Monograph |
Corresponding author: Somsak Panha ( somsak.pan@chula.ac.th ) Academic editor: Pavel Stoev
© 2016 Warut Siriwut, Gregory Edgecombe, Chirasak Sutcharit, Piyoros Tongkerd, Somsak Panha.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Siriwut W, Edgecombe GD, Sutcharit C, Tongkerd P, Panha S (2016) A taxonomic review of the centipede genus Scolopendra Linnaeus, 1758 (Scolopendromorpha, Scolopendridae) in mainland Southeast Asia, with description of a new species from Laos. ZooKeys 590: 1-124. https://doi.org/10.3897/zookeys.590.7950
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The centipede genus Scolopendra in mainland Southeast Asia is reviewed taxonomically based on morphological characters, informed by a molecular phylogenetic analysis using sequences from three mitochondrial and nuclear genes (COI, 16S rRNA and 28S rRNA). Eight nominal species of Scolopendra, namely S. morsitans Linnaeus, 1758, S. subspinipes Leach, 1816, S. dehaani Brandt, 1840, S. multidens Newport, 1844, S. calcarata Porat, 1876, S. japonica Koch, 1878, S. pinguis Pocock, 1891, and S. dawydoffi Kronmüller, 2012, are redescribed together with some revision of type materials. Geographical variation in each species has been compiled with reference to samples that span their distribution ranges in Southeast Asia and some parts of neighbouring areas such as East Asia, the Indian Ocean, and Africa. Comparative study of traditional taxonomic characters from external morphology provides further information to distinguish some closely related species. Scolopendra cataracta Siriwut, Edgecombe & Panha, sp. n., is described from the southern part of Laos, with additional records in Thailand and Vietnam. The phylogenetic framework for Southeast Asian Scolopendra recognizes S. calcarata + S. pinguis, S. morsitans, and a S. subspinipes group that unites the other six species as the main clades. Within the S. subspinipes group, two monophyletic groups can be distinguished by having either slender or short, thick ultimate leg prefemora and different numbers of apical spines on the coxopleuron. Scolopendra arborea Lewis, 1982, is placed in subjective synonymy with S. dehaani. A survey of external morphology of the genital segments confirms its potential for improving species identification in Scolopendra. Some observations on biology and behaviour are recorded based on field surveys in this area.
Chilopoda , Scolopendra , systematics, distribution, phylogeny, species diversity
The genus Scolopendra Linnaeus, 1758, is among the predominant centipede groups in tropical regions. These animals are generalist feeders that play an important role as one of the top carnivorous invertebrates in soil ecosystems. In several Asian countries, Scolopendra has symbolic status or figures in superstitions, and is used commercially in traditional medicine (
The taxonomic study of Scolopendra dates back to the late 19th to mid-20th centuries (
To date, 99 described species of Scolopendra have been recorded (
The validity of various scolopendrid species has been ambiguous because their distributions have not been comprehensively documented and/or because the diagnostic value of particular taxonomic characters has been unclear (
In this work, we review Scolopendra species in mainland Southeast Asia. The type material of some species has been re-described and, where available, type material is photographed and illustrated. All species are compared with the most closely allied congeners to provide distinguishing taxonomic characters. Variability in morphological characters is recorded in order to document geographical variation. The description of a new species is based on specimens from three SE Asian countries. Molecular phylogeny of three standard genes is analyzed, adding new samples to previous work on Scolopendra, to test the monophyly of each species and to determine the phylogenetic position of new species. An identification key to Scolopendra is presented and distribution ranges for species are updated.
Material examined. Specimens were collected mainly throughout mainland Southeast Asia, principally from Thailand, Laos, Cambodia, Myanmar, Singapore and Malaysia and kept at Chulalongkorn UniversityMuseum of Zoology, Bangkok, Thailand. Examination of additional Southeast Asian and other Oriental regional collections including available type material was based on both identified and previously undetermined specimens in several museums. All specimens were observed by using either a LEICA MZ 16A, Nikon SMZ25 or Olympus stereo-microscope. Morphological characters were photographed using montaged image stacks. Each morphological feature was serially captured with a Canon 700d linked to an automated calibration program, either Cell’D imaging or Helicon Focus on a desktop PC. In addition, illustration of some morphological variation was made by free-hand drawings.
Institutional abbreviations:
Behaviour, biology and distribution. Feeding behaviour and brooding of eggs and hatchlings was observed and photographed both in the field and the laboratory. Characteristics of habitats and brood chambers are discussed in
The distributional ranges of all SE Asian Scolopendra species were reinvestigated based on field sampling, museum collections, and literature records from this region. Localities cited in the descriptions are arranged geographically and are separated in two sections:
I. A determined locality refers to the corrected name of a locality. In cases of inaccurate spelling and outdated names on old labels, we provide the corrected name in square parentheses based on resources from the internet and/or historical notes. Spellings of new collection localities in Thailand were transcribed by the Thai Romanization program (
II. An undetermined locality refers to a name that is localized at only a regional scale such as by region or country.
Distribution maps for each species include the records from recent field surveys and specimens from museum collections that provide sufficiently detailed locations. Each of those localities is marked by a filled symbol. Some localities from previous taxonomic work are included using a blank symbol. All undetermined localities and some specimens which were attributed only to a region, island or country have been excluded from the distribution maps.
Species identification and description. Morphological terminology follows the standardized terminology (
Abbreviations for terminology applied to morphology used in descriptions and some comparison tables are as follow: PS, paramedian suture; AP, apical spine; SAP, subapical spine; LS, lateral spine; DS, dorsal spine; V, ventral spine; VL, ventro-lateral spine; VM, ventro-median spine; M, median spine; DM, dorso-median spine; SP, spine on prefemoral process; ULBS, ultimate leg-bearing segment. Abbreviated terminology for the genital region used in figures is as follows: IM, intermediate membrane; AV, anal valve; GP, gonopod; LA, lamina adanalis; LS, lamina subanalis; PN, penis; SGS I, sternite of genital segment 1; SGS II, sternite of genital segment 2; TGS, tergite of genital segment.
The list of synonyms for each Scolopendra species follows Chilobase (http://chilobase.biologia.unipd.it/). Diagnoses are revised from
Phylogenetic reconstruction. Southeast Asian and some temperate Asian Scolopendra sequences were obtained from GenBank, based on previous phylogenetic analyses (
Scolopendra sequences used in phylogenetic reconstruction in present study. Abbreviation names of voucher ID codes refer to museum collections as follow:
Species | Taxon locality | Voucher ID number | COI | 16S | 28S | Reference |
---|---|---|---|---|---|---|
Scolopendra morsitans Linnaeus, 1758 | Khon Kaen, Thailand |
|
KR705662 | KR705600 | KR705724 | 1 |
Nan, Thailand |
|
KR705661 | KR705599 | KR705723 | 1 | |
Chonburi, Thailand |
|
KR705660 | KR705598 | KR705722 | 1 | |
Surin, Thailand |
|
KR705666 | KR705604 | KR705728 | 1 | |
Chiang Mai, Thailand |
|
KR705665 | KR705603 | KR705727 | 1 | |
Sa Kaeo, Thailand |
|
KR705664 | KR705602 | KR705726 | 1 | |
Sisophon, Cambodia |
|
KR705663 | KR705601 | KR705725 | 1 | |
Singapore |
|
KR705636 | KR705574 | KR705698 | 1 | |
Scolopendra subspinipes Leach, 1816 | Papua New Guinea |
|
KF676528 | KF676488 | – | 3 |
Martinique |
|
HQ402554 | HQ402502 | – | 3 | |
Scolopendra cingulata Latreille, 1829 | Spain |
|
HM453310 | HM453220 | – | 3 |
Scolopendra dehaani Brandt, 1840 | Lopburi, Thailand |
|
KR705689 | KR705627 | KR705751 | 1 |
Ayutthaya, Thailand |
|
KR705688 | KR705626 | KR705750 | 1 | |
Lan Island, Rayong, Thailand |
|
KR705684 | KR705622 | KR705746 | 1 | |
Sa Kaeo, Thailand |
|
KR705682 | KR705620 | KR705744 | 1 | |
Trad, Thailand |
|
KR705681 | KR705619 | KR705743 | 1 | |
Sichang Island, Chonburi, Thailand |
|
KR705683 | KR705621 | KR705745 | 1 | |
Chiang Mai, Thailand |
|
KR705659 | KR705597 | KR705721 | 1 | |
Chiang Mai, Thailand |
|
KR705658 | KR705596 | KR705720 | 1 | |
Maehongson, Thailand |
|
KR705657 | KR705595 | KR705719 | 1 | |
Maehongson, Thailand |
|
KR705656 | KR705594 | KR705718 | 1 | |
Sakon Nakhon, Thailand |
|
KR705655 | KR705593 | KR705717 | 1 | |
Mahasarakarm, Thailand |
|
KR705651 | KR705589 | KR705713 | 1 | |
Loei, Thailand |
|
KR705653 | KR705591 | KR705715 | 1 | |
Ubon Ratchathani, Thailand |
|
KR705652 | KR705590 | KR705714 | 1 | |
Phatthalung, Thailand |
|
KR705641 | KR705579 | KR705703 | 1 | |
Nakhon Si Thammarat, Thailand |
|
KR705639 | KR705577 | KR705701 | 1 | |
Ranong, Thailand |
|
KR705637 | KR705575 | KR705699 | 1 | |
Phang Nga, Thailand |
|
KR705640 | KR705578 | KR705702 | 1 | |
Chumphon, Thailand |
|
KR705638 | KR705576 | KR705700 | 1 | |
Uthai Thani, Thailand |
|
KR705632 | KR705570 | KR705694 | 1 | |
Prachuap Khiri Khan, Thailand |
|
KR705628 | KR705566 | KR705690 | 1 | |
Kanchanaburi, Thailand |
|
KR705631 | KR705569 | KR705693 | 1 | |
Ratchaburi, Thailand |
|
KR705630 | KR705568 | KR705692 | 1 | |
Tak, Thailand |
|
KR705629 | KR705567 | KR705691 | 1 | |
Siem Reap, Cambodia |
|
KR705687 | KR705625 | KR705749 | 1 | |
Srisophon, Cambodia |
|
KR705686 | KR705624 | KR705748 | 1 | |
Attapue, Laos |
|
KR705678 | KR705616 | KR705740 | 1 | |
Champasak, Laos |
|
KR705673 | KR705611 | KR705735 | 1 | |
Luang Prabang, Laos |
|
KR705677 | KR705615 | KR705739 | 1 | |
Phongsaly, Laos |
|
KR705676 | KR705614 | KR705738 | 1 | |
Perak, Malaysia |
|
KR705669 | KR705607 | KR705731 | 1 | |
Kelantan, Malaysia |
|
KR705668 | KR705606 | KR705730 | 1 | |
Perak, Malaysia |
|
KR705667 | KR705605 | KR705729 | 1 | |
Scolopendra multidens Newport, 1844 | Qiang Binh, Vietnam |
|
KF676540 | KF676485 | – | 3 |
Scolopendra calcarata Porat, 1876 | Kanchanaburi, Thailand |
|
KR705650 | KR705588 | KR705712 | 1 |
Wat Mae Long, Mae Chaem, Chiang Mai, Thailand |
|
KU512629 | KU512632 | KU512635 | This study | |
Lan Sang Waterfall, Mueang, Tak, Thailand |
|
KU512630 | KU512633 | KU512636 | This study | |
Scolopendra japonica Koch, 1878 | Matsumoto, Japan |
|
KR705679 | KR705617 | KR705741 | 1 |
Xieangkhuang, Laos |
|
KR705671, KR705670 | KR705609, KR705608 | KR705733, KR705732 | 1 | |
Phongsaly, Laos |
|
KR705675, KR705674 | KR705613, KR705612 | KR705737, KR705736 | 1 | |
Scolopendra pinguis Pocock, 1891 | Bo Kaeo, Laos |
|
KR705646 | KR705584 | KR705708 | 1 |
Kanchanaburi, Thailand |
|
KR705646 | KR705584 | KR705708 | 1 | |
Nan, Thailand |
|
KR705644 | KR705582 | KR705706 | 1 | |
Xieangkhuang, Laos |
|
KR705643 | KR705581 | KR705705 | 1 | |
Huaphun, Laos |
|
KR705642 | KR705580 | KR705704 | 1 | |
Chiang Mai, Thailand |
|
KR705649 | KR705587 | KR705711 | 1 | |
Mae Hong Son, Thailand |
|
KR705648 | KR705586 | KR705710 | 1 | |
Scolopendra dawydoffi Kronmüller, 2012 | Trad, Thailand |
|
KR705680 | KR705618 | KR705742 | 1 |
Nakhon Ratchasima, Thailand |
|
KR705654 | KR705592 | KR705716 | 1 | |
Nakhon Ratchasima, Thailand |
|
KR705635, KR705634 | KR705573, KR705572 | KR705697, KR705696 | 1 | |
Scolopendra cataracta sp. n. | Tad E-tu Waterfall, Bolaven Plateau, Pakse, Champasak, Laos | Holotype |
KR705672 | KR705610 | KR705734 | 1 |
Tad-Yueang Waterfall, Mueang Singh, Luang Namtha, Laos | Paratype |
KR705633 | KR705571 | KR705695 | 1 | |
Kao Sok National Park, Surat Thani, Thailand | Paratype |
KU512631 | KU512634 | KU512637 | This study | |
Cormocephalus monteithi Koch, 1983 | Queensland, Australia |
|
HM453309.1 | AF370861.1 | HM453274 | 3 |
Digitipes kalewaensis Siriwut, Edgecombe & Panha, 2015 | Kalewa, Sagaing, Burma |
|
KP204116 | KP204112 | – | 2 |
Otostigmus astenus Kohlrausch, 1878 | Fiji / Vanuatu |
|
HM453312 | HM453221 | – | 3 |
Sterropristes violaceus Muadsub & Panha, 2012 | Similan, Thailand |
|
KF676519 | KF676477 | – | 3 |
The phylogenetic tree from the updated concatenated DNA dataset aggregates studied specimens into eleven monophyletic groups within Scolopendrinae that are compatible with morphological identification (Fig.
Maximum likelihood tree for Scolopendra in mainland Southeast Asia: colours for clades correspond to species and outgroups; black and white circles indicate statistical support values in both ML and BI analyses or only ML or BI analysis, respectively. Numbers at nodes are bootstrap support and posterior probability. Specimen codes in parentheses following localities correspond to
Molecular marker | Length | Parsimony informative sites | Variable sites | Conserved sites |
---|---|---|---|---|
COI | 611 | 260 | 302 | 309 |
16S | 446 | 214 | 271 | 175 |
28S | 683 | 149 | 188 | 450 |
Genetic distance between Scolopendra species in mainland Southeast Asia and outgroups; upper right and lower left distance collected from COI and 16S partial gene pairwise comparisons.
Species | 16S | ||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | ||
COI | S. dehaani 1 | 0.143 | 0.159 | 0.148 | 0.216 | 0.216 | 0.210 | 0.159 | 0.142 | 0.185 | 0.253 | 0.259 | 0.265 | 0.294 | |
S. cataracta 2 | 0.159 | 0.164 | 0.138 | 0.238 | 0.213 | 0.199 | 0.156 | 0.158 | 0.175 | 0.259 | 0.262 | 0.288 | 0.300 | ||
S. dawydoffi 3 | 0.171 | 0.190 | 0.164 | 0.209 | 0.220 | 0.217 | 0.123 | 0.127 | 0.083 | 0.222 | 0.292 | 0.285 | 0.293 | ||
S. subspinipes 4 | 0.167 | 0.194 | 0.177 | 0.229 | 0.238 | 0.212 | 0.162 | 0.166 | 0.174 | 0.257 | 0.271 | 0.303 | 0.291 | ||
S. pinguis 5 | 0.219 | 0.247 | 0.215 | 0.224 | 0.171 | 0.240 | 0.246 | 0.230 | 0.233 | 0.234 | 0.280 | 0.298 | 0.322 | ||
S. calcarata 6 | 0.228 | 0.240 | 0.212 | 0.230 | 0.219 | 0.241 | 0.237 | 0.223 | 0.250 | 0.255 | 0.270 | 0.278 | 0.315 | ||
S. morsitans 7 | 0.204 | 0.230 | 0.204 | 0.233 | 0.244 | 0.242 | 0.214 | 0.215 | 0.231 | 0.274 | 0.323 | 0.341 | 0.290 | ||
S. japonica 8 | 0.202 | 0.227 | 0.163 | 0.208 | 0.218 | 0.219 | 0.232 | 0.123 | 0.135 | 0.258 | 0.287 | 0.297 | 0.307 | ||
S. cingulata 9 | 0.188 | 0.197 | 0.177 | 0.214 | 0.224 | 0.228 | 0.221 | 0.181 | 0.131 | 0.261 | 0.274 | 0.291 | 0.322 | ||
S. multidens 10 | 0.196 | 0.203 | 0.107 | 0.178 | 0.216 | 0.214 | 0.229 | 0.163 | 0.188 | 0.230 | 0.304 | 0.326 | 0.323 | ||
Cormocephalus 11 | 0.236 | 0.263 | 0.234 | 0.238 | 0.269 | 0.239 | 0.267 | 0.243 | 0.231 | 0.233 | 0.266 | 0.296 | 0.322 | ||
Digitipes 12 | 0.234 | 0.216 | 0.227 | 0.255 | 0.249 | 0.252 | 0.289 | 0.260 | 0.237 | 0.229 | 0.279 | 0.213 | 0.246 | ||
Sterropristes 13 | 0.234 | 0.246 | 0.228 | 0.249 | 0.233 | 0.240 | 0.245 | 0.217 | 0.232 | 0.242 | 0.244 | 0.205 | 0.199 | ||
Otostigmus 14 | 0.217 | 0.237 | 0.222 | 0.237 | 0.250 | 0.232 | 0.251 | 0.248 | 0.267 | 0.236 | 0.261 | 0.219 | 0.208 |
Genetic distance under pairwise sequence comparison within populations of Scolopendra species in mainland Southeast Asia.
Species | COI | 16S |
---|---|---|
Scolopendra dehaani | 0.083 | 0.052 |
Scolopendra cataracta | 0.165 | 0.087 |
Scolopendra dawydoffi | 0.019 | 0.010 |
Scolopendra subspinipes | 0.146 | 0.109 |
Scolopendra pinguis | 0.184 | 0.113 |
Scolopendra calcarata | 0.114 | 0.072 |
Scolopendra morsitans | 0.086 | 0.068 |
Scolopendra japonica | 0.128 | 0.061 |
Three main clades are identified in mainland Asian Scolopendra (Fig.
The remaining Scolopendra species may be divided into two groups, one consisting of S. morsitans and another including former subspecies of S. subspinipes sensu
All samples of S. cataracta united as a clade with S. dehaani to the exclusion of S. subspinipes (Fig.
In this region, nine species are identified from our survey. The taxomomic boundaries between species were based on information from both morphology and molecular analysis. Two other species of Scolopendra were not included in this paper, namely S. mirabilis (Porat, 1876), and S. hardwickei Newport, 1845. In the case of S. mirabilis, an African-central Asian species, the single known specimen in SE Asia may be introduced, being found on an island in a coastal area of northern Vietnam (
1 | Tergite of ultimate leg-bearing segment with median suture | S. morsitans Linnaeus, 1758 |
– | Tergite of ultimate leg-bearing segment without median suture | 2 |
2 | Sternal paramedian sutures complete | 3 |
– | Sternal paramedian sutures incomplete | 4 |
3 | Coxopleural process with three or more apical spines, thick prefemur of ultimate leg with at least two ventro-lateral and four spines on prefemoral process; average ratio of width:length of ultimate leg prefemur1:2 | S. japonica Koch, 1878 |
– | Coxopleural process with 1–2 apical spines, slender prefemur of ultimate leg with two ventro-lateral and two spines on prefemoral process; ratio of width:length of ultimate leg prefemur 1:3 | S. subspinipes Leach, 1816 |
4 | Ultimate leg prefemur with at least one ventro-lateral spine | 5 |
– | Ultimate leg prefemur without ventro-lateral spines | S. dehaani Brandt, 1840 |
5 | Coxopleural process with one lateral and one dorsal spine; ultimate leg prefemur with numerous small scattered spines | 6 |
– | Coxopleural process without lateral and dorsal spines; ultimate leg prefemur with a few enlarged spines in rows | 8 |
6 | Legs 21 with a tarsal spur | S. calcarata Porat, 1876 |
– | Leg 21 without tarsal spurs | 7 |
7 | Sternites of anterior body segments with complete paramedian sutures | S. gracillima sternostriata Schileyko, 1995 |
– | All sternites with incomplete paramedian sutures, reaching only 20–30% on anterior part of sternites | S. pinguis Pocock, 1891 |
8 | Complete paramedian sutures on tergites | 9 |
– | Short, incomplete paramedian sutures confined to anterior and posterior parts of tergites | S. cataracta Siriwut, Edgecombe & Panha, sp. n. |
9 | Cephalic plate and tergite 1 densely punctate; tergites with short median sulcus on posterior part | S. multidens Newport, 1844 |
– | Cephalic plate and tergite 1 sparsely punctate; tergites without median sulcus on posterior part | S. dawydoffi Kronmüller, 2012 |
Scolopendra
morsitans
Linnaeus, 1758: 638.
Scolopendra
brandtiana
Gervais, 1837: 16.
Scolopendra
bilineata
Brandt, 1840: 155.
Scolopendra
crassipes
Brandt, 1840: 153.
Scolopendra
erythrocephala
Brandt, 1840: 155.
Scolopendra
limbata
Brandt, 1840: 154.
Scolopendra
platypus
Brandt, 1840: 153.
Scolopendra
elegans
Brandt, 1841: 21.
Scolopendra
fulvipes
Brandt, 1841: 22.
Scolopendra
morsitans
scopoliana
C.L. Koch, 1841: 222, pl. 11.
Scolopendra
angulipes
Newport, 1844: 97.
Scolopendra
leachii
Newport, 1844: 97.
Scolopendra
longicornis
Newport, 1844: 97.
Scolopendra
platypoides
Newport, 1844: 97.
Scolopendra
tuberculidens
Newport, 1844: 97.
Scolopendra
algerina
Newport, 1845: 387.
Scolopendra
fabricii
Newport, 1845: 384.
Scolopendra
formosa
Newport, 1845: 383.
Scolopendra
richardsoni
Newport, 1845: 385.
Scolopendra
tigrina
Newport, 1845: 381.
Scolopendra
varia
Newport, 1845: 380.
Scolopendra
tongana
Gervais, 1847: 275.
Scolopendra
infesta
Koch, 1847: 169.
Scolopendra
planipes
Koch, 1847: 168.
Scolopendra
pella
Wood, 1861: 13.
Scolopendra
porphyratainia
Wood, 1861: 15.
Scolopendra
brachypoda
Peters, 1862: 529, pl. 33, fig. 2.
Scolopendra
mossambica
Peters, 1862: 527, pl. 33, fig. 4.
Scolopendra
compressipes
Wood, 1862: 31.
Scolopendra
modesta
Wood, 1862: 29.
Scolopendra
carinipes
Humbert & Saussure, 1870: 204.
Scolopendra
afzelii
Porat, 1871: 1146.
Scolopendra
attenuata
Porat, 1871: 1148.
Scolopendra
chlorocephala
Porat, 1871: 1149.
Scolopendra
cognata
Porat, 1871: 1145
Scolopendra
intermedia
Porat, 1871: 1145.
Scolopendra
picturata
Porat, 1871: 1144.
Scolopendra
pilosella
Porat, 1871: 1148.
Scolopendra
saltatoria
Porat, 1871: 1151.
Scolopendra
vaga
Porat, 1871: 1151.
Scolopendra
wahlbergi
Porat, 1871: 1150.
Eurylithobius
slateri
Butler, 1876: 446.
Scolopendra
impressa
Porat, 1876: 12.
Scolopendra
morsitans
procera
Haase, 1887: 53, pl. 33, fig. 53.
Scolopendra
morsitans
sulcipes
Haase, 1887: 54, pl. 33, fig. 54.
Scolopendra
morsitans
calcarata
Daday, 1891: 150.
Scolopendra
grandidieri
Saussure & Zehntner, 1902: 302, pl. 3, fig. 13, pl. 12, fig. 6.
Scolopendra
lineata
Saussure & Zehntner, 1902: 308, pl. 15, fig. 19.
Scolopendra
spinosella
Saussure & Zehntner, 1902: 308, pl. 2, fig. 11.
Scolopendra
morsitans
fasciata
Attems, 1930a: 372.
Scolopendra
morsitans
amazonica
Bücherl, 1946: 135.
Trachycormocephalus
jodhpurensis
Khanna, 1977: 154, figs 5–8.
India.
Thailand —
Cambodia —
Laos —
Myanmar —
Vietnam —
Brunei —
Philippines —
Indonesia —
China —
Japan —
Indian and Middle Asian Territory —
Africa —
Australia —
South Pacific —
Undetermined locality —
17–23 antennal articles, 5–8 basal articles glabrous dorsally. Each tooth-plate with 5–6 teeth. Tergites 7(12)-20 with paramedian sutures. Tergite of ultimate leg-bearing segment with median suture. Complete paramedian sutures on sternites 2–20. Coxopleural process with 3–4 apical and 0–1 lateral spines. Ultimate leg prefemora with three rows of ventral spines (2–6 VL, 3 V, 2–6 VM), 2–6 M, 2–6 DM and 0–8 spines on prefemoral process. One tarsal spur on legs 1–19 (in Southeast Asia).
Body length up to 12.7 mm (In Australian populations according to
Antenna usually with 18–20 articles (sometimes 17, 21 or 23 on one side), basal 5–7 glabrous dorsally (Figs
Anterior margin of T1 underlying cephalic plate (Fig.
Scolopendra morsitans (
Coxopleural process moderately long or short, usually with 4–5 apical and 0–1 lateral spines (Fig.
All legs without setae and tibial spurs. One tarsal spur on legs 1–19 (20 in some African and Indian populations). Ultimate legs: thick and moderately long, with ratios of lengths of prefemur and femur 1.2:1, femur and tibia 1.3:1, tibia and tarsus 2 1.7:1.; tarsus 1 and tarsus 2 2.8:1. In male, lateral margin of prefemora, femora and tibia marginated dorsally. Prefemoral spines (Figs
Genital segments well developed, reaching longer than the distance between posterior margin of sternite of ultimate leg-bearing segment and distal part of coxopleural process (Fig.
Colouration. Scolopendra morsitans demonstrates colour variation among its populations in SE Asia. Previously, colour variation has been recorded in African, Australian and Taiwanese populations (
Colour morph 1. Dichromatic. Cephalic plate, T1 and tergite of ultimate leg-bearing segment orange, the remaining tergites brownish. Posterior borders and lateral margins of tergites dark. Antenna bright orange. Pleuron of leg-bearing segments with pale grey integument, pleurites orange. Legs 1–21 orangish or yellow. Ultimate legs orangish or light brown.
Colour morph 2. Dichromatic. Cephalic plate, T1 and tergite of ultimate leg-bearing segment dark brown or blackish, the remaining tergites brownish. Posterior borders and lateral margins of tergites dark. Antenna dark blue. Pleuron of leg-bearing segments with pale grey integument, pleurites orange or brown. Legs 1–20 yellowish or pale. Ultimate legs blackish or brown.
Scolopendra morsitans is morphologically varied and subsumes many synonyms that are now attributed to geographical and/or ontogenetic variation. Intraspecific variation has been studied in Africa (
Geographical variation in several populations of Scolopendra morsitans in Old World territory including Australia. ? insufficient data.
Character | India3 | Burma1 | Indochina1,2 | Malay Archipelago1,2 | Philippines1 | East Asia1,6 | Australia5 | Africa2,4 |
---|---|---|---|---|---|---|---|---|
Number of antennal articles | 19–20 | 18–22 | 20–21 | 18–20 | 17–19 | 18–20 | 17–23 | 17–21 |
Number of glabrous articles | 6–9 | 6–7 | 5 | 6–7 | 6–7 | 5–7? | 3–8 | 5–7 |
Teeth on tooth-plate | 5 | 4–5 | 3–7 | 4–5 | 4–5 | 5 | 3–6 | 5 |
First tergite with complete paramedian sutures | 2 | 3 | 4–5 | 2–3 | 2–3 | 3 | 2–4 | 2–4 |
First tergite with margination | 7–17 | 6–17 | 5–14 | 6–13 | 12–13 | 10–14 | 5 | 2–7 (15) |
Tergite surface | smooth | smooth | smooth | smooth | smooth | smooth | smooth | smooth |
Median furrow on tergite of ULBS | present | present | present | present | present | present | present | present |
Paramedian sutures on sternites | incomplete | incomplete | incomplete | incomplete | incomplete | incomplete | incomplete | incomplete |
Sternite of ULBS | ? | without depression | without depression | without depression | without depression | without depression | without depression | ? |
Spines on coxopleural process |
AP: 3–5 LS: 0–1 |
AP: 3–5 LS: 1 |
AP: 1–5 SAP: 0–1 LS: 0–1 |
AP: 3–5 LS: 0–1 |
AP: 4–5 LS: 0–1 |
AP: 4 LS: 0–1 |
AP: 2–6 |
AP: 2–6 LS: 0–1 |
Spine formula on prefemora of ultimate legs | VS: 3 rows DS: 4–5 2 rows (4–5) SP: 4–10 |
V: 5–10 (3 rows) M: 2–4 DM: 2–4 SP: 3–7 |
V: 5–10 (3 rows) M: 0–6 DM: 0–5 SP: 2–6 |
V: 5–10 (3 rows) M: 2–4 DM: 2 SP: 3–7 |
V: 9–10 (3 rows) M: 2–4 DM: 2 SP: 3–7 |
V: 7–9 (3 rows) M: 0–3 DM: 2–6 SP: 0–4 |
VL: 2–6 VM: 2–8 M: 2–6 DM: 2–6 SP: 4–8 |
VL: 6–12 M: 2–6 DM: 3–6 SP: 3–8 |
Legs with one tarsal spur | 1–19(20) | 1–19 | 1–19 | 1–19 | 1–19 | 1–19 | 1–19 | 1–19(20) |
Our phylogenetic analysis corroborates the monophyly of SE Asian populations of this species (Fig.
This is one of the oldest described centipede species and it is distributed worldwide in the tropics.The native distribution is difficult to determine because of assumed introduction in several areas.
Scolopendra
subspinipes
Leach, 1816: 383.
Scolopendra
audax
Gevais, 1837: 50.
Scolopendra
septemspinosa
Brandt, 1840: 152.
Scolopendra
borbonica
Blanchard, 1829: 7, pl. 1.
Scolopendra
sexspinosa
Newport, 1844: 96,
Rhombocephalus
gambiae
Newport, 1845: 392.
Scolopendra
ceylonensis
Newport, 1845: 391.
Scolopendra
flava
Newport, 1845: 392.
Scolopendra
gervaisii
Newport, 1845: 390.
Scolopendra
lutea
Newport, 1845: 392.
Scolopendra
ornata
Newport, 1845: 392.
Scolopendra
placeae
Newport, 1845: 390.
Scolopendra
planiceps
Newport, 1845: 391.
Scolopendra
rarispina
Gervais, 1847: 270.
Scolopendra
sandwichiana
Gervais, 1847: 276.
Scolopendra
mactans
Koch, 1847: 16,
Scolopendra
sulphurea
Koch, 1847: 156,
Scolopendra
byssina
Wood, 1861: 10.
Scolopendra
cephalica
Wood, 1861: 12.
Scolopendra
cephalica
gracilis
Wood, 1861: 13.
Scolopendra
dinodon
Wood, 1861: 12.
Scolopendra
gracilipes
Wood, 1861: 12.
Scolopendra
parvidens
Wood, 1861: 13.
Scolopendra
plumbeolata
Wood, 1861: 14.
Scolopendra bispinipes Wood, 1862: 28. Brölemann 1909: 25.
Scolopendra
nesuphila
Wood, 1862: 31.
Scolopendra
repens
Wood, 1862: 31.
Scolopendra
elongata
Porat, 1871: 1143.
Rhombocephalus
smaragdinus
Butler, 1876: 446.
Scolopendra
damnosa
Koch, 1878: 789.
Scolopendra
mutilans
Koch, 1878: 791. Haase 1881: 47, pl. 3, fig. 47.
Scolopendra
aurantiipes
Tömösváry, 1885: 67.
Scolopendra
variispinosa
Tömösváry, 1885: 67.
Scolopendra
rugosa
Meinert, 1886: 202.
Scolopendra
meyeri
Haase, 1887: 49, pl. 3, fig. 50.
Scolopendra
macracanthus
Bollman, 1889: 213.
Scolopendra
flavicornis
Tömösváry, 1885: 67.
Scolopendra
subspinipes
gracilipes
Daday, 1891: 149.
Scolopendra
subspinipes
molleri
Verhoeff, 1892: 199.
Scolopendra
polyodonta
Daday, 1893: 5.
Scolopendra
machaeropus
Attems, 1901: 136.
Scolopendra
aringensis
Sinclair, 1901: 529, pl. 31, fig. 46, pl. 32, Figs 67, 85, 86, 93.
Scolopendra
subspinipes
mutilans
Kraepelin, 1903: 263.
Scolopendra
subspinipes
gastroforeata
Muralewicz, 1913: 201.
Scolopendra
subspinipes
piceoflava
Attems, 1934: 51.
Scolopendra
subspinipes
fulgurans
Bücherl, 1946: 148,
Not designed. The whereabouts of the holotype are unknown.
Specimens referred to S. subspinipes Leach, 1816: Malaysia —
Singapore —
Indonesia —
Philippines —
China —
Japan —
South Asia —
Africa —
Jamaica —
Madagascar —
Rodrigues —
Mauritius —
Seychelles —
Comoros —
Pacific Islands —
United Kingdom —
Caribbean Sea —
Central and South America —
Undetermined locality —
Specimens referred to S. mutilans Koch, 1878: Japan — Syntype
China —
Korea —
Undetermined locality —
17–19 antennal articles, 6 basal articles glabrous dorsally. Each tooth-plate with (4)5-7 teeth. Tergites 3(4)-20 with paramedian sutures. Complete tergite margination on TT14 (17)-21. Tergite of ultimate leg-bearing segment without depression or median suture. Complete paramedian sutures on sternites 2(3)-20. Coxopleural process with 2 apical spines, without lateral and dorsal spine. Ultimate leg prefemora with 2 VL, 1–2 M, 0–3 DM and prefemoral process with 1–6 spines. One tarsal spur on legs 1–19 or 20.
Body length up to 16 cm. Reddish brown colouration on entire body. Cephalic plate and segments monochromatic or dichromatic. Tergites reddish brown; dark band on posterior border of tergites. Cephalic plate with small punctae on anterior part; median sulcus present. Posterior part of cephalic plate without paramedian sulci.
Antenna usually with 18–19 articles (16–17 articles on one side in some specimens), basal 6 subcylindrical and glabrous dorsally (Fig.
Anterior margin of T1 underlying cephalic plate (Figs
Coxopleural process moderately long, with two apical and 0–1 subapical spines; pore-free area extending 30–70% length from distal part of coxopleural process to margin of sternite of ultimate leg-bearing segment (Figs
All legs without setae and tibial spur. One tarsal spur on legs 1–19 or more commonly 1–20. Ultimate legs: moderately long and slender, with ratios of lengths of prefemur and femur 1.4:1, femur and tibia 1.2:1, tibia and tarsus 2 1.4:1.; tarsus 1 and tarsus 2 2:1. Prefemoral spines: 2 VL, 1–2 M, 0–3 DM and prefemoral process with 1–6 spines (Figs
Genital segments well developed, reaching longer than distance between posterior margin of sternite of ultimate leg-bearing segment and distal part of coxopleural process. Sternite of genital segment 1 round and convex posteriorly, with median suture. In male, sternite of genital segment 2 attached to penis. Tergite of genital segment without small setae. Gonopods with small setae in male. Penis with apical bristle.
Recently, the taxonomic validity of S. subspinipes and its former subspecies has been evaluated both by morphology (
Scolopendra subspinipes (Syntype
Morphological comparison of Scolopendra subspinipes populations from different geographical regions. ? insufficient data.
Character | Bay of Bengal and Indian Ocean1 | Malay Peninsula1 | Vietnam 2 | Indonesia1 | Philippines1 | Taiwan 3 | China1 | Japan1 |
---|---|---|---|---|---|---|---|---|
Number of antennal articles | 18 | 18–19 | 18–19 | 17–19 | 17–18 | 18–19 | 16–18 | 15–19 |
Number of glabrous articles | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 |
Teeth on tooth-plate | 5+5 | 4+6, 5+5, 6+5, 6+6, 7+7 | 4–9 (each side?) | 5+5, 6+7, 7+7, 6+10 | 5+5, 10+5, 7+7 | 5+5 | 5+5 | 4+5, 5+5, 6+5, 6+6 |
First tergite with complete paramedian sutures | 3 | 3 | 2(9) (poorly defined in some specimens) |
3–4 | 3–4 | 4–6 | 3–6 | 3–6 |
First tergite with margination | 5 | 4–10 | 14–15 | 4–11 | 3–5 | 5–9 | 5–8 | 5–16 |
Tergite surface | smooth | smooth | smooth | smooth | smooth | smooth | smooth | smooth |
Median furrow on tergite of ULBS | absent | absent | absent | absent | absent | absent | absent | absent |
Paramedian sutures on sternites | complete | complete | complete | complete | complete | incomplete | complete | complete |
Sternite of ultimate leg-bearing segment | without pit | pit-like median furrow | ? | pit-like median furrow | without pit | ? | pit-like median furrow | pit-like median furrow |
Spines on coxopleural process | AP: 2 |
AP: 2 SAP: 0–1 |
AP: 2 |
AP: 2 SAP: 0–1 |
AP: 2 SAP: 0–1 |
AP: 2–3 |
AP: 1–2 SAP: 0–1 |
AP: 0–2 SAP: 0–1 |
Spine formula on prefemora of ultimate legs |
VL: 2 M: 1 DM: 1 SP: 2 |
VL: 2–3 M: 1–2 DM: 1–2 SP: 2–3 |
VL: 0–3 M: 0–2 DM: 0–3 SP: 1≥ |
VL: 2 M: 1–2 DM: 1–2 SP: 2–5 |
VL: 2 M: 1–2 DM: 1–2 SP: 2 |
VL: 2 VM: 1 DM: 2 SP: 2–4 |
VL: 2–3 M: 1 DM: 1 SP: 1–2 |
VL: 0–3 M: 0–2 DM: 0–2 SP: 1–6 |
Legs with one tarsal spur | 1–20(R) | 1–19(20) | 1–19 | 1–19 (20) | 1–19 (20) | 1–20 | 1–20 | 1–19(20) |
S. subspinipes piceoflava, another former subspecies of S. subspinipes from Sulawesi, Indonesia, is currently treated as a synonym of S. subspinipes (see
Syntypes
Scolopendra subspinipes (Syntypes
Tomohon, Sulawesi, Indonesia [Tomohon, North Sulawesi Province, Indonesia].
Body length 16.7 cm in male and 17.1 and 16.5 cm in female syntypes. Preserved male still exhibiting traces of its colouration pattern: cephalic plate and segments dark greenish or brown. Antenna yellowish. Tergites with yellowish or pale colour on posterior margin. All legs blue-greenish, distal part yellow. Cephalic plate without small punctae on anterior part, median sulcus present. Posterior part of cephalic plate without paramedian sulci.
Antenna usually with 17–19 articles, basal 6 subcylindrical and glabrous both on dorsal and ventral sides. Antennae reach segment 4 (Fig.
Anterior margin of T1 underlying cephalic plate (Fig.
Coxopleural process long (Fig.
All legs without setae and tibial spur. One tarsal spur on legs 1–20. Ultimate legs: slender and long (Fig.
Sternite of genital segment 1 round and convex posteriorly, with median suture. In male, sternite of genital segment 2 attached to penis. Tergite of genital segment without small setae. Gonopods present in male.
Based on examination of the syntypes, we corroborate the assignment of this nominal subspecies to the S. subspinipes group. Some morphological characters that appear, however, not to be identical with S. subspinipes are the sharpness and length of the coxopleural process, which bears one or two strong apical spines, the ratio of ultimate leg podomeres, and the colouration pattern on the tergites that is clearly distinct from other geographical populations of S. subspinipes (the posterior part of the tergites exhibiting a yellowish colouration). On the other hand, the syntypes of S. subspinipes piceoflava also display morphological variation between each other with respect to the number of prefemoral spines on the ultimate legs: a male specimen has 4–6 spines on the prefemoral process whereas VL, M and DM spines are absent in one female specimen. The latter is similar to S. dehaani but it is possible that this absence may be due to regeneration in this individual. However, without additional material and lacking molecular data with which to test relationships among morphological similar species, we tentatively accept S. subspinipes piceoflava as a junior synonym of S. subspinipes as proposed by
Previous studies regarded S. subspinipes s.l. as a cosmopolitan species in tropical regions (
Scolopendra
dehaani
Brandt, 1840: 152,
Scolopendra
subspinipes
dehaani
— Pocock 1891: 409,
Scolopendra
subspinipes
—
Scolopendra
childreni
Newport, 1844: 96.
Scolopendra
concolor
Newport, 1845: 394.
Scolopendra
inermis
Newport, 1845: 393.
Scolopendra
inermipes
Koch, 1847: 153.
Scolopendra
silhetensis
Newport, 1845: 393.
Scolopendra
lucasii
Gervais, 1847: 270.
Scolopendra
horrida
Koch, 1847: 154.
Scolopendra
limicolor
Wood, 1861: 12.
Scolopendra
bispinipes
Wood, 1862: 28.
Scolopendra
fissispina
Koch, 1865: 891.
Scolopendra
nudipes
Tömösváry, 1885: 67.
Scolopendra
foveolata
Verhoeff, 1937: 220.
Scolopendra arborea Lewis, 1982: 389. Lewis 2010: 97. Syn. nov.
Java, Indonesia.
Thailand —
Laos —
Cambodia —
Myanmar —
Malaysia —
Singapore —
Indonesia —
China —
Indian Territory —
Mexico —
Undetermined locality —
17–21 antennal articles, 6 basal articles glabrous dorsally. Each tooth-plate with 5–6 teeth. Tergites 3(4)–20 with paramedian sutures. Complete tergite margination from T7. Tergite of ultimate leg-bearing segment without depression or suture. Complete or incomplete paramedian sutures on sternites. Coxopleural process with two apical spines, absent lateral and dorsal spines. Ultimate leg prefemora with 0–1 M, 0–1 DM and prefemoral process with 1–4 spines. One tarsal spur on legs 1–19(20).
Body length up to more than 25 cm (collections from Java,
Antenna usually with 18 articles (17, 19 and 21 articles in some specimens), basal 6 subcylindrical and glabrous dorsally (Fig.
Brooding and feeding behaviours: A Scolopendra dehaani exhibiting simple coiling with cluster of embryonic stadia (photograph by Natdanai Likhitrakarn) B Scolopendra morsitans exhibiting double coiling with post-embryonic stadia C–D Scolopendra dehaani preying on snail-eating snake Pareas carinatus D Flexibility of trunk segments during predation.
Anterior margin of T1 underlying cephalic plate (Fig.
Coxopleural process moderately long or short with two apical spines (atypically 0–1 spines); pore-free area extending 90–100% length from distal part of coxopleural process to margin of sternite of ultimate leg-bearing segment (Fig.
All legs without setae and tibial spur. One tarsal spur on legs 1–19 or 20 in equal frequency. Ultimate legs: moderately long and slender, with ratios of lengths of prefemur and femur 1.1:1, femur and tibia 1.2:1, tibia and tarsus 2 1.6:1.; tarsus 1 and tarsus 2 2.3:1. Prefemora long and slender, flattened dorsally (Fig.
Genital segments well developed, reaching longer than distance between posterior margin of sternite of ultimate leg-bearing segment and distal part of coxopleural process. Sternite of genital segment 1 round and convex posteriorly, with median suture (Figs
Colouration. This species is among those that exhibited the most varied colouration patterns.
Colour morph 1: Dichromatic. All segments including cephalic plate dark brownish. Posterior border of tergites with a dark band. Antenna reddish brown. Pleuron with pale grey integument, all pleurites brownish. Legs chestnut brown, tibiae and tarsi dark purplish.
Colour morph 2: Dichromatic. All segments brown or yellowish orange. Posterior border of tergites with a dark band. Antenna yellowish orange. Pleuron with pale grey integument, pleurites pale grey. Legs dark brownish, tibiae and tarsi dark purplish.
Colour morph 3: Monochromatic. All segments including cephalic plate reddish brown. Antenna reddish brown. Pleuron with pale grey integument, pleurites brownish. Legs 1–18 yellowish, tibiae and tarsi reddish brown. Leg 20 and ultimate legs entirely reddish brown.
Colour morph 4: Dichromatic. Cephalic plate, TT1–2 and 19–21 bright reddish, rest of tergites brown. Posterior border of tergites with a dark band. Antenna yellowish or bright orange. Pleuron with pale grey integument, pleurites orange. Legs 1–19 yellowish, without secondary colouration on distal part. Leg 20 and ultimate legs entirely reddish.
Colour morph 5: Dichromatic. All segments including cephalic plate cherry reddish. Posterior border of tergites with a dark band. Antenna reddish or orange. Pleuron with grey integument, pleurites orange. All legs reddish.
This is the largest centipede in Asia. A consistent character that is treated as diagnostic for this species is the absence of ventral spines on the ultimate leg prefemur. Scolopendra dehaani possesses characters of the S. subspinipes s.l. sensu Lewis (2010) but after morphological survey (
Morphotype 1: Complete paramedian sutures on tergites and sternites. This is the typical form of S. dehaani, according to
Morphotype 2: Paramedian sutures complete on tergites, confined to 20–30% length of sternites. This morphotype is observed in a specimen apparently from Bangladesh.
Morphotype 3: Paramedian sutures lacking on tergites on all segments (Fig.
Based on these morphotypes, we surmise that this variability might have a geographic basis and could suggest evidence of cryptic speciation. Molecular data are presently lacking for morphotype 3 in particular and, presently, we apply the specific name S. dehaani Brandt, 1840, throughout the entire geographic range. A morphological comparison through the species’ geographical range is given in Table
Morphological survey of geographical variation of Scolopendra dehaani populations from different regions.
Characters | Geographical distribution area | ||||||
---|---|---|---|---|---|---|---|
Java1,2 | Thailand, Laos and Cambodia1 | Burma1 | Malay Peninsula1 | India3 | China- Japan1 | Mexico1 | |
Number of antennal articles | 14–18 | 18–21 | 17–19 | 17–19 | 18–19 | 18 | 18 |
Number of glabrous articles | 6 | 6 | 6 | 6 | 6 | 6 | 6 |
Teeth on tooth-plate | 5+4,5+5 | 5+5 | 5+5,5+6,6+6 | 5+5,6+6,7+8,8+10 | 6+6 | 6+7 | 6+6 |
First tergite with complete paramedian sutures | incomplete | 4 | 2–6 | 3–4 | 4 | 3–5 or incomplete | incomplete |
First tergite with margination | 7–10 | 13 | 10–14 | 7–13 | 5 | 8 | 7 |
Tergite surface | smooth | smooth | smooth | smooth | smooth | smooth | smooth |
Median furrow on tergite of ULBS | absent | absent | absent | absent | absent | absent | absent |
Extent (percentage) of paramedian sutures on sternites | 10–15% (rarely with complete PS) | 90–100% | 100% | 30–100% | 80–100% | 80–100% | 20% |
Sternite of ultimate leg-bearing segment | with or without pit- like median depression | with pit- like median depression | with or without pit- like median depression | with or without pit-like median depression | without pit-like median depression | without pit-like median depression | without pit-like median depression |
Spines on coxopleural process |
AP: 2 SAP: 0–1 |
AP: 2 | AP: 2 | AP: 0–2 | AP: 2 | AP: 2 |
AP: 1 SAP: 0–1 |
Spinulation formula on prefemora of ult. legs |
M: 0–2 DM: 0–2 SP: 2–4 |
M: 1 DM: 1 SP: 2–3 |
M: 0–2 DM: 0–2 SP: 2 |
M: 1 DM: 1 SP: 1–3 |
DM: 1 SP: 2 |
M: 0–1 DM: 1–2 SP: 1–2 |
M: 0–1 DM: 1 SP: 2 |
Legs with one tarsal spur | 1–20 | 1–19 | 1–19 (20) | 1–19 (20) | 1–20 | 1–20 | 1–20 |
One new subjective synonym is proposed for S. dehaani. The holotype and sole known specimen of S. arborea Lewis, 1982 (
Widespread species in the Southeast Asian mainland and some islands (Fig.
Scolopendra
multidens
Newport, 1844: 97,
Scolopendra rugosa Meinert, 1886: 202.
Scolopendra
subspinipes
multidens
—
Not designated.
Holotype
17–18 antennal articles, 6 basal articles glabrous dorsally. Each tooth-plate with 5–10 teeth. Tergites 2(3)–20 with paramedian sutures. Complete tergite margination from TT12 (14)–21. Tergite of ultimate leg bearing segment without depression or suture. Paramedian sutures 20–60% on anterior part of sternites. Coxopleural process with 2–3 apical spines. Ultimate leg prefemora with 2 VL, 1 M, 1–2 DM and 1–3 spines on prefemoral process. One tarsal spur on legs 1–19.
Body length 11.4 cm in syntype. Dried holotype brownish on entire body. Cephalic plate with small punctae; median sulcus present. Posterior part of cephalic plate without paramedian sulci.
Antennae with 17–18 articles, basal 6 subcylindrical and glabrous dorsally on left side, 6 articles glabrous ventrally. Antennae reach segment 4. Forcipular trochanteroprefemoral process with denticles in two groups, one apical and three inner. Tooth-plates quadrate, with 5 teeth (Fig.
Anterior margin of T1 underlying cephalic plate (Fig.
Coxopleural process moderately long or short with 1–2 apical and 1–2 subapical spines; pore-free area extending 80% length from distal part of coxopleural process to margin of sternite of ultimate leg-bearing segment (Fig.
All legs without setae and tibial spur. One tarsal spur on legs 1–19; holotype lacking leg 20. Ultimate legs (Fig.
Genital segments well developed, reaching longer than distance between posterior margin of sternite of ultimate leg-bearing segment and distal part of coxopleural process (Fig.
Colouration. According to
Morphological characters are similar to S. subspinipes sensu Chao, 2008. The validity of S. multidens as a separate species was re-established by the absence of gonopods on the first genital segment in males (
Widespread species in Asia (Fig.
Scolopendra
calcarata
Porat, 1876: 10.
China.
Holotype:
Thailand —
17 antennal articles, 4–6 basal articles glabrous dorsally. Each tooth-plate with 5–8 teeth. Tergites 3–20 with paramedian sutures. Complete margination only on tergite of ultimate leg-bearing segment. Tergite of ultimate leg-bearing segment without depression or median suture. Incomplete paramedian sutures on sternites. Coxopleural process with 3–4 apical, 0–3 subapical and 0–1 lateral spines, without dorsal spine. Ultimate leg prefemora with 4–12 VL, 0–12 VM, 1–5 M, 2 DM, prefemoral process with 3–5 spines. One tarsal spur on legs 1–21.
Body length up to 5.3 cm. Blackish colouration on most of dorsal part of body. Cephalic plate dichromatic; anterior part of cephalic plate dark blue or black, posterior margin green yellowish (Fig.
Antenna usually with 17 articles, basal 4–6 subcylindrical, glabrous dorsally (Fig.
Anterior margin of T1 underlying cephalic plate (Figs
Coxopleural process moderately long, with 3–4 apical, 0–3 subapical and 0–1 lateral spines, without dorsal spine. Pore-free area extending 50–75% length from distal part of coxopleural process to margin of sternite of ultimate leg-bearing segment (Figs
All legs with small setae, without tibial spur. One tarsal spur on legs 1–21. Ultimate legs moderately long and slender, with ratios of lengths of prefemur and femur 1.2:1, femur and tibia 1.2:1, tibia and tarsus 2 1.4:1.; tarsus 1 and tarsus 2 2.6:1. Prefemoral spines (Figs
Genital segments well developed, reaching longer than distance between posterior margin of sternite of ultimate leg-bearing segment and distal part of coxopleural process. Sternite of genital segment 1 round and convex posteriorly, with median suture (Fig.
This montane species was sometimes collected together with other scolopendrids such as species of Otostigmus and Rhysida. External phenotypic characters are similar to S. pinguis but the unique, diagnostic character that permits species identification is the presence of a tarsal spur on the ultimate legs, which is atypical for Scolopendra. However, this character has been reported in some individuals of a few other Scolopendra species in Southeast Asia, notably S. subcrustalis (see
Morphological similarity between S. calcarata and S. pinguis is indicated by several characteristics, including the number of antennal articles, the shape of teeth on the forcipular tooth-plates (these being in the form of minute denticles), and the number of spines on the coxopleural process. In addition, the habitat preferences of these two species resemble each other, both of them being found only in montane territory, and they also show similar dichromatic colouration patterns. There is no evidence from our survey that these two species are distributed sympatrically. These characters are consistent with the molecular phylogeny, which resolves these two species as sister taxa.
This species is quite rare in tropical mainland Southeast Asia, usually distributed along mountain ranges in the western territory of Thailand (Fig.
Scolopendra
japonica
Koch, 1878: 790.
Scolopendra
subspinipes
japonica
–
Otostigmus
politoides
Attems, 1953: 147.
Otostigmus
puncticeps
Attems, 1953: 146, figs 16–17.
Japan.
Syntype:
Laos —
Japan —
Indonesia —
China —
Undetermined —
Diagnosis. 17–19 antennal articles, 6 basal articles glabrous dorsally. Each tooth-plate with 4–6 teeth. Tergites (3)4–20 with paramedian sutures. Complete margination from TT(10)12–21. Tergite of ultimate leg-bearing segment without depression or suture. Complete paramedian sutures on sternites 2–20. Coxopleural process with 3 apical spines. Ultimate leg prefemora with 2–3 VL, 1–2 M, 1–3 DM and prefemoral process with 0–5 spines. One tarsal spur on legs 1–19(20).
Body length up to 12.9 cm. Two colour morphs; morph 1 with antenna and legs 1–20 yellowish, morph 2 with antenna and legs 1–20 reddish. All tergites greenish brown. Cephalic plate with median sulcus. Paramedian sulci or sutures absent on posterior part.
Antenna usually with 18 articles (atypically with 17 or 19), basal 6 subcylindrical and glabrous dorsally (Fig.
Anterior margin of T1 underlying cephalic plate (Fig.
Coxopleural process moderately long, usually with three apical spines (Fig.
All legs without setae and tibial spur. One tarsal spur on legs 1–19 (on leg 20 in one spm.). Ultimate legs: moderately thick and long (Figs
Genital segments well developed, reaching longer than distance between posterior margin of sternite of ultimate leg-bearing segment and distal part of coxopleural process. Sternite of genital segment 1 round and convex posteriorly, with median suture (Fig.
The validity of Scolopendra japonica at the species level was defended by
The sympatric distribution of this species and former subspecies of S. subspinipes as well as other Asian temperate Scolopendra complicates morphological delimitation of species boundaries except using the three phenotypic characters discussed above. In this paper, we compared taxonomic characters based on collections in the
Morphological comparison of Scolopendra japonica populations in the present study and the related species S. cingulata.
Characters | S. japonica | S. cingulata | ||
---|---|---|---|---|
Japan (Syntypes) | Japan-China | Laos | ||
Number of antennal articles | 17–19 | 17–19 | 12–18 | 17–22 |
Number of glabrous articles | 6 | 6 | 6 | 6 |
Teeth on tooth-plate | 5+5 | 4+5, 5+5 | 6+6, 5+5 | 4+4,5+5 |
First tergite with complete paramedian sutures | 3–4 | 3–4 | 4 | 2–3 |
First tergite with margination | 10–13 | 11–15 | 12 | 7–12 |
Tergite surface | smooth | smooth | short median furrow on posterior part of TT7–19 | smooth |
Median furrow on tergite of ULBS | absent | absent | absent | absent |
Paramedian sutures on sternites | complete | complete | complete | complete |
Sternite of ultimate leg-bearing segment | sides converging posteriorly | sides converging posteriorly | sides converging posteriorly | sides converging posteriorly |
Spines on coxopleural process |
AP: 0–3 SAP: 0–1 |
AP: 0–3 SAP: 0–1 |
AP: 3 | AP+SAP: 1–5 |
Spine formula on prefemora of ultimate legs |
VL: 2–3 M: 1–2 DM: 1–3 SP: 0–3 |
VL: 2–3 M: 1–2 DM: 1–2 SP: 1–5 |
VL: 2–3 M: 1 DM: 2 SP: 3–4 |
VL: 1–2 M+DM: 4–8 SP: 1–11 |
Legs with one tarsal spur | 1–19 (20) | 1–19 (1–20) | 1–19 | 19 |
In the current phylogenetic framework of Scolopendra, S. japonica is resolved in the same clade as S. cingulata Latreille, 1829. The two species are morphologically similar despite their markedly disjunct distributions, i.e., S. cingulata in the Mediterranean versus S. japonica in East Asia (Table
Probably distributed throughout the temperate zone of East Asia including mainland and insular territory (Fig.
Scolopendra
pinguis
Pocock, 1891b: 411,
1,000–2,000 ft., Carin Mountain, Cheba District, Burma [Kayah-Karen Mountains, Myanmar].
Type material. This species was described based on one specimen and the holotype was probably destroyed. It was collected during a field expedition to Burma (Myanmar) by Leonardo Fea, the assistant zoologist at the Museo Civico di Storia Naturale di Genova, Genova, Italy. In 1891, Pocock published on the myriapods of Burma based on Fea and Oates’s collections. Subsequently most of Oates’s collection was deposited in the
Thailand —
Laos —
17 antennal articles, 3–4 basal articles glabrous dorsally. Each tooth-plate with 6 teeth. Tergites 3–20 with paramedian sutures. Complete tergite margination from TT16 (18)–21. Tergite of ultimate leg-bearing segment without depression or suture. Paramedian sutures on anterior 10–30% of sternites. Coxopleural process with 3–7 apical + subapical, 1–2 lateral and 0–1 dorsal spines. Ultimate leg prefemora with 6–12 VL, 1–12 VM, 2–3 M, 3–4 DM and prefemoral process with 3–4 spines. One tarsal spur on legs 1–19(20).
Body length up to 85 mm. Darkish blue colouration on entire body. Cephalic plate dichromatic in some populations. Tergites dark blue or nearly black. Cephalic plate with small punctae on anterior part, median sulcus present. Posterior part of cephalic plate without paramedian sulci.
Antenna usually with 17 articles, basal 3–4 subcylindrical and glabrous dorsally (Fig.
Anterior margin of T1 underlying cephalic plate (Figs
Scolopendra pinguis (
Coxopleural process moderately long or short with 3 apical, 0–3 subapical, 0–1 dorsal and 1 lateral spine(s). Pore-free area extending 30–45% length from distal part of coxopleural process to margin of sternite of ultimate leg-bearing segment (Figs
All legs with small setae on tarsus 2. Tibial spur absent on all legs. One tarsal spur on legs 1–19(20). Ultimate legs: thick and moderately long (Fig.
Genital segments well developed, reaching longer than distance between posterior margin of sternite of ultimate leg-bearing segment and distal part of coxopleural process. Sternite of genital segment 1 round and convex posteriorly, with median suture (Fig.
Colouration. According to
Colour morph 1A: Monochromatic, all segments including cephalic plate dark blue. Antenna dark blue on basal part, light blue on distal part. Pleuron with pale blue integument, all pleurites black. All legs blue, legs 19–21 dark blue. This morph has been found only in Thailand.
Colour morph 1B: Monochromatic, all segments including cephalic plate dark blue. Antenna dark blue on basal part, light blue on distal part. Pleuron with pale blue integument, all pleurites black. Most legs yellowish on prefemur, other podomeres dark blue with yellowish band bordering articulations; last three legs dark blue or black. This morph has been found only in Thailand.
Colour morph 2A: Dichromatic, cephalic plate dark blue on anterior part, yellowish on posterior part and T1. Antenna dark blue on basal part, light blue on distal part. Pleuron with pale blue integument, all pleurites black. All legs dark blue or black. This morph has been found both in Thailand and North-Central Laos.
Colour morph 2B: Dichromatic, cephalic plate dark blue on anterior part, yellowish on posterior part and T1. Antenna dark blue on basal part, light blue on distal part. Pleuron with pale blue integument, all pleurites black. All legs yellowish on prefemur, other podomeres light blue with yellowish band bordering articulations. This morph has been found both in Thailand and North-Central Laos.
Discussion. Scolopendra pinguis has not been revised since Pocock (1891) described the holotype from Burma. Two additional records from Batavia-Buitenzorg (Bogor, Java) expanded its geographical distribution across Southeast Asia (
Morphological comparison of Scolopendra pinguis, Scolopendra gracillima Attems, 1898 and S. calcarata. ? Character not present or insufficient data.
Characters | S. pinguis | S. gracillima | S. calcarata | |||||
---|---|---|---|---|---|---|---|---|
Pocock (1891, 1893) |
|
This study | Attems (1930) |
|
|
|
This study | |
Number of antennal articles | 17 | 17 | 17 | 17 | 17 | 17 | 17 | 17 |
Number of glabrous articles | 3 | 3 | 4 | 5 | 6 | 4 | 5–6 | 4 |
Teeth on tooth-plate | ? | ? | 6+6 | 5+5 | 4+4, 5+5 | 10–12 (in total) | 5+5, 6+6 | 5+5 |
First tergite with complete paramedian sutures | T3 | T3 | T3 | ? | T3 | ? | T3 | T3 |
First tergite with margination | 20–21 | 20–21 | T16–18 | 20–21 | only 21 | T12–14 | TT15–21 | only T21 |
Tergite surface | ? | ? | smooth | punctate on TT3(4)–19(20) | ? | smooth | ? | smooth |
Median furrow on tergite of ULBS | absent | absent | absent | absent | absent | present | absent | absent |
Extent (percentage) of paramedian sutures on sternites | nearly complete on anterior part of body | nearly complete on anterior part of body | incomplete (20–30%) | incomplete | complete only on anterior part of body | ? | incomplete | incomplete (20–50%) |
Sternite of ultimate leg-bearing segment | wide, sides converging | wide, sides converging | sides converging posteriorly | narrow, posterior margin rounded | narrow, posterior margin rounded | ? | trapeziform with shallow depression | sides converging |
Spines on coxopleural process |
AP+SAP: 5 L: 1 |
AP: 3 SAP: 0–3 LS: 1 DS: 1 |
AP: 3 SAP: 0–3 LS: 1–2 DS: 0–1 |
AP+SAP: 3–5 LS: 1 |
AP+SAP: 5–6 LS: 1 |
AP+SAP: 5 |
AP+SAP: 5 LS: 1 DS: 1 |
AP: 3–4 SAP: 0–3 LS: 1 DS: 1 |
Spine formula on prefemora of ult. legs |
V: 14–22 D: 2–8 |
V: 14–22 D: 2–8 (5–8) |
VL: 6–12 VM: 1–12 M: 2–6 DM: 3–4 |
VL: 6–8 VM: 3–4 M: 3–4 D: 1–3 SP:4–5 |
VL: 7 VM: 5–6 D: 3–4 SP: 4 |
V: 9 M: 5 D: 7 SP: 5 |
VL: 9–12 VM: 11–12 M: 2–3 D: 2–3 |
VL: 4–7 VM: 0–3 M: 1–2 DM+SP: 3–4 |
Legs with one tarsal spur | 1–20 (?) | 1–19 | 1–19 (20) | 1–20 | 1–20 | 1–21 | 1–21 | 1–21 |
A native species in Southeast Asia, distributed along the montane ranges between the Thailand-Burma borders (Fig.
Scolopendra
subspinipes
cingulatoides
Attems, 1938: 335, fig. 307,
Scolopendra
dawydoffi
Kronmüller, 2012: 22, table 1, fig. 4E [new replacement name].
Two localities were reported in the original description, Hagiang, Haut Tonkin [Hà Giang Province, northern Vietnam], and Thakek, Laos [Thakhek, Khammouane Province, Laos].
Syntypes
Scolopendra dawydoffi (
Thailand —
Malaysia —
17–18 antennal articles, 6 basal articles glabrous dorsally. Each tooth-plate with 5–10 teeth. Tergites 2(3)-20 with paramedian sutures. Complete tergite margination from TT11–21. Tergite of ultimate leg-bearing segment without depression or suture. Paramedian sutures on anterior 15–60% of sternites. Coxopleural process with 2–3 apical+subapical spines. Ultimate leg prefemora with 1–2 VL, 0–2 M, 0–2 DM, prefemoral process with 1–5 spines. One tarsal spur on legs 1–19.
Body length up to 16.2 cm (14.7 and 15.1 cm in syntypes). Reddish colouration on entire body. Cephalic plate and tergites dichromatic. Cephalic plate and tergites reddish orange; posterior border of tergites with dark band. Cephalic plate with small punctae; median sulcus present. Posterior part of cephalic plate without paramedian sulci.
Antenna usually with 18 articles (sometimes 17 on one side in some specimens), basal 6 subcylindrical and glabrous dorsally (Fig.
Anterior margin of T1 underlying cephalic plate (Fig.
Scolopendra dawydoffi (
Coxopleural process moderately long or short with two apical spines and one subapical spine (atypically only two apical spines; Fig.
All legs without setae and tibial spur. One tarsal spur on legs 1–19. Ultimate legs: thick and moderately long (Figs
Genital segments well developed, reaching longer than distance between posterior margin of sternite of ultimate leg-bearing segment and distal part of coxopleural process. Sternite of genital segment 1 round and convex posteriorly, with median suture (Fig.
This species is distinguished from S. subspinipes by its short, robust ultimate legs and three apical/subapical spines on the coxopleural process. The characteristic of incomplete paramedian sutures on the sternites further distinguishes it from S. subspinipes and S. japonica (which have complete paramedian sutures on the sternites). However, S. dawydoffi is similar to S. multidens in the absence of gonopods in the male. The distribution of S. dawydoffi is restricted to mainland Southeast Asia whereas S. multidens occurs in temperate regions of Asia, including both inland and insular parts. A specimen identified as S. multidens from Vietnam is genetically differentiated from Thai populations (see discussion of S. multidens above for molecular arguments in favour of the two taxa being separate species). Moreover, to test the hypothesis that characteristics of S. dawydoffi might indicate affinities to the cingulata group (with reference to the Mediterranean species S. cingulata;
Morphological comparison of Scolopendra dawydoffi and Scolopendra multidens; data from present study and previous taxonomic studies, i.e.,
Characters | S. dawydoffi | S. multidens | ||
---|---|---|---|---|
Laos (Syntypes) | Thailand | China (Holotype) | Hong Kong and Taiwan | |
Number of antennal articles | 18 | 17–18 | 7/4, damaged | 17–19 |
Number of glabrous articles | 6 | 6 | 6 | 6 |
Teeth on tooth-plate | 6+6 | 5+5, 7+6, 10+9 | 5+7 | 7+7 |
First tergite with complete paramedian sutures | 3 | 2–3 | 2 | 2–4 |
First tergite with margination | 11 | 12–14 | 9 | 13 |
Tergite surface | smooth | smooth | short median furrow on posterior part | short transverse groove on anterior-lateral part (TT2–20) |
Median furrow on tergite of ULBS | absent | absent | absent | absent |
Extent (percentage) of paramedian sutures on sternites | incomplete (15–35%) | incomplete (20–60%) | incomplete (40–60%) | incomplete (20–100%) |
Sternite of ultimate leg-bearing segment | sides converging posteriorly | sides converging posteriorly | sides converging posteriorly | sides converging posteriorly |
Spines on coxopleural process |
AP: 1 SAP: 2 |
AP: 1–2 SAP: 1–2 |
AP: 1 SAP: 2 |
AP: 1–2 SAP: 1–2 |
Spine formula on prefemora of ultimate legs |
VL: 2 M: 2 DM: 2 SP: 2–3 |
VL: 0–2 M: 0–1 DM: 0–2 SP: 1–5 |
VL: 3 M: 2 DM: 2 SP: 2–3 |
VL: 2 M: 2 DM: 2 SP: 1–3 |
Legs with one tarsal spur | 1–19 | 1–19 | 1–19 (20?) | 1–19 (20L) |
Podomeres of ultimate legs with shallow groove on posterior part | prefemur and femur | prefemur and femur | prefemur and femur | prefemur and femur |
Gonopods on genital segment | ? (female) | absent | ? | absent |
This species was formerly reported from only two occurrences in Southeast Asia, one in each of Laos and Vietnam (
Holotype
From “cataract”, meaning waterfall, for the type locality at Tad E-tu Waterfall.
18–19 antennal articles, 6 basal articles glabrous dorsally. Cephalic plate punctate. 5–6 teeth on tooth-plate. Tergites 7(14)-20 with paramedian sutures, all incomplete, present only on anterior and posterior parts. Tergite of ultimate leg-bearing segment without depression or suture. Paramedian sutures confined to anterior 15–20% of sternites. Coxopleural process with 1–3 apical+subapical spines, 0–1 dorsal spine, without lateral spine. Ultimate leg prefemora with 1–2 VL, 1–2 M, 0–2 DM and prefemoral process with 1–3 spines. Tarsal spur on legs 1–19(20).
(variation of paratypes is given in parentheses). Body length 12.8 cm (up to ca. 20 cm long in paratype
Antenna usually with 19 articles (atypically with 18 articles on one side), basal 6 glabrous dorsally (Fig.
Anterior margin of T1 underlying cephalic plate (Fig.
Scolopendra cataracta sp. n.: A Tergites 9–11 B Tergite of ultimate leg-bearing segment C–D Sternite of ultimate leg-bearing segment and coxopleura E–H Variation in ventral spines on ultimate leg prefemora F Lateral view of coxopleuron G Dorsal view of ultimate legs. Holotype
Scolopendra cataracta sp. n.: A Lateral view of coxopleuron B Tergite of ultimate leg-bearing segment C Sternite of ultimate leg-bearing segment, coxopleura and ultimate leg prefemora. D Asymmetry of spines on coxopleural process E Dorsal view of ultimate leg prefemora. Holotype
Coxopleural process (Fig.
All legs without setae and tibial spur. One tarsal spur on legs 1–20 (1–19 in one paratype,
Genital segments well developed (Figs
The paratype collected in Thailand in 2001 (
This species exhibits an atypical characteristic for scolopendromorphs, namely incomplete paramedian sutures on the tergites. Very few members in only three genera share this character, these being within Scolopocryptops Newport, 1844, Scolopendra and Rhysida Wood, 1862. Within Scolopendra, only two described species, S. hainanum from Hainan Island, China, and “S. subspinipes piceoflava” (treated above as a synonym of S. subspinipes following
S. cataracta differs from S. hainanum by the short paramedian sutures on the sternites versus being nearly complete in S. hainanum, the number of spines on the coxopleural process (three vs one or two), and the number of VL spines on the ultimate leg prefemora. The two species can also be distinguished by their colouration patterns and their distributions, though the latter are closely associated. For these reasons, we regard S. cataracta as distinct from S. hainanum, and its sampled populations group as monophyletic for each partial gene analysis (see Table
Morphological comparison of Scolopendra cataracta sp. n. and some related species in the adjacent sub-regions. ? Character not present, R right side, * data from
Characters | S. cataracta sp. n. | S. subspinipes piceoflava (Syntypes) | S. hainanum* | |||
---|---|---|---|---|---|---|
Laos (Holotype) | Thailand (Paratype) | Vietnam (Paratypes) | Laos (Paratype) | |||
Number of antennal articles | 19 | 19 | 19 | 18(19) | 17–18 | 17–19 |
Number of glabrous articles | 6 | 6 | 6 | 6 | 6 | 6 |
Teeth on tooth-plate | 6+6 | 6+6 | 5+5, 6+6 | 6+6 | 5+5, 6+6 | 6+6,7+7 |
First tergite with complete paramedian sutures | incomplete (short PS on anterior and posterior part) | incomplete (short PS on posterior parts from TT5–20) | incomplete (short PS on anterior and posterior parts from TT3–20) | incomplete (short PS on anterior and posterior part) | complete, starting from T3 | incomplete, starting from T3–4 |
First tergite with margination | 7 | 14 | 7 and 10 | 12 | 5–7 | 5 |
Tergite surface | smooth | smooth | smooth | smooth | punctate | smooth |
Extent (percentage) of paramedian sutures on sternites | 15–20% | 20–45% | 10–25% | 15% | complete | nearly complete |
Sternite of ultimate leg-bearing segment | sides converging posteriorly | sides converging posteriorly | sides converging posteriorly, with pit like-furrow | sides converging posteriorly | sides converging posteriorly, with pit like-furrow | sides converging posteriorly, with pit like-furrow |
Spines on coxopleural process |
AP: 3(2) DP: 0–1 |
AP: 1–2 |
AP: 1 SAP: 2 |
AP: 3 DP: 1 |
AP: 1 |
AP: 1–2 |
Spine formula on prefemora of ultimate legs |
VL: 2 M: 1 DM: 0–1 SP: 2 |
VL: 1 M: 0–1 DM: 1–2 SP: 1 |
VL: 2 M: 1–2 DM: 1–2 SP: 2–3 |
VL: 1 M: 0 DM: 0 SP: 2 |
V: 0–2 SP: 2–5 |
VL: 1 VM: 1 DM: 2 SP: 2 |
Legs with one tarsal spur | 1–20 | 1–20 | 1–19, 20(R) | 1–19 | 1–20 | 1–19 |
Gonopods on genital segment | female specimen | ? | female specimens | ? | present | absent |
All localities are in mainland territory. The currently known distribution (Fig.
The authors sincerely thank all members of the Animal Systematics Research Unit and Department of Biology, Faculty of Science, Chulalongkorn University, for field collecting and technical support. We give special thanks to T. Krutchuen for her excellent drawings and to T. Asami, K. Inkhavilay and B. Ng for collecting in Japan, Laos and Malaysia. We also extend thanks to N. Akkari, H. Sattmann, V. Stagl and E. Schiller at