Corresponding author: Pavel Stoev (
Academic editor: Robert Mesibov
The millipede genus
Callipodidans are still poorly documented in Southeast Asia, with only 15 species, four genera and three families being hitherto known in the region (
The family
The third family,
The biospeleological explorations of Dr Louis Deharveng and Mrs Anne Bedos (Muséum National d’Histoire Naturelle, Paris, hereafter MNHN) in caves in Vietnam, and the active collecting work of Dr Peter Jäger (Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt, hereafter SMF) and Mr Helmut Steiner (Hanau, Germany) in Laos revealed new material of Callipodida which was kindly offered to us for study. All examined specimens turned out to belong to new species of
All material treated in the paper is preserved in 70% ethanol and is shared between the MNHN, SMF and the National Museum of Natural History, Sofia (NMNHS). All photographs were taken with a Leica DFC 420 digital camera mounted on a Leica MZ16A stereomicroscope. Automontage Pro software from Syncroscopy was used for image-stacking 3D focus expansion. Drawings were made with the aid of a camera lucida mounted on Leica-WILD M10 and Leica-MZ16 microscopes. All illustrations were processed and additionally cleaned up with Adobe Photoshop CS.
All species descriptions are automatically exported at the time of publication to a wiki platform (
(based on
Holotype: ♂, 69 PT + telson, Vietnam, Hai Phong Province, Cat Ba Island, Hoa Cuong Cave near Gia Luang,
This species was found in two moderately long, humid caves. Tien Duc is approximately 100 m long, while Hoa Cuong is 100–120 m long. In Tien Duc, the specimen was found on non-humid walls. Both caves host a rich cave fauna, including unidentified cambalopsid and haplodesmid millipedes (L. Deharveng, in lit.).
For Cat Ba Island, the type locality.
Differs from
Maximal length
Width of PT 2=3<1=4<5<6<7. PT higher than broad, ratio: 1.05 : 1. Dorsal side of collum and PT2–3 smooth, lateral sides ribbed. Crests poorly developed, broad and flattened anteriorly, abruptly narrowing and more pronounced posteriorly; 5+5 between the ozopores on midbody PT. Ozopores small, barely visible on most PT, lying between crests 5 and 6 in midbody PT. Paraprocts divided into smaller dorsal and larger ventral sclerites. Dorsal sclerite surmounted with two macrosetae in a vertical row. Spinnerets: long and slender, ending with a long seta.
All legs white-yellowish, long and slender, ending with a long claw. Tarsal pads very poorly developed, present on leg-pairs 3–12. No particular modifications on coxae of pregonopodal legs, prefemora of legs 4–7 swollen. Leg-pair 9 (
Chaetotaxy: unknown, all setae broken off.
Gonopods (
unknown.
Holotype: ♂, 70 PT + telson, Vietnam, Quang Binh Province, Dong Hoi, Cha Noi: Hang Cha Noi (cave),
The species was found in several caves of moderate length (Hong Ruc about 50 m, Troc and Cha Noi more than 200 m), which seem to have never been surveyed previously. A road was built inside Hang Cha Noi during the War, and remains of rusted ammunitions were observed inside the cave at the time of collection. A rich cave fauna was found inside the Grotte de Troc, including two other species of Diplopoda, an undescribed cambalopsid, and
Named after the French zoologist Louis Deharveng, a passionate explorer of the caves of southeastern Asia, who together with A. Bedos collected this species.
Differs from congeners by having almost equally subdivided paraprocts, long antennae, strongly swollen and long gonocoxal process
Maximal length:
Width of PT: 1=2=3<4<5<6<7. PT higher than broad, ratio: 1.06 : 1. Dorsal side of collum and PT 2–3 smooth. Crests poorly developed, flattened, 5+5 between the ozopores on midbody PT, anterior part of crests broad, abruptly narrowing posteriorly. Ozopores small, barely visible on PT 5–6, lying on crest 6 in midbody PT, missing on the last 4 PT. Paraprocts divided into two almost equal-sized dorsal and ventral sclerites (
All legs white-yellowish, long and slender, ending with a long claw. Tarsal pads very poorly developed, present on leg-pairs 3–12. No particular modifications on coxae of pregonopodal legs. Prefemora of legs 4–7 swollen. Leg-pair 9 (
Chaetotaxy: unknown, all setae broken off.
Gonopods (
70–74 PT + telson; body colour darker, lateral sides light brownish; crests more pronounced than in males; second leg-pair unmodified (
Holotype: ad. ♂, 61 PT + telson; length
Three of the adult specimens were found in a cave situated at the foot of a limestone hill overgrown with sparse vegetation (
View of the type locality of
Named after Peter Jäger, curator of Arachnida and Myriapoda at SMF, who has been actively exploring the fauna of Laos since 2003 and collected the species.
Differs from congeners by the following set of characters: reduced eyes, composed of 10–11 ocelli; well expressed, narrow, pleurotergal crests; 59–61 PT; white-yellowish body and antennae; gonopods: process
Maximal length
Width of PT: 2–4<1=5<6<7<8<9<10. PT higher than broad, ratio: 1.06 : 1. Dorsal side of collum and PT 2–3 smooth; complete crests series appearing from PT 4 onwards. Crests at midbody PT well apart from each other; 6+6 (lateralmost one less pronounced), no secondary crest series, all crests well expressed (ribbed), narrow, equally broad along the whole length of metazonite, not touching anteriorly (
All legs white-yellowish, very long and slender, ending with a long claw. Tarsal pads poorly developed, present only on leg-pairs 3–12. No particular modifications on coxae of pregonopodal legs. Prefemora of posterior legs less swollen than others. Leg-pairs 4–7: coxa subquadrate; prefemur strongly swollen. Leg-pair 9 (
Chaetotaxy (
Chaetotaxy of anterior PT in
Anterior setae | Posterior setae | |
---|---|---|
Collum | 5+5 | - |
PT 2 | 5+5 | - |
PT 3 | 5+5 | - |
PT 4 | 5+5 | - |
PT 5 | 5+5 | - |
PT 6 | 5+5 | - |
Gonopods (
58–59 PT in adults; middle PT slightly broader than those of the male.
Only known from Xiao Cave, Hekou Yaozu Autonomous County, Yunnan Province, China.
Although callipodidans are often found in caves, especially in Southeast Asia and southern Europe, there are no species among them possessing an eye reduction similar to that observed in
Holotype: ♂; 71 PT + telson, Laos, Luang Phrabang Province, Ponsai District, Ben Nambo (Thapo) Village, Tham Gia (Bat cave) (E-48-001/07),
For detailed descriptions of the cave and its exact locality see
Named after the German biospeleologist Helmut Steiner, an active explorer of the caves of Laos, who collected the species.
Males: Differs from congeners by the following set of characters: 67–71 PT in adults; head and PT 1–4 white, remaining PT mottled light brown–grayish, antennomeres 2–6 brown; antennae moderately long, extending beyond posterior edge of PT7 when folded backwards; eyes black, well delineated, composed of 33–35 ocelli; midbody PT with 3+3 crests between ozopores; all crests flattened, almost equally broad along metazonal length, only slightly narrowed posteriorly and touching each other anteriorly. Gonopods: differ from those of congeners by the laterally narrowed gonocoxal process
Maximal length:
Width of PT: 2–3<1=4<5<6<7<8<9<10. PT slightly higher than broad; ratio: 1.07 : 1. Dorsal side of collum and PT 2–3 smooth, ribbed only laterally; complete crests series appearing from PT 4 onwards. Midbody PT with 3+3 crests between ozopores; no secondary crest series, all crests flattened, almost equally broad along the metazonal length, only slightly narrowed posteriorly and touching each other anteriorly. Ozopores on midbody PT lying between crests 3 and 4, visible from sixth to last but two PT. Paraprocts divided into smaller dorsal and bigger ventral sclerites. Each dorsal sclerite with a pair of macrosetae in a vertical line. Spinnerets: long and slender, ending with a long seta each. All setae on telson dark brown, contrasting with the whitish background.
All legs white-yellowish, moderately long and slender, ending with a long claw. Tarsal pads poorly developed, present only on leg-pairs 3–12. No particular modifications on coxae of pregonopodal legs. Prefemora of legs 4–7 swollen. Leg-pair 9 (
Chaetotaxy: pleurotergal setae barely visible, minute, one each on PT 1 and 2, others presumably broken off.
Gonopods (
unknown.
Only known from the type locality in Thailand, Saraburi Province, Muang District, Sriwilai Cave Temple,
This species is easily distinguished from congeners by its strikingly snow-white anterior pleurotergites and antennal tips, contrasting with a generally dark brown body (
ad. ♀, Vietnam, Lang Son Province, Huu Lung Area, Snake cave, 19.III.1989, P. Beron leg. (NMNHS); 1 juv., same province and area, cave at 97 km N from Hanoi, 20.III.1989, P. Beron leg. (NMNHS).
The genus
In contrast, peripheral characters such as the relative lengths of antennomeres (
Graph showing antennomere lengths in the new species.
9th leg in
Relative length of antennae in
Species | Antennae reaching back |
---|---|
|
beyond posterior edge of PT9 |
|
beyond posterior edge of PT10 |
|
beyond posterior edge of PT10 |
|
beyond posterior edge of PT8* |
|
beyond posterior edge of PT7 |
|
slightly beyond posterior edge of PT4 |
A close-up photograph of the gonopodal region of
Distribution of the species of genus
1(4) | body and antennae uniformly yellow-whitish; eye unpigmented, with less than 20 ocelli | 2 |
2(3) | 55–56 PT; 16 ocelli; crests low, flattened; cannula of gonopods short and coiled; Xiao cave, China |
|
3(2) | 59–61 PT, 10–11 ocelli; crests well-developed, ribbed; cannula long and straight; caves in Khammouan Province, Laos | |
4(1) | either body or antennae with brown pigment; eye black, with more than 30 ocelli | 5 |
5(6) | body brown with light middorsal band stretching from PT 5 to the body end; first four PT, anterior part of head and antennal articles 6 and 7 snow white; eye with 45–50 ocelli; antennae short, slightly extending beyond posterior edge of PT 4, femoroid with two slender acicular and one shorter and subfalcate terminal projections; surroundings of Sri Wilai Temple, Thailand |
|
6(5) | body generally white-yellowish, sometimes mottled gray-brownish; antennal articles 2–5 brown; antennae long, extending beyond the posterior edge of PT 7 when folded backwards; eye with less than 40 ocelli; femoroid with three slender acicular and one shorter and subfalcate terminal projections | 7 |
7(8) | length of antennae |
|
8(7) | length of antennae more than 7 mm; 5+5 crests between ozopores on midbody PT; gonocoxal process |
9 |
9(10) | body length |
|
10(9) | body length |
We cordially thank Peter Jäger, Helmut Steiner, Louis Deharveng and Anne Bedos for the granted material. Nesrine Akkari has greatly contributed to the production of photographs and assembling the images, for which we are extremely thankful. P. Jäger and G. Brovad (Natural History Museum of Denmark) provided figures 13 and 24. This project was made possible with funding to PS from Deutsche Forschungsgemeinschaft and the EC FP6 Integrated Infrastructure Initiative Synthesys (DK-TAF).