Corresponding authors: Sergei I. Golovatch(
Academic editor: Robert Mesibov
The genus
The present paper provides a new review of
New material derives from several provinces of northern Thailand taken between 2006 and 2010. All holotypes, as well as most of the paratypes and non-types, are in the collection of the Museum of Zoology, Chulalongkorn University, Bangkok, Thailand (CUMZ), some duplicates also being donated to the collections of the Natural History Museum of Denmark, University of Copenhagen, Denmark (ZMUC), and of the Zoological Museum, State University of Moscow, Russia (ZMUM), as indicated in the text.
Coloration was photographed in the laboratory (both live and alcohol material) for all of the encountered species. Material was then fixed, preserved in 75% ethanol and studied in the lab using a standard Olympus stereomicroscope. Scanning electron micrographs (SEM) were taken using a JEOL, JSM-5410 LV microscope. After SEM examination of the gonopods, they were returned to alcohol.
The following species of
The gonopod telopodite in
♂ (CUMZ), Thailand, Tak Province, Umphang District, near Umphang City, ca 490 m, 16°2'20N, 98°52E, 6.07.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn.
1 ♂, 1 ♀, 2 juv. (CUMZ), same locality, together with holotype. 6 ♂, 4 ♀ (CUMZ), 3 ♂ (ZMUC), 3 ♂ (ZMUM), Tak Province, Umphang District, Doi Hua Mod, 900 m, 16°3'14N, 98°49'16E, 5.06.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn. 6 ♂, 1 ♀ (CUMZ), same Province, same District, Cave Ta Ko Bi, ca 530 m, 16°03'14N, 98°49'14E, 5.07.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn.
To emphasize the spiniform processes
Differs from congeners in both processes
Length 26 mm (holotype), 25–29 mm (♂), 33–38 mm (♀), width of midbody pro- and metazona 2.0 and 2.9 mm (holotype), 1.8–2.4 and 3.1–3.2 mm (♂), 2.4–2.7 and 3.3–3.8 mm (♀), respectively. Coloration of live animals black-brown (
Clypeolabral region of head very densely, vertigial region sparsely setose. Epicranial suture distinct. Antennae long and slender, reaching behind segment 4 (♂) or 3 (♀) dorsally. In width, head < collum < segments 3–4 < 2 < 5–16 (♂), or head = segment 3 < 4 < collum < segments 5–16 (♀); thereafter body gradually and gently tapering towards telson (
Tegument generally rather smooth and shining, but prozona very finely rugulose, metaterga often rugose (
Sterna moderately setose, without modifications; a deeply notched sternal lobe between ♂coxae 4 (
Gonopods (
♂ (CUMZ), Thailand, Mae Hong Son Province, Pangmapha District, near Cave Pha Mon, 19.07.2008, leg. S. Panha, J. Sutcharit & N. Likhitrakarn.
1 ♂, 2 ♀ (CUMZ), same lcality, together with holotype. 1 ♂ (CUMZ), same District, Mae Lana crossroads, 19.07.2008, leg. S. Panha, J. Sutcharit & N. Likhitrakarn.
To emphasize the large size of this species
Differs from congeners in the large size, coupled with a short spiniform process
Length 41 mm (holotype), 40–42 mm (♂), 38–39 mm (♀), width of midbody pro- and metazona 3.0 and 4.5 mm (holotype), 2.8–3.0 and 4.3–4.5 mm (♂), 3.6–3.8 and 4.7–5.0 mm (♀), respectively. Coloration in alcohol dark brown to black-brown (
All characters as in
Antennae short and slender (
Tegument generally rather smooth and either dull (only in places modestly shining) or shining (
Sterna moderately setose, without modifications; a slightly notched sternal lobe between ♂ coxae 4 (
Gonopods (
♂ (CUMZ), Thailand, Chiang Mai Province, Fang District, Doi Phahom Pok National Park, 6.07.2009, leg. A. Pansook.
1 ♂, 1 ♀ (CUMZ), 1 ♂ (ZMUC), 1 ♂ (ZMUM), same locality, together with holotype.
To emphasize the extremely long spines
Differs from congeners in process
Length ca 30 mm (holotype), 27–30 mm (♂), 32.5 mm (♀), width of midbody pro- and metazona 2.0 and 2.9 mm (holotype), 1.9–2.4 and 2.8–3.3 mm (♂), 2.5 and 3.0 mm (♀), respectively. Coloration of live animals, as well as of alcohol material black-brown (
All characters as in
Antennae rather short and slender, reaching behind to end of segment 3 (♂) dorsally. Collum with paraterga like rudimentary flaps, especially poorly developed in ♀. In width, head < collum = segments 3–4 < 2 < 5–16 (♂) (
Metaterga with two transverse rows of rather long setae: 2+2 in anterior and 2(3)+2(3) in posterior row, the latter often abraded, but then readily traceable as insertion points on low longitudinal ridges or tubercles (
Sterna moderately setose, without modifications; an entire, linguiform, sternal lobe between ♂ coxae 4 (
Gonopods (
♂ (CUMZ), Thailand, Nan Province, Pua District, Ton Tong Waterfall, ca 1130 m, 19°10'52N, 101°5'45E, 10.10.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn.
To emphasize the velum-shaped end of gonopod lobe
Differs from congeners except
Length ca 28 mm, width of midbody pro- and metazona 2.0 and 2.6 mm, respectively. Coloration of live animal and alcohol material rather uniformly dark brown to blackish (
All characters as in
Clypeolabral region of head very densely setose, but vertigial region bare. Antennae short and barely reaching behind segment 2 dorsally. In width, head = segments 3 and 4 < collum < segment 2 < 5–16; thereafter body gradually and gently tapering towards telson (
Metaterga with two transverse rows of long setae: 2+2 in anterior and 2(3)+2(3) in posterior row, the latter often abraded, but then readily traceable as insertion points. Axial line thin, in places incomplete, but readily visible on both halves of metaterga. Paraterga strongly developed (
Sterna rather densely setose, without modifications except for a subquadrate, setose, sternal lobe between coxae 4 (
Gonopods (
♂ (CUMZ), Thailand, Chiang Mai Province, Chom Thong District, Doi Inthanon National Park, summit, 2520 m, 18°34'29N, 98°28'48E, 12.10.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn.
1 ♂ (CUMZ), same locality, together with holotype.
To emphasize the close resemblance to
Very similar to
Length 31 mm (holotype) or 31.5 mm (♂), width of midbody pro- and metazona 2.4 and 3.2 mm (holotype) or 2.3 and 3.4 mm (♂), respectively. Coloration of live animals and alcohol material uniformly blackish-brown (
All characters as in
Antennae rather short and slender, reaching behind segment 3 dorsally. In width, head = segment 3 < collum < segments 2 and 4 < 5–16; thereafter body gradually and gently tapering towards telson (
Paraterga on collum like large rounded flaps (
A low, only slightly divided, setose lobe between coxae 4 (
Gonopods (
The following seven species have been illustrated in additional detail to confirm their identities, as well as to provide further information concerning both their variation and distribution.
2 ♂ (CUMZ), Thailand, Chiang Mai Province, Chom Thong District, Doi Inthanon National Park, Siriphum Waterfall, ca 1320 m, 18°32'49N, 98°30'57E, 13.10.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn; 2 ♂ (CUMZ), same locality, main road, 10 km before summit, ca 1700 m, 18°31'15N, 98°30'1E, 13.10.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn.
This strictly topotypic material fully agrees with the original description (
3 ♂ (CUMZ), Thailand, Chiang Mai Province, Mueang Chiang Mai District, Doi Suthep National Park, ca 1300 m, 18°48'9N, 98°54'11E, 20.04.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn; 4 ♂, 3 ♀, 2 juv. (CUMZ), same Province, WiangKaen District, Doi Phatang, 6.07.2009, leg. S. Panha & J. Sutcharit.
This partly topotypic material fully agrees with the original description (
1 ♂ (CUMZ), Thailand, Chiang Mai Province, Chom Thong District, Doi Inthanon National Park, main road, 10 km before summit, ca 1700 m, 18°31'15N, 98°30'1E, 13.10.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn.
This strictly topotypic material fully agrees with the original description (
4 ♂, 7 ♀, 1 juv. (CUMZ), Thailand, Chiang Mai Province, Mueang Chiang Mai District, Doi Suthep National Park, ca 1300 m, 18°48'9N, 98°54'11E, 22.10.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn.
This represents a second record of this species, the type locality being Doi Inthanon National Park in the same province. Our material almost fully agrees with the original description (
2 ♂ (CUMZ), Thailand, Chiang Mai Province, Chom Thong District, Doi Inthanon National Park, summit, 2520 m, 18°34'29N, 98°28'48E, 12.10.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn.
This strictly topotypic material fully agrees with the original description (
4 ♂ (CUMZ), Thailand, Chiang Mai Province, Phrao District, Buathong Waterfall forest park, 510 m, 19°4'10N, 99°4'46E, 29.09.2009, leg. N. Likhitrakarn.
This near-topotypic material fully agrees with the original description (
5 ♂, 3 ♀, 1 juv. (CUMZ), Thailand, Chiang Mai Province, Chom Thong District, Doi Inthanon National Park, Siriphum Waterfall, ca 1320 m, 18°32'49N, 98°30'57E, 13.10.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn; 2 ♂, 2 ♀ (CUMZ), same province, Wiang Kaen District, Doi Phatang, 25.10.2008, leg. S. Panha & J. Sutcharit; 5 ♂, 2 ♀ (CUMZ), Lampang Province, Ngao District, Thum Pha Thai, 23.10.2008, leg. S. Panha, J. Sutcharit & N. Likhitrakarn; 1 ♂, 1 ♀ (CUMZ), Chiang Rai Province, Mueang Chiang Rai District, Ban Pang Rim Kon, 10.07.2006, leg. S. Panha; 1 ♂ (CUMZ), same province, Thoeng District, Phucheefah, 10.07.2006, leg. S. Panha & J. Sutcharit; 2 ♂, 1 ♀ (CUMZ), same province, Wiang Kaen District, Doi Pha Tang, 10.07.2006; 1 ♂, 2 ♀ (CUMZ), Phayao Province, Chiang Kham District, Nam Min Waterfall, 23.10.2008, leg. S. Panha & J. Sutcharit; 2 ♂, 2 ♀ (CUMZ), Phrae Province, Rong Kwang District, Tham Pha Nang Khoi, ca 280 m, 18°22'10N, 100°21'12E, 9.10.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn; 2 ♂, 2 ♀ (CUMZ), same locality, 29.09.2010, leg. J. Sutcharit & P. Pimvichai; 9 ♂, 6 ♀, 2 juv. (CUMZ), Nan Province, Pua District, Ton Tong Waterfall, ca 1130 m, 19°12'36N, 101°4'14E, 10.10.2009, leg. S. Panha, J. Sutcharit & N. Likhitrakarn.
This species has long been known as perhaps the most widespread and common congener in northern Thailand, also showing considerable variation both in body texture and gonopod structure (
1 | Most ♂ prefemora evidently swollen laterally ( |
2 |
– | All ♂ prefemora normal, not bulged laterally | 25 |
2 | Surface of metaterga virtually smooth, at best extremely faintly rugulose in certain places and/or with a few barely traceable (setiferous) tubercles near caudal margin (setae mostly broken off) | 3 |
– | Surface of metaterga mostly rugulose to coarsely rugose/tuberculate | 6 |
3 | Paraterga moderately developed ( |
|
– | Paraterga relatively well-developed, radio of ♂ midbody prozonite to metazonite width over 1:1.2. Transverse sulcus on metaterga starting from segment 4 or 5, always fully developed and reaching base of paraterga on segment 5 | 4 |
4 | Calluses without incisions. Gonopod postfemoral lobe |
|
– | Calluses mostly with 1–2 incisions. Gonopod postfemoral lobe |
5 |
5 | Metatergal surface entirely smooth, polished, without tubercles. Midline wanting. Pleurosternal carinae relatively weak, as small teeth only on a few anteriormost segments; ♂ legs without adenostyles (= tubercles). Gonopods with three rather small, spiniform processes near base of lobe |
|
– | Metaterga at best only very faintly rugulose near waist, near sulcus and/or at base of paraterga, with 2–3 weak, oblong tubercles near rear margin. Midline mostly traceable at least on anterior halves of metaterga. Pleurosternal carinae more strongly developed; most ♂ postfemora and tibiae tuberculiferous. Gonopods with only two larger outgrowths near base of lobe |
|
6 | Metaterga without evident setiferous tubercles, only sometimes with very small, rudimentary wrinkles or knobs | 7 |
– | Metaterga with evident setiferous tubercles | 11 |
7 | Body larger: 38–42 mm long, 2.8–3.8 and 4.3–5.0 mm wide on pro- and metaterga, respectively. Gonopod with a short spiniform process |
|
– | Body smaller. Gonopod otherwise | 8 |
8 | Both processes |
|
– | Gonopod otherwise | 9 |
9 | Gonopod process |
|
– | Gonopod otherwise | 10 |
10 | ♂ legs shorter, ca 1.2–1.3 times as long as midbody height ( |
|
– | ♂ legs longer, ca 1.6–1.7 times as long as midbody height ( |
|
11 | Most metaterga with a pattern of 2+2 and 2+2 setiferous tubercles in two rows, rear row somewhat less strongly developed than fore one |
|
– | Most metaterga with rear row of setiferous tubercles or wrinkles more strongly developed than fore row, the latter (next to) wanting | 12 |
12 | Transverse sulcus on metaterga starting from segment 4, either fully or almost fully developed there, always fully developed from segment 5 | 13 |
– | Transverse sulcus on metaterga starting only from segment 5 | 16 |
13 | Transverse sulcus fully developed and reaching base of paraterga already from segment 4. Gonopod tooth |
|
– | Transverse sulcus fully developed only from segment 5. Gonopod tooth |
14 |
14 | Paraterga 2 caudolaterally rather broadly rounded. Gonopod relatively simple, process |
|
– | Paraterga 2 caudally pointed. Gonopods more complex | 15 |
15 | Coloration dark brown, without cingulate pattern. Sternal lamina between ♂ coxae 4 low and distinctly bimodal ( |
|
– | Coloration pale, with a cingulate pattern. Sternal lamina between ♂ coxae 4 high, subquadrate. Gonopod tooth |
|
16 | Paratergal corner protruding caudad beyond rear contour only from segment 15, being obtusangular or subrectangular and lying more or less within the contour until segment 14 |
|
– | Paratergal corner protruding caudad before segment 14, mostly pointed | 17 |
17 | Pattern of tergal setation on segments 18 and/or 19: 2+2 and 5+5 in two rows | 18 |
– | Pattern of tergal setation at least on segments 5–19: 2+2 and 4+4 in two rows | 21 |
18 | Pattern of tergal setation 2+2 and 5+5 on both segments 18 and 19. Paraterga 2 caudally pointed. Epiproct with pre-apical incisions very close to apical knobs. Sternal lamina between ♂ coxae 4 an unusually low, even ridge. Adenostyles on midbody ♂ postfemora and, to a lesser extent, tibiae exceptionally prominent |
|
– | Pattern of tergal setation 2+2 and 5+5 on segment 19. Paraterga 2 more or less narrowly rounded. Pre-apical incisions on epiproct better removed from tip. Sternal lamina between ♂ coxae 4 concave medially. Ventral adenostyles on ♂ legs less prominent | 19 |
19 | Body smaller: width ca 2.0 mm. Sternal lamina between ♂ coxae 4 as a pair of separate, setiferous tubercles ( |
|
– | Body larger: width over 3.0 mm. Sternal lamina between ♂ coxae 4 single. Ventral adenostyles on ♂ legs more prominent. Gonopod with a spine near base of process |
20 |
20 | Sternal lamina between ♂ coxae 4 high, emarginate ( |
|
– | Sternal lamina between ♂ coxae lower, slightly concave. Adenostyles on ♂ postfemora and tibiae less strongly developed. Gonopods more complex, spine |
|
21 | Paraterga 2 pointed caudally. Sternal lamina between ♂ coxae 4 exceptionally densely setose, low, concave ventrally ( |
|
– | Paraterga 2 more or less narrowly rounded caudally. Sternal lamina between ♂ coxae 4 higher and less strongly setose. Gonopod outgrowths |
22 |
22 | Sternal lamina between ♂ coxae 4 with a straight ventral margin. Pleurosternal carinae poorly developed, in ♂ slightly projecting caudad beyond rear margin only until segments 8–10 | 23 |
– | Sternal lamina between ♂ coxae 4 slightly concave ventrally. Pleurosternal carinae better developed, in ♂ slightly projecting caudad beyond rear margin at least till segment 15 | 24 |
23 | Body smaller: width up to 3.1–3.2 mm. Mid-dorsal line very clear on both halves of metaterga. Gonopods relatively simple, with both |
|
– | Body larger: width 4.0–5.3 mm. Mid-dorsal line not so well-developed at least on rear halves of metaterga. Gonopods more complex, with both |
|
24 | Metatergum 19 slightly rugulose posteriorly. Calluses on segment 2 with three, on following paraterga with two, incisions. Gonopods extremely complex, with numerous spiniform outgrowths |
|
– | Metatergum 19 entirely smooth. Calluses with two and three incisions on poreless and poriferous paraterga, respectively. Gonopod less strongly differentiated |
|
25 | Either most of ♂ sterna with oblique tubercles or spines, or only anterior sterna with small cones near coxae | 26 |
– | Neither spines nor tubercles on ♂ sterna | 28 |
26 | Only rear sternum on most of ♂ segments with a pair of small spines. Metaterga mostly with 2+2 and 3+3 setiferous tubercles in two transverse rows. Gonopod process |
|
– | Fore and rear sterna of most of ♂ segments with a pair of tubercles and spines, respectively. Fore row of tergal setae not borne on tubercles, rear row on 2+2 tubercles | 27 |
27 | Transverse sulcus starting from metatergum 4, fully developed from metatergum 5. ♂ tarsal brushes missing. Northern Vietnam |
|
– | Transverse sulcus starting from metatergum 3, still underdeveloped on metatergum 4, fully developed from metatergum 5. ♂ tarsal brushes present only on a few anteriormost legs. Yunnan, China |
|
28 | Metaterga entirely smooth and polished, devoid of evident tubercles, at best extremely faintly rugulose near transverse sulcus | 29 |
– | Metaterga rather clearly rugose/tuberculate/granulate, posterior row of setae at least partly borne on tubercles | 32 |
29 | Transverse sulcus on metaterga starting from segment 4, but fully developed and reaching base of paraterga only from segment 5. Ventral adenostyles on ♂ legs: a distal knob on femur, a distomedial knob on postfemur, and a parabasal knob on both tibia and tarsus | 30 |
– | Transverse sulcus on metaterga starting only from segment 5. Pattern of ♂ leg adenostyles otherwise | 31 |
30 | Head a little wider than collum and subequal in width to segment 3. Paraterga caudally considerably acutangular and beak-shaped only from segment 14. Gonopod process |
|
– | Head a little narrower than collum and subequal in width to segment 2. Paraterga caudally beak-shaped already from segment 7, especially strongly so from segment 12. Gonopod process |
|
31 | Larger species: body width 3.1 mm. Pattern of tergal setation: 2+2 and 3+3 to 6+6 in two rows, rear row easily traceable due to insertion points. Metaterga very finely rugulose only near transverse sulcus. Epiproct unusually broad. Pads instead of adenostyles on ♂ femora, postfemora, tibiae (all distally) and tarsi (almost entirely) |
|
– | Small species: width 1.6 mm. Only a single row of 2+2 tergal setae. Metaterga entirely smooth. Epiproct not so wide. Adenostyles on ♂ legs present |
|
32 | Metatergal surface polished and smooth except for conspicuous tubercles in two rows | 33 |
– | Metaterga at least partly rugulose/rugose to granular; at most one row of tubercles | 35 |
33 | Paraterga very poorly developed, rounded, low, projecting slightly caudad beyond rear contour like small knobs only on segments 18 and 19. Calluses virtually devoid of incisions. Transverse sulcus on metaterga poorly developed, starting already from segment 2, although fully developed only from segment 5. A paramedian pair of denticles between ♂ coxae 5 behind a prominent, subquadrate lamina between ♂ coxae 4. Gonopod process |
|
– | Paraterga better developed, protruding caudad beyond rear contour at least from segment 5. Calluses always at least with one lateral incision. Transverse sulcus starting only from segments 3–5. Dentiform tubercles between ♂ coxae 5 missing. Gonopod process |
34 |
34 | Paraterga acutangular caudally and pointed beak-like already from collum. Tergal setiferous tubercles: 3+3 and 5+5 on segments 16–19. Adenostyle pattern on ♂ legs: a distal knob on femora and a parabasal knob on most of postfemora, tibiae, and tarsi. Gonopod process |
|
– | Paraterga acutangular caudally and pointed beak-like only from segment 4. 2+2 and 4+4 tergal setiferous tubercles on segments 16–19. Adenostyle pattern on ♂ legs: a proximal finger-shaped tubercle crowned with a bunch of setae only on femora 6, 8, and 9. Gonopod process |
|
35 | Transverse sulcus on metaterga starting and fully developed from segment 5. Ventral adenostyles present on all ♂ podomeres except coxa | 36 |
– | Transverse sulcus on metaterga starting from segment 4, but fully developed only from segment 5. Ventral tubercles only on some of ♂ telopoditomeres | 37 |
36 | Metaterga rugulose also in front of transverse sulcus, at rear margin with several oblong tubercles. Sternal lamina between ♂ coxae 4 like a pair of setiferous knobs. Neither gonopod lobe |
|
– | Metaterga rugose only behind transverse sulcus, without evident turbercles at rear margin. Sternal lamina between ♂ coxae 4 single. Both gonopod lobe |
|
37 | Metaterga modestly rugulose only near transverse sulcus, posteriorly neither granular nor microtuberculate. Calluses broad. Gonopod process |
38 |
– | Metaterga distinctly rugose-granular/microtuberculate even on fore halves. Calluses narrow. Gonopod process |
39 |
38 | Caudal corner of paraterga pointed from segment 3. Pleurosternal carinae particularly well-developed, surpassing rear contour until segment 16 or 17. Adenostyles often present on ♂ prefemora, pattern as in |
|
– | Caudal corners of paraterga mainly narrowly rounded, pointed only from segment 15. Pleurosternal carinae less strongly developed |
|
39 | Coloration dark, brown. Sternal lamina between ♂ coxae 4 like a pair of separate, setiferous tubercles preceded by another pair of very small tubercles between coxae 3. Gonopod with lobe |
|
– | Coloration uniformly pale. Sternal lamina between ♂ coxae 4 single, not accompanied by additional tubercles in front or behind. Gonopod lobe |
40 |
40 | Mostly 3+3 tubercles at rear margin of metaterga. Sternal lamina between ♂ coxae 4 distinctly emarginate. Larger adenostyles close to midlength on ♂ postfemora and tibiae, femora with a distal knob. Gonopod process |
|
– | Mostly 4+4 tubercles at rear margin of metaterga. Sternal lamina between ♂ coxae trapeziform. Larger adenostyles on both postfemora and tibiae more distal, femoral knob missing. Gonopod process |
|
In Thailand, all
Distribution of
Almost all
Within
We would like to extend our deep gratitude to all of the collectors involved, mainly staff of the Animal Systematics Research Unit of Chulalongkorn University, Bangkok. The first author is deeply obliged to the programme Strategic Scholarships for Frontier Research Network for the Joint Ph.D. Program Thai Doctoral degree from the Office of the Higher Education Commission, CHE–RG under the Limestone Biodiversity Project and the BRT Programme (TRF/BIOTECH Thailand Biodiversity Research and Training Program) for their financial support.