AnimaliaScolopendromorphaScolopendridaeSiriwutWarutEdgecombeGregory D.SutcharitChirasakTongkerdPiyorosPanhaSomsakA taxonomic review of the centipede genus Scolopendra Linnaeus, 1758 (Scolopendromorpha, Scolopendridae) in mainland Southeast Asia, with description of a new species from LaosZookeys17520162016590112410.3897/zookeys.590.7950 Scolopendra dehaani Brandt, 1840Figs 7A, 18, 19, 20, 21, 22, 23, 24and 25A Scolopendra dehaani Brandt, 1840: 152, 1841: 59. Newport 1844: 96, 1845: 394. Kohlrausch 1881: 47. Meinert 1886: 203. Flower 1901: 21. Siriwut et al. 2015a: 7, table 1, figs 1–2. Kronmüller 2012: 24, figs 3C, 4D, table 1. Scolopendra subspinipes dehaani — Pocock 1891: 409, 1894: 312. Silvestri 1895: 714. Kraepelin 1903: 260. Attems 1907: 80, 1914a: 106, 1930b: 31, 1938: 334, 1953: 138. Takakuwa 1942c: 41. Wang 1955b: 16, 1956: 158, 1965b: 449. Schileyko 1998: 268, 2007: 75. Chao and Chang 2003: 2. Lewis 2010b: 113. Tran et al. 2013: 230. Scolopendra subspinipesJangi and Dass 1984: 30, figs 2–6. Scolopendra childreni Newport, 1844: 96. Kraepelin 1903: 260. Scolopendra concolor Newport, 1845: 394. Daday 1889: 150, 1891: 188. Kraepelin 1903: 260. Scolopendra inermis Newport, 1845: 393. Koch 1863: 64, pl. 29, fig. 55. Scolopendra inermipes Koch, 1847: 153. Kraepelin 1903: 260. Scolopendra silhetensis Newport, 1845: 393. Kraepelin 1903: 260. Scolopendra lucasii Gervais, 1847: 270. Kraepelin 1903: 260. Scolopendra horrida Koch, 1847: 154. Kraepelin 1903: 260. Scolopendra limicolor Wood, 1861: 12. Kraepelin 1903: 260. Scolopendra bispinipes Wood, 1862: 28. Kraepelin 1903: 260. Scolopendra fissispina Koch, 1865: 891. Kraepelin 1903: 260. Scolopendra nudipes Tömösváry, 1885: 67. Kraepelin 1903: 260. Scolopendra foveolata Verhoeff, 1937: 220. Würmli 1972: 91. Scolopendra arborea Lewis, 1982: 389. Lewis 2010: 97. Syn. nov.Type locality.

Java, Indonesia.

Material.

ThailandCUMZ 00325, one spm., Pha Mon Cave, Pang Ma Pha, Mae Hong Son (19°30'01.6"N, 98°16'43.5"E). CUMZ 00324, one spm., Wat Ban Mai, Mae Hong Son (19°17'55.3"N, 97°59'13.5"E). CUMZ 00323, one spm., Wat Tham Chiangdao, Chiang Mai (19°23'36.8"N, 98°55'42.6"E). CUMZ 00377, one spm., Wat Tham Pak Piang, Chiang Dao, Chiang Mai (19°24'10.498"N, 98°55'52.691"E). CUMZ 00378, one spm., Tham Chiang Dao, Chiang Dao, Chiang Mai (19°23'35.758"N, 98°55'44.412"E). CUMZ 00379, five spms., Huai Hong Khrai, Doi Saket, Chiang Mai (18°51'26.107"N, 99°13'21.827"E). CUMZ 00346, one spm., Hui Hong Khrai, Chiang Mai (18°50'58.6"N, 99°13'18.9"E). CUMZ 00380, one spm., Suan Hin Maharat Stone Park, Long, Phrae (18°9'15.072"N, 99°59'10.343"E). CUMZ 00381, one spm., Lai Nan, Wieng Sa, Nan (18°36'39.969"N, 100°53'17.242"E). CUMZ 00382, two spms., Si Nan National Park, Na Noi, Nan (18°21'56.97"N, 100°49'56.393"E). CUMZ 00288, one spm., Phusang Waterfall, Phusang, Phayao (19°40'05.0"N, 100°23'25.1"E). CUMZ 00243, Hub Pa-Tat, Larnsak, Uthai Thani. CUMZ 00276, one spm., Wat Tham Erawan, Ban Rai, Uthai Thani (15°02'01.5"N, 99°27'16.6"E). CUMZ 00329, one spm., Wat Phothikhun, Maesot, Tak (16°44'39.2"N, 98°36'17.2"E). CUMZ 00374, two spms., Wat Tham Namphu Khao Rong Kwang, Lansak, Uthai Thani (15°25'59.212"N, 99°35'7.924"E). CUMZ 00375, two spms., Wat Tham Khao Chakkachan, Chum Ta Bong, Nakhon Sawan (15°35'48.533"N, 99°32'38.758"E). CUMZ 00376, one spm., Wat Khao Huai Lung, Banphot Phisai, Nakhon Sawan (15°55'41.844"N, 99°52'20.407"E). CUMZ 00257, one spm., Wat Huai Lung, Banphot Phisai, Nakhon Sawan (15°55'29.6"N, 99°52'28.0"E). CUMZ 00256, one spm., Bang Ban, Ayutthaya (14°21'51.4"N, 100°29'22.3"E). CUMZ 00282, one spm., Wang Kanlueang Waterfall, Chai Badan, Lopburi (15°06'49.4"N, 101°06'38.8"E). CUMZ 00289, one spm., Wat Khao Somphot, Chai Badan, Lopburi (15°09'42.2"N, 101°16'49.5"E). CUMZ 00286, one spm., Chaloem Phra Kiat, Saraburi (14°40'11.9"N, 100°53'09.4"E). CUMZ 00371, one spm., Chet Sao Noi Waterfall, Muak Lek District, Saraburi (14°43'35.463"N, 101°11'22.63"E). CUMZ 00372, one spm., Phu Khae Botanical Garden, Chaloem Phra Kiat, Saraburi (14°40'10.455"N, 100°53'13.401"E). CUMZ 00373, one spm., Wat Tham Welu Wan, Suphanburi (14°57'6.824"N, 99°38'57.176"E). CUMZ 00364, one spm., Wat Mai Luak, Sai Yok, Kanchanaburi (14°26'16.384"N, 98°52'19.822"E). CUMZ 00365, two spms., Wat Tha Thung Na, Sai Yok, Kanchanaburi (14°29'40.864"N, 98°50'21.332"E). CUMZ 00268, one spm., Wat Namtok, Saiyok, Kanchanaburi (14°13'47.5"N, 99°03'59.8"E). CUMZ 00270, 00361, two spms., Wat Tham Lijia, Sangkhaburi, Kanchanaburi (15°04'12.6"N, 98°33'59.0"E). CUMZ 00362, two spms., Wat Khao I-San, Ratchaburi (13°22'57.897"N, 99°46'21.374"E). CUMZ 00363, one spm., Central Botanical Garden, Ratchaburi (13°36'24.229"N, 99°39'37.584"E). CUMZ 00253, one spm., Tham Khao Bin, Ratchaburi (13°35'35.6"N, 99°40'02.3"E). CUMZ 00241, one spm., Wat Huailat, Phu Ruea, Loei, Thailand (17°27'00.4"N, 101°24'45.4"E). CUMZ 00383, one spm., Wat Pa Huai Lat, Phu Ruea, Loei (17°26'20.14"N, 101°25'17.223"E). CUMZ 00277, two spms. and CUMZ 00384, five spms., Tham Pha Pu, Loei (17°34'41.5"N, 101°42'39.1"E). CUMZ 00292, one spm., Wat Tham Pak Khaew Chiang Khan, Loei (17°52'34.0"N, 101°40'20.8"E). CUMZ 00247, one spm., Ban Dong Savanh, Phang Khon, Sakon Nakhon (17°28'17.9"N, 103°30'12.6"E). CUMZ 00266, one spm. and CUMZ 00392, one spm., Kham Hom Waterfall, Sakon Nakhorn (17°07'19.9"N, 104°01'07.4"E). CUMZ 00393, one spm., Wat Nikhom Kaset, Mukdahan (16°48'26.312"N, 104°42'41.205"E). CUMZ 00394, one spm., Pak Maenam Songkhram, Nakhon Phanom (17°38'23.43"N, 104°27'13.517"E). CUMZ 00395, one spm., Cha Naen Waterfall, Bueng Kan (18°12'54.298"N, 103°53'18.422"E). CUMZ 00385, one spm., Tham Pu Lup, Chum Phae, Khon Kaen (16°47'48.98"N, 102°37'31.026"E). CUMZ 00386, two spms., Wat Tham Phra, Tha Khantho, Kalasin (16°51'46.069"N, 103°15'57.202"E) CUMZ 00249, two spms., Ban Dan Chang, Tha Kantho, Kalasin (14°49'29.3"N, 99°41'36.5"E). CUMZ 00271, one spm., Wat Tham Phupha, Thakantho, Kalasin (16°48'05.7"N, 103°12'37.5"E). CUMZ 00275, two spms., Tha Tum, Mueang, Maha Sarakham (16°10'32.2"N, 103°26'59.6"E). CUMZ 00387, one spm., Wat Pa Khok Hinlat, Mueang, Maha Sarakham (16°10'4.82"N, 103°28'57.255"E). CUMZ 00287, one spm., Ban Phon Thong, Kaset Wisai, Roi-Et (15°39'59.6"N, 103°33'10.9"E). CUMZ 00369, one spm., Wat Pa Ka Thon, Wang Nam Khiao, Nakhon Ratchasima (14°24'38.479"N, 101°41'3.581"E). CUMZ 00370, two spms., Wat Tham Phrommachan Thamma Ram, Pak Chong, Nakhon Ratchasima (14°34'29.046"N, 101°16'39.524"E). CUMZ 00388, five spms., Khao Kradong Forest Park, Buriram (14°56'26.209"N, 103°5'21.1"E). CUMZ 00389, one spm., Phanom Sawai Forest Park, Surin (14°45'39.325"N, 103°22'2.067"E). CUMZ 00390, one spm., Mueang, Surin (14°52'9.914"N, 103°30'21.877"E). CUMZ 00284, one spm., and CUMZ 00391, six spms., Pa Son Nongkhu, Sangkhla, Surin (14°40'55.7"N, 103°45'51.9"E). CUMZ 00248, two spms., Kaeng Lamduan, Ubon Ratchathani (14°26'15.0"N, 105°06'06.7"E). CUMZ 00279, one spm., Nang Rong Waterfall, Nakhon Nayok (14°19'52.5"N, 101°19'09.1"E). CUMZ 00252, one spm., Si Chang Island, Chonburi (13°09'08.1"N, 100°48'29.3"E). CUMZ 00258, two spms., Wat Khao Maidaeng, Sriracha, Chonburi (12°56'56.0"N, 101°02'11.9"E). CUMZ 00320, one spm., Lan Island, Chonburi (12°55'05.8"N, 100°46'43.8"E). CUMZ 00285, one spm., Juang Island, Sattahip, Chonburi (12°31'46.4"N, 100°57'18.4"E). CUMZ 00269, two spms., Samet Island, Ban Phe, Rayong (12°34'04.3"N, 101°27'23.3"E). CUMZ 00291, one spm., Wat Khao Sarp, Rayong (12°36'46.8"N, 101°23'18.8"E). CUMZ 00254, one spm., Wat Khao Chakan, Sa Kaeo (13°39'38.0"N, 102°05'02.7"E). CUMZ 00366, one spm., Wat Tham Khao Phrachan, Khao Chakan, Sa Kaeo (13°34'37.66"N, 102°5'39.42"E). CUMZ 00367, one spm., Tham Phet Pho Thong, Khlong Hat, Sa Kaeo (13°39'15.671"N, 102°29'28.066"E). CUMZ 00368, two spms., Wat Khao Phrom Suwan, Watthana Nakhon, Sa Kaeo (13°54'10.886"N, 102°30'49.344"E). CUMZ 00322, one spm., Tha Sen Waterfall, Trad (12°07'59.1"N, 102°42'22.6"E). CUMZ 00360, seven spms., Wat Khao Ma Rong, Bang Saphan, Prachuab Khiri Khan (11°12'23.041"N, 99°30'1.442"E). CUMZ 00326, one spm., Kreab Cave, Langsuan, Chumphon (9°49'01.8"N, 99°02'15.6"E). CUMZ 00359, one spm., Ban Tham Thong, Pathio, Chumphon (10°55'59.984"N, 99°29'27.521"E).CUMZ 00293, one spm., Tham Khao Kriab, Pathio, Chumphon (9°49'01.3"N, 99°02'17.9"E). CUMZ 00354, one spm., Tham Khao Phlu, Lamae, Chumphon (9°43'41.103"N, 99°6'10.996"E). CUMZ 00355, one spm., Wat Tham Khao Kriap, Chumphon (9°48'58.347"N, 99°2'10.364"E). CUMZ 00356, one spm., Ton Phet Waterfall, Kapong, Ranong (9°43'36.056"N, 98°37'7.853"E). CUMZ 00357, one spm., Wat Khao Sap, Ranong (9°57'50.532"N, 98°38'28.765"E) CUMZ 00358, one spm., Bok Krai Waterfall, Ranong (10°22'31.47"N, 98°51'15.943"E). CUMZ 00278, one spm., Bok Krai Waterfall, Kraburi, Ranong (10°22'34.6"N, 98°51'18.3"E). CUMZ 00262, one spm., King Rama V Stone park, Kraburi, Ranong (10°29'36.7"N, 98°54'35.7"E).CUMZ 00251, three spms., Sai Rung Waterfall, Takua Pa, Phang Nga (7°26'26.2"N, 99°48'47.5"E). CUMZ 00261, one spm., Surin Islands, Phang Nga (9°26'58.4"N, 97°52'37.8"E). CUMZ 00273, two spms., Khao Phlai Dam, Sichon, Nakhon Si Thammarat (9°05'33.4"N, 99°54'25.0"E). CUMZ 00281, two spms., Klong Phot Waterfall, Nop Phitam, Nakhon Si Thammarat (7°48'37.8"N, 99°12'20.0"E). CUMZ 00351, one spm., Wat Tha Li-Phong, Chian Yai, Nakhon Si Thammarat (8°5'20.669"N, 100°7'59.627"E). CUMZ 00352, one spm., Tham Khamin, Ban Na San, Surat Thani (8°49'9.678"N, 99°22'19.029"E). CUMZ 00265, one spm., Kao Sok Resort, Phanom, Surat Thani (8°54'18.9"N, 98°31'21.1"E). CUMZ 00353, two spms., Ban Laem Thong, Chaiya, Surat Thani (9°24'33.236"N, 99°17'37.035"E). CUMZ 00267, one spm., Wat Huai To, Mueang, Krabi (8°13'37.6"N, 98°53'01.7"E). CUMZ 00263, two spms., Lanta Island, Krabi (7°29'55.7"N, 99°05'20.4"E). CUMZ 00283, 10 spms., Rog Nai Island, Koh Lanta, Krabi (7°13'12.7"N, 99°04'12.7"E). CUMZ CUMZ 00260, one spm., Wat Tham Suea, Krabi (8°07'26.1"N, 98°55'27.2"E). CUMZ 00250, two spms., Ron Waterfall, Khlong Thom, Krabi (7°56'07.4"N, 99°12'37.5"E). CUMZ 00255, one spm., Tham Rue Si, Kantang, Trang (7°28'35.9"N, 99°29'04.0"E). CUMZ 00259, one spm., Chao Mai Beach, Sikao, Trang (7°26'27.8"N, 99°20'45.0"E). CUMZ 00350, two spms., Thung Khai Botanical Garden, Trang (7°28'6.826"N, 99°38'8.117"E). CUMZ 00346, one spm., Wat Tham Khao Chin, Satun (6°38'33.161"N, 100°5'8.924"E). CUMZ 00347, one spm., Tham Ton Din, Kuan Don, Satun (6°43'16.848"N, 100°9'53.819"E). CUMZ 00348, one spm., Boriphat Waterfall, Rata Phum, Songkhla (6°59'41.991"N, 100°8'47.947'E). CUMZ 00349, one spm., Wat Hua Khao Chaison, Phatthalung (7°26'11.723"N, 100°7'44.344"E) 00264, one spm., Tham Malai, Phatthalung (7°38'58.3"N, 100°06'23.7"E). CUMZ 00244, two spms., Wat Tham Phuthakodom, Sinakharin, Phatthalung (7°33'37.2"N, 99°53'07.3"E). CUMZ 00274, five spms., Tham Wang Thong, Khuan Kanun, Phatthalung (7°40'55.1"N, 100°00'56.8"E). CUMZ 00280, one spm., Tham Su Mano, Srinakarin, Phatthalung (7°35'12.3"N, 99°52'04.3"E). NHMUK, one spm., Trang, leg. Dr. David A. Wamell. NHMUK, two spms., Lamphun. NHMUK 1898.4.5-28.32, two spms., Bangkok, two spms., Ko Si Chang Island and one spm., Dong Phaya Fai, leg. S.S. Flower. NHMUK 1898.6.28.17-18, two spms., Kabin, Siam [Sa Kaeo, Thailand], leg. H. Way. NHMUK, one spm., Siam [Thailand], leg. W.H. Hillman. NHMW Inv. No. 774, one spm., Xieng-Zang near Mekong, Hinter-Indien [Chiang Saen, Chiang Rai, Thailand], January 1886.

LaosCUMZ 00335, one spm., Ban Bun Tai, Phongsaly (21°24'18.148"N, 101°57'27.294"E). CUMZ 00396, one spm., Mueang Phong Sali, Phongsaly (21°30'26.611"N, 102°10'27.28"E). CUMZ 00399, one spm., Luang Prabang (19°53'10.2"N, 102°08'16.2"E). CUMZ 00246, one spm., Ban Xai Na Pho, Xekong (15°25'28.3"N, 106°36'34.8"E). CUMZ 00245, one spm., Ban Ka Soam, Attapue (14°49'15.8"N, 106°49'05.0"E). CUMZ 00332, one spm., Thung Hin Tang 39 km from Vietnamese border, Attapue (14°45'57.353"N, 107°7'55.516"E). CUMZ 00397, one spm., Tad-Fan, Pakse, Champasak (15°11'25.735"N, 106°7'39.408"E). CUMZ 00398, one spm., Khon Phapaeng Waterfall, Champasak (13°56'53.2"N, 105°56'27.1"E).

CambodiaCUMZ 00330, one spm., Angkor Wat, Siem Reap (13°24'45.5"N, 103°52'14.7"E). CUMZ 00331, one spm., Wat Tham Ban Kele, Srisophon (13°36'05.5"N, 102°57'09.3"E).

MyanmarNHMUK 1889.7.15.2, 16 juveniles and one adult spms., Thayelinyo, Burma, leg. E.W. Oates. NHMUK, eight spms., Mandalay, Burma, leg. Oates. NHMUK, one juvenile spm., Cqimana, Uflu, Burma [undetermined]. NHMUK 1889.7.15.10, one spm., Moulmein, Burma [Mawlamyaing District, Mon State], leg. E.W. Oates. NHMUK 1889.7.15.3, five spms., Tondwingyi, Burma. NHMUK 1889.5.15.5, one spm., Teikjyi near Rangoon [Yangon], Burma. NHMUK, two spms., Rangoon [Yangon], Burma, leg. Oates. NHMUK, one spm., Owen Island, Mergui Archipelago, Burma, leg. Dr. Anderson. NHMUK 1889.5.15.6, one spm., Ihawawaddy River [Irrawaddy River], Burma, leg. E.W. Oates (Cpt). NHMUK 1889.5.15.7, one spm., South Tenasserim Range, Burma, leg. E.W. Oates.

MalaysiaNHMUK 1885.8.15.3.6, two spms., Kualin, Kedah, Malaysia, leg. S.S. Flower. NHMW Inv. No. 773, one spm., Penang, Siam [now Malaysia], leg. Skind, 1/2/1882. CUMZ 00336, one spm., Gumpung Baru, Gunung Genting, Perak (4°41'39.9"N, 100°52'46.0"E). CUMZ 00338, one spm., Klinic Desa, Kampung Panit Luar, Perak (4°56'17.9"N, 100°59'00.1"E). NHMUK 1887.71, fifty-two spms., Perak, Malaysia, leg. L. Way Esq. NHMUK 1889.5.26.1, one spm., Perak, Malaysia, leg. J.H. Leech. CUMZ 00337, Gua Musang, Kelantan (4°52'11.3"N, 102°00'40.6"E). NHMUK 1904.7.14.1-3, two spms., Kuala Kenting, Kelantan, Malaysia don. Imperial Institute. NHMUK(E) 200054 Chilo.1952.9.8.616, one spm., 2000 ft. in Koyan Forest, Sarawak, Oxford University Sarawak Expedition.

SingaporeNHMUK, one spm., Singapore, labelled “Scolopendra subspinipes var. concolor”, by Senniry. NHMW Inv. No. 775, one spm., Singapore, 1/1/1887, Aurora Expedition, leg. Swoboda, det. Attems C.

IndonesiaNHMUK 1893.5.13.2.30, two spms., east coast of Sumatra, Indonesia, leg. Mrs. Findlay. NHMW Inv. No. 8603, one spm., Java, Indonesia, with old label “Soumarak near Java”, leg. Plason A., det. Attems C., 1874. NHMW Inv. No. 772, one spm., Java, 1882. NHMW Inv. No. 8602, one spm., West Java, leg. Ausbeute Kainy, det. Attems C. NHMW Inv. No. 37U, one spm., Java, Novara Expedition. NHMW, one spm., Java, 1/2/1882, leg. Breitensl. NHMW Inv. No. 1648, one spm., Java, 1/5/1879, don. Moscovics. NHMW Inv. No. 8604, one spm., Java, Indonesia, don. Moscovics, det. Attems C. in 1/5/1879. NHMW Inv. No. 8600, one spm., Buitenzorg [Bogor], Java, Indonesia, leg. Müller, det. Attems C. NHMW Inv. No. 8605, six spms., Java, Indonesia. NHMW Inv. No. 8598, one spm., Java, Indonesia, leg. Adensamer, det. Attems C. NHMW Inv. No. 778, one spm., Java, Indonesia, don. Moscovics, det. Attems C. NHMW Inv. No. 779, one spm., Java, Indonesia, Novara Expedition.

ChinaNHMUK, one spm., Hong Kong by G. Browning.

Indian TerritoryNHMW Inv. No. 768, one spm., India or Aracan, leg. Stoliczka in 1873. NHMUK, one spm., Chamba, Himalaya in Schlogin.

MexicoNHMUK, one spm., Mexico, leg. California Academy of Science [probably introduced, if not mislabeled].

Undetermined localityNHMUK, 1894.8.23.7, one spm., unknown locality, leg. C. Hose Esq. NHMUK 1913.6.18.903, two spms., Côte Malaga, Koch’s collection; NHMUK, one spm., near mouth of Patam R. [possibly Karandag (Patam) River, Bangladesh].

Diagnosis.

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).

Composite description.

Body length up to more than 25 cm (collections from Java, NHMW). Varied colouration; cephalic plate and segments monochromatic. Tergites usually brownish-orange with or without dark band on posterior border of tergites. Cephalic plate with small punctate on anterior part; median sulcus present. Posterior part of cephalic plate without paramedian sulci.

Antenna usually with 18 articles (17, 19 and 21 articles in some specimens), basal 6 subcylindrical and glabrous dorsally (Fig. 21A), 5–5.5 glabrous ventrally. Antennae reach segment 3–4. Forcipular trochanteroprefemoral process bearing denticles in two groups, one apical and 2–3 inner. Tooth-plates wider than long or nearly equivalent, 4–5 teeth on each side (Figs 21B, 23C–D; rarely 3, 7, 8 or 10). Tooth-plate with straight, transverse basal suture. Coxosternite smooth, without median suture (Fig. 21C, 23A). Article 2 of second maxillary telopodite with spur.

Colouration changes and patterns during developmental stages of Scolopendra dehaani: A Juvenile stage B Sub-adult stage C Adult stage (specimen from northern Thailand).

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.

Scolopendra dehaani (CUMZ 00282): A Cephalic plate and trunk segments 1–2 B Tooth-plates C Forcipular segment D–E Spiracles 3, 5 and 8, respectively.

Anterior margin of T1 underlying cephalic plate (Fig. 21A). Complete paramedian sutures from T4; margination typically starting on TT9–12 (14 in one spm. from Burma; NHMUK 1889.5.1.7). Tergite surface (Figs 22A, 23B, 24D) smooth, without median sulci. Tergite of ultimate leg-bearing segment (Figs 22E, 23E) curved posteriorly, without median furrow or depression; ratio of width: length of tergite of ultimate leg-bearing segment 0.84:1. Paramedian sutures usually complete on sternites (Figs 22B, 24A), atypically confined to anterior 30–50% (Fig. 23F). Surface of sternite smooth (atypically with small pit on posterior part). Sternite of ultimate leg-bearing segment (Fig. 22D) with sides converging posteriorly; surface of sternite without depression. Pore-field on coxopleuron terminating well beneath margin of tergite of ultimate leg-bearing segment, anterior part of pore area widest (Figs 22C, 24E).

Scolopendra dehaani (CUMZ 00268 and 00282): A Tergites 10–11 B Sternites 10–11 C Lateral view of coxopleuron D Sternite of ultimate leg-bearing segment and coxopleura E Tergite of ultimate leg-bearing segment F–G ventral and dorsal view of ultimate leg prefemora.

Scolopendra dehaani (CUMZ 00365): A Forcipular segment B Tergites 9 and 10 C–D Variation in teeth on tooth-plates E Tergite of ultimate leg-bearing segment F Sternites 9 and 10.

Scolopendra dehaani (CUMZ 00365): A Sternites 9–11 B–C Spines on ultimate leg prefemora (dorsal and ventral views, respectively) D Tergite 9–11 E Lateral view of coxopleuron.

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. 22D).

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. 22G). Prefemoral spines (Figs 22F–G, 24B–C): 0 VL, 0–1 M, 0–1 DM and prefemoral process with 1–4 spines. Posterior margin of prefemur with shallow median groove.

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 7A, 25A). In male, sternite of genital segment 2 attached to penis. Tergites of genital segments without small setae. Gonopods with or without small setae. Penis with apical bristle.

Genital segment(s) in some preserved Scolopendra specimens: A Scolopendra dehaani (male) B Scolopendra calcarata (female) C Scolopendra pinguis (male) D Scolopendra morsitans (male).

Colouration. This species is among those that exhibited the most varied colouration patterns. Chao (2008) suggested two major types of Scolopendra colouration, referred to as monochromatic and dichromatic. The variability in colouration of Scolopendra dehaani has been reported in Southeast Asia over the past century (Flower 1901). In the present study, the colour patterns of growth stages have been photographed (Fig. 19). Recently, ontogenetic variation including specific changes in colouration has been discussed with reference to geographical distribution (Siriwut et al. 2015a). These findings indicated that the colouration change might involve a heritable component among populations. A descriptive classification of colouration in Scolopendra dehaani is as follows:

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.

Discussion.

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 Scolopendra subspinipes s.l. sensu Lewis (2010) but after morphological survey (Kronmüller 2012) and molecular delimitation (Siriwut et al. 2015b) a species rank has been conferred to this name. The morphological variability in the extent of paramedian sutures recorded in some specimens might demand further examination. Three morphotypes of Scolopendra dehaani within the examined collections can de delimited as follow:

Morphotype 1: Complete paramedian sutures on tergites and sternites. This is the typical form of Scolopendra dehaani, according to Attems (1930b) and Jangi and Dass (1984), observed throughout the geographic range of the species.

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. 24D) and confined to 10–20% length of sternites (Fig. 23F). This morphotype has been observed in spec. from Hong Kong and northern India and is common in Java.

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 Scolopendra dehaani Brandt, 1840, throughout the entire geographic range. A morphological comparison through the species’ geographical range is given in Table 7. In addition, we recorded some brooding and feeding behaviour of individuals in their natural habitat. Scolopendra dehaani exhibited double coiling when guarding offspring in the brood chamber (Fig. 20A), similar to Scolopendra morsitans (Fig. 20B). Two stages of feeding behaviour have been observed, which may be described as late foraging (Fig. 20D) and consumption stages (Fig. 20C). The centipede attached itself to the posterior part of the body of the snail-eating snake Pareas carinatus Boie, 1828 and then advanced to the anterior part of the prey’s body, stabbing the snake several times. The posterior part of the centipede hung twisted with a palm trunk by using the locomotory and ultimate legs. The period of consumption of the prey lasted approximately one to two hours after the initial recording; all somatic tissue was completely eaten by the centipede.

Morphological survey of geographical variation of Scolopendra dehaani populations from different regions.

CharactersGeographical distribution area
Java1,2Thailand, Laos and Cambodia1Burma1Malay Peninsula1India3China- Japan1Mexico1
Number of antennal articles14–1818–2117–1917–1918–191818
Number of glabrous articles6666666
Teeth on tooth-plate5+4,5+55+55+5,5+6,6+65+5,6+6,7+8,8+106+66+76+6
First tergite with complete paramedian suturesincomplete42–63–443–5 or incompleteincomplete
First tergite with margination7–101310–147–13587
Tergite surfacesmoothsmoothsmoothsmoothsmoothsmoothsmooth
Median furrow on tergite of ULBSabsentabsentabsentabsentabsentabsentabsent
Extent (percentage) of paramedian sutures on sternites10–15% (rarely with complete PS)90–100%100%30–100%80–100%80–100%20%
Sternite of ultimate leg-bearing segmentwith or without pit- like median depressionwith pit- like median depressionwith or without pit- like median depressionwith or without pit-like median depressionwithout pit-like median depressionwithout pit-like median depressionwithout 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 spur1–201–191–19 (20)1–19 (20)1–201–201–20

Note: each superscript number refers to data from recent and earlier studies as follow: 1 = This study, 2 = Brandt (1840), 3 = Jangi and Dass (1984).

One new subjective synonym is proposed for Scolopendra dehaani. The holotype and sole known specimen of Scolopendra arborea Lewis, 1982 (NHMUK 1952.9.8.616) is approximately 40 mm in length, from an elevation of 2,000 ft. at Koyan Forest, Sarawak. It is here regarded as an immature specimen of Scolopendra dehaani. Lewis (1982, 2010) noted that morphological characteristics of Scolopendra arborea are similar to some other Asian and one Pacific island species, namely Scolopendra dehaani, Scolopendra puensis Jangi & Dass, 1984, Scolopendra gracillima, Scolopendra hardwickei Newport, 1844, and Scolopendra metuenda Pocock, 1895. In its original description, Scolopendra arborea was noted to have a colouration pattern similar to juveniles of Scolopendra dehaani, i.e., the cephalic plate and last two segments black, the other tergites bright orange, the legs bright blue with blackish lateral lines. Its taxonomic characters appear very close to Scolopendra dehaani, the apparent differences being the number of spines on the coxopleural process (one in Scolopendra arborea versus two in Scolopendra dehaani) and tergite margination starting from T20 or only T21 margination (reaching more forward to anterior segments in Scolopendra dehaani). According to our study, the number of apical spines on the coxopleural process of adults of Scolopendra dehaani is quite strictly two, but on some occasions a single minute apical spine is present in small sub-adult and juvenile specimens. Morover, our re-examination of the holotype of Scolopendra arborea indicated that tergite margination starts from T15, which decreases the distinction from Scolopendra dehaani. An unusual morphological feature of Scolopendra arborea is the expansion of the peritrema on the spiracle of segment 3 to cover part of the tri-crescentic flaps. This atypical feature may be ontogenetic variation or the effect of muscle contraction or extension around the spiracle margin which might be affected by fixation. Without other significant diagnostic characters to distinguish this species from Scolopendra dehaani, which is abundant on the mainland and on some islands in Southeast Asia, we synonymise Scolopendra arborea with Scolopendra dehaani.

Distribution.

Widespread species in the Southeast Asian mainland and some islands (Fig. 18). The first occurrence was reported from Java, Indonesia. In this study, we provide all recorded localities of this species in Asian territory, together with some localities in which the species might be introduced, as follows: Southeast Asia: probably this is the native distribution area of this species according to population abundance and genetic structure. Population density is high throughout mainland territory especially in synanthropic areas but populations are more scattered in montane areas. The currently known distribution is as follows: Thailand (entirely), Laos (Vientiane, Khammouane, Phongsaly and Savannakhet), Cambodia (Siem Reap), Vietnam (fide Tran et al. 2013: Poulo Condore, Quang Binh, Nghe An, Dak Lak, Kon Tum, Ba Ria), Myanmar (Mergui Archipelago, Rangoon (Yagon) and Mandalay), Malaysia (Kelantan, Perak, Penang, Kedah, Selangor, Johore and Sarawak), Singapore, Indonesia (Kota Bogor (Java), Solok, Si-Rambe?, Singgalang Mountain (Sumatra), Anambas and Ambon islands) and Philippines. East Asia: China (Hong Kong and Hainan), Japan (Okinawa) and Taiwan (Hualien). South Asia: India (Himachal Pradesh (Chamba Himalaya), West Bengal (Calcutta), Karnakata, Andaman and Nicobar Islands), Bangladesh and Sri Lanka. Central America: Mexico (probably introduced).

Genital segments in some live Scolopendra specimens: A Scolopendra dehaani (male) B Scolopendra dawydoffi (male) C Scolopendra morsitans (male) D Scolopendra japonica (female) E Scolopendra pinguis (male) F Scolopendra pinguis (female).

Distribution map of Scolopendra dehaani and Scolopendra subspinipes in Southeast Asia and China-Japan Sea (small map): Filled and blank colours refer to localities from the present study and in the literature, respectively.

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