptilien der Insel Zypern. Bonn, Diplomarbeit, 

Friedrich-Wilhelms Universität, 200 pp. 
Plötner, J., Ohst, T., Böhme, W. & Schreiber, R. 2001. Divergence in mitochondrial DNA of 

Near Eastern water frogs with special reference to the systematic status of Cypriote and 

Anatolian populations (Anura, Ranidae). Amphibia-Reptilia 22 (4): 397-412. 
Sattou, N. & Nei, M. 1987. The neighbor-joining method: a new method for reconstructing phy- 

logenetic trees. Molecular Biology and Evolution 4: 406-425. 
Schätti, B. 1985. Eine neue Zornnatter aus Zypern, Coluber cypriensis n. sp. Revue suisse de 

Zoologie 92 (2): 471-477. 
Schatti, B. 1986. Morphological evidence for a partition of the snake genus Coluber (pp. 235- 

238). In: Rocek, Z. (ed.). Studies in Herpetology (Proceedings of the European 

Herpetological Meeting, 1985). Prague, Charles University, XXVIII + 754 pp. 
Schatti, B. 1987. The phylogenetic significance of morphological characters in the Holarctic 

racers of the genus Coluber (Reptilia, Serpentes). Amphibia-Reptilia 8 (4): 401-415. 
Schatti, B. 1988. Systematik und Evolution der Schlangengattung Hierophis Fitzinger, 1843. 

Zürich, Inaugural-Dissertation, 50 pp. 
Schatti, B. 1993. Coluber Linnaeus, 1758 - Zornnattern (pp. 69-71). In: Böhme, W. (ed.). 

Handbuch der Reptilien und Amphibien Europas, Band 3/1. Wiesbaden, Aula-Verlag, 

479 pp. 
Schätti, B. & Sigg, H. 1989a. Die Herpetofauna der Insel Zypern. 1. Die herpetologische 

Erforschung und Amphibien. Herpetofauna (Weinstadt) 11 (61): 9-18. 
Schätti, B. & Sigg, H. 1989b. Die Herpetofauna der Insel Zypern. 2. Schildkröten, Echsen und 

Schlangen. Herpetofauna (Weinstadt) 11 (62): 17-26. 
Schätti, B. & Utiger, U. 2001. Hemerophis, a new genus for Zamenis socotrae Günther, and a 

contribution to the phylogeny of Old World racers, whip snakes, and related genera 

(Reptilia: Squamata: Colubrinae). Revue suisse de Zoologie 108 (4): 919-948. 
Schätti, B. & Vanni, S. 1986. Intraspecific variation in Coluber viridiflavus Lacépède, 1979, 

and validity of its subspecies (Reptilia, Serpentes, Colubridae). Revue suisse de Zoologie 

93 (1): 219-232. 
Schätti, B. & Wilson, L. D. 1986. Coluber. Catalogue of American Amphibians and Reptiles 

399: 1-4. 
Schmidt, W. F. 1960. Zur Struktur und Tektonik der Insel Cypern. Geologische Rundschau 50: 

375-395. 
Spitzenberger, F. 1979. Die Säugetierfauna Zyperns. Teil II: Chiroptera, Lagomorpha, 

Carnivora und Artiodactyla. Annalen des Naturhistorischen Museums Wien 82: 439-465. 
Swofford, D. L. 1998. PAUP*: phylogenetic analysis using parsimony (and other methods). 

Version 4.0b 10 for Mac. Sunderland, Sinauer. 
Szczerbak, N. N. & Böhme, W 1993. Coluber caspius Gmelin, 1789 - Kaspische Pfeilnatter 

oder Springnatter (pp. 83-96). In: Böhme, W. (ed.). Handbuch der Reptilien und 

Amphibien Europas, Band 3/1. Wiesbaden, Aula-Verlag, 479 pp. 
Utiger, U. 2002. Molecular phylogeny of the Oriental colubrid genus Coelognathus Fitzinger. 

1843, and a contribution to the systematics of Holarctic racers and ratsnakes (Reptilia: 

Squamata: Serpentes). University of Zürich, Ph. D. thesis [part.], 73 pp. 
Utiger, U, Helfenberger, N., Schätti, B., Schmidt, C, Ruf, M. & Ziswtler, V 2002. 

Molecular systematics and phylogeny of Old and New World ratsnakes, Elaphe auct., 

and related genera (Reptilia, Squamata, Colubridae). Russian Journal of Herpetology 9 

(2): 105-124. 



238 U. UTIGER & B. SCHÄTTI 



APPENDIX. Genbank numbers of specimens examined by Schätti & Utiger (2001) used in this 
study (COI/12S). Coelognathus flavolineatus (AY039200/AY039162); Eirenis modestus 
(AY039181/AY039143); Hemerophis socotrae (AY039167/ AY039129); Hemorrhois hippo- 
crepis (AY039194/AY039156), H. ravergieri (AY039193/AY039155); Hierophis caspius 
(AY039164/AY039126), H. gemonensis (AY039183/AY039145), H. jugularis (AY039188/ 
AY039150), H. schmidti (AY039197/AY039159), H. viridiflavus (AY039175/AY039137); Pla- 
tyceps collaris (AY039195/AY039157), P. najadum (AY039179/AY039141), P. ventromacu- 
latus (AY039172/AY039134); Spalerosophis diadema (AY039182/AY039144). 



REVUE SUISSE DE ZOOLOGIE 
Tome 111 — Fascicule 1 

Pages 

James, Samuel W., Hong, Yong & Kim, Tae Heung. New earthworms of 
Pheretima and Pithemera (Oligochaeta: Megascolecidae) from Mt. 
Arayat, Luzon Island, Philippines 3-10 

Gil de Pertierra, Alicia A. Redescription of Monticellia magna (Rego, dos 
Santos & Silva, 1974) (Eucestoda: Monticelliidae) parasite of Pime- 
lodus spp. (Pisces: Siluriformes) from Argentina, and morphological 
study of microtriches 1 i_ 20 

Loureiro, Marcelo, Azpelicueta, Maria de las Mercedes & Garcia, Gra- 
cida. Austrolebias arachan (Cyprinodontiformes, Rivulidae), a new 
species of annual fish from northeastern Uruguay 21-30 

Puthz, Volker. Edaphus comellinii sp. n. aus Sri Lanka (Coleoptera: Sta- 

phylinidae). 88. Beitrag zur Kenntnis der Euaesthetinen 31-33 

Salgado Costas, José Maria & Fresneda, Javier. Two new taxa of Lepto- 
dirini (Coleoptera: Leiodidae, Cholevinae) from the Cantabrian 
cornice (Asturias, Spain). Biogeographical observations 35-55 

Papp, Laszló. Description of the first apterous genus of Sciomyzidae 

(Diptera), from Nepal 57-62 

Pace, Roberto. Specie nuove o poco note di Homalotini, Silusini, Bolito- 
charini, Diestotini e Autaliini della Cina e della Thailandia (Cole- 
optera, Staphylinidae) 63-76 

Scheller, Ulf. Pauropoda (Myriapoda) from Vietnam (Pauropoda and 

Symphyla of the Geneva Museum XIII) 77-101 

Tilbury, Colin R. & Mariaux, Jean. A re-evaluation of the generic 
assignment of Bradypodìon spinosum (Matschie, 1892) and some 
considerations on the genus Rhampholeon Günther, 1874 103-110 

De Chambrier, Alain, Rego, Amilcar A. & Mariaux, Jean. Redescription 
of Brooksiella praeputialis and Goezeella siluri (Eucestoda: proteo- 
cephalidea), parasites of Cetopsis coecutiens (Siluriformes) from the 
Amazon and proposition of Goezeella dankbrooksi sp. n 111-120 

Kottelat, Maurice. Salaria economidisi, a new species of freshwater fish 
from Lake Trichonis, Greece, with comments on variation in S. fluvia- 
tilis (Teleostei: Blenniidae) 121-137 

Schwendinger, Peter J. & Martens, Jochen. A taxonomic revision of the 
family Oncopodidae IV The genus Oncopus Thorell (Opiliones, 
Laniatores) 139-174 

Baehr, Martin. Colasidia wau, a new leleupidiine species from Papua New 

Guinea (Insecta, Coleoptera, Carabidae, Zuphiinae) 175-181 

Merz, Bernhard. Revision of the Minettia fasciata species-group (Diptera, 

Lauxaniidae) 183-211 

Mirande, Juan Marcos, Aguilera, Gaston & Azpelicueta, Maria de las 
Mercedes. A new species of Astyanax (Characiformes, Characidae) 
from the upper rio Bermejo basin, Salta, Argentina 213-223 

Utiger, Urs & Schätti, Beat. Morphology and phylogenetic relationships 
of the Cyprus racer, Hierophis cypriensis, and the systematic status of 
Coluber gemonensis gyarosensis Mertens (Reptilia: Squamata: Colu- 
brinae) 225-238 



REVUE SUISSE DE ZOOLOGIE 

Volume 111 — Number 1 

Pages 

James, Samuel W., Hong, Yong & Kim, Tae Heung. New earthworms of 
Pheretima and Pithemera (Oligochaeta: Megascolecidae) from Mt. 
Arayat, Luzon Island, Philippines 3-10 

Gil de Pertierra, Alicia A. Redescription of Monticellia magna (Rego, dos 
Santos & Silva, 1974) (Eucestoda: Monticelliidae) parasite of Pime- 
lodus spp. (Pisces: Siluriformes) from Argentina, and morphological 
study of microtriches 11-20 

Loureiro, Marcelo, Azpelicueta, Maria de las Mercedes & Garcia, Gra- 
cida. Austrolebias arachan (Cyprinodontif ormes, Rivulidae), a new 
species of annual fish from northeastern Uruguay 21-30 

Puthz, Volker. Edaphus comellinii sp. n. from Sri Lanka (Coleoptera: Sta- 

phylinidae). 88th Contribution to the Knowledge of Euaesthetinae . . . 31-33 

Salgado Costas, José Maria & Fresneda, Javier. Two new taxa of Lepto- 
dirini (Coleoptera: Leiodidae, Cholevinae) from the Cantabrian 
cornice (Asturias, Spain). Biogeographical observations 35-55 

Papp, Laszló. Description of the first apterous genus of Sciomyzidae 

(Diptera), from Nepal 57-62 

Pace, Roberto. New or little known species of Homalotini, Silusini, 
Bolitocharini, Diestotini and Autaliini from China and Thailand (Cole- 
optera, Staphylinidae) 63-76 

Scheller, Ulf. Pauropoda (Myriapoda) from Vietnam (Pauropoda and 

Symphyla of the Geneva Museum XIII) 77-101 

Tilbury, Colin R. & Mariaux, Jean. A re-evaluation of the generic 
assignment of Bradypodion spinosum (Matschie, 1892) and some 
considerations on the genus Rhampholeon Günther, 1874 103-110 

De Chambrier, Alain, Rego, Amilcar A. & Mariaux, Jean. Redescription 
of Brooksiella praeputialis and Goezeella siluri (Eucestoda: proteo- 
cephalidea), parasites of Cetopsis coecutiens (Siluriformes) from the 
Amazon and proposition of Goezeella dankbrooksi sp. n 111-120 

Kottelat, Maurice. Salaria economidisi, a new species of freshwater fish 
from Lake Trichonis, Greece, with comments on variation in S. fluvia- 
tilis (Teleostei: Blenniidae) 121-137 

Schwendinger, Peter J. & Martens, Jochen. A taxonomic revision of the 
family Oncopodidae IV The genus Oncopus Thorell (Opiliones, 
Laniatores) 139-174 

Baehr, Martin. Colasidia wau, a new leleupidiine species from Papua New 

Guinea (Insecta, Coleoptera, Carabidae, Zuphiinae) 175-181 

Merz, Bernhard. Revision of the Minettia fasciata species-group (Diptera, 

Lauxaniidae) 183-211 

Mirande, Juan Marcos, Aguilera, Gaston & Azpelicueta, Maria de las 
Mercedes. A new species of Astyanax (Characif ormes, Characidae) 
from the upper rio Bermejo basin, Salta, Argentina 213-223 

Utiger, Urs & Schätti, Beat. Morphology and phylogenetic relationships 
of the Cyprus racer, Hierophis cypriensis, and the systematic status of 
Coluber gemonensis gyarosensis Mertens (Reptilia: Squamata: Colu- 
brinae) 225-238 

Indexed in Current Contents, Science Citation Index 



PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE 

CATALOGUE DES INVERTÉBRÉS DE LA SUISSE, N°s 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 DECHINODERMES ACTUELS 
CONSERVÉS DANS LES COLLECTIONS NATIONALES SUISSES, 
SUIVI D'UNE 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 
(COTENTIN-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. LObl, 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. Schätti & 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.— 



Volume 1 1 1 - Number 1 - 2004 
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 con- 
sidered 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 (high quality printing, if possible by a laser-printer), on one side only 
and double-spaced, on A4 (210 x 297 mm) or equivalent 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 disk (3,5"), or 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 formatted 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 
summarise the contents and conclusions of the paper and name all newly described taxa. The abstract is followed 
by less than 10 key-words, separated by hyphens, which are suitable for indexing. Terms already mentioned in the 
title of the paper have not to be repeated as key-words. 

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 Nomen- 
clature 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 and distribution, description 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 
symbols 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: 
Penare», E. 1888. Recherches sur le Ceratium macroceros. Thèse, Genève, 43 pp. 
Penare», 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 halftones. If possible, 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 
electronically scanned figures are submitted on disk or CD-R (never in MS-Powerpoint format !). this should be 
clearly indicated on the print-out of these figures enclosed with the print-out of the text. The following minimum 
resolutions should be respected: scanned line drawings: 600 dpi; scanned photographs: 300 dpi; digital photographs: 
5 millions of pixels. 

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-1211 Genève 6, Switzerland. 
Phone: +41 22 418 63 33 - Fax: +41 22 418 63 01. E-mail: volker.mahnert @ mhn.ville-ge.ch 
Home page RSZ: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 




SMITHSONIAN INSTITUTION jjjjggn 

3 9088 01260 2157 



]