Sandracottus hunteri (Crotch, 1872), 2025

Hendrich, Lars & Brancucci, Michel, 2025, Revision of the Oriental and Australasian diving beetle genus Sandracottus Sharp, 1882 (Coleoptera, Dytiscidae, Dytiscinae), ZooKeys 1223, pp. 87-147 : 87-147

Sandracottus hunteri (Crotch, 1872) stat. rev.

Figs 7 , 18 , 30 , 37 , 38 , 40 , 41 , 53

Dytiscus fasciatus Fabricius, 1775: 825 (type locality India).

Hydaticus hunteri Crotch, 1872: 205 (type locality China) (replacement name for Dytiscus fasciatus Fabricius ).

Sandracottus fasciatus (Fabricius, 1775): Régimbart 1899: 333 (descr.); Zimmermann 1920: 234 (cat.); Hendrich and Balke 1995: 47 (ecol., cat.).

Sandracottus mixtus (non Blanchard, 1843): Hendrich 1995: 48 (ecol.); Park et al. 2009: 47 (check list); Ghosh and Nilsson 2012: 19 (cat.); Lee et al. 2014: 36. (faun.); Suprayitno et al. 2022: 135 (check list, faun.); Atthakor et al. 2018: 93 (faun.); Ghosh and Gupta 2012: 110 (faun.); Alarie et al. 2023: 307 (larval descrip.); Hájek and Nilsson 2024: 91 (cat.).

Sandracottus fasciatus var. crucialis Régimbart, 1899: 333 (type locality Vietnam, Myanmar); Vazirani 1977: 85 (syn.).

Sandracottus manipurensis Vazirani, 1969: 277 (type locality Manipur, Assam, India); Mukhopadhyay and Ghosh 2004: 368 (cat.); Ghosh and Nilsson 2012: 19 (cat.); Hájek and Nilsson 2024: 91 (cat.) (syn. nov.).

Sandracottus hunteri (Crotch, 1872): Sharp 1884: 447 (faun., check list); Jäch et al. 2012 (faun.); Suprayitno et al. 2017: 35 (faun., ecology).

Type material of Dytiscus fasciatus .

Holotype, 1 male: “ Dyt. Fasciatus Fabr. Pag. [page] 293 No 7 ” [handwritten label], “ Compared with modern examples in Brit. Mus. R. A. G. March 1926 ” [printed label with red letters], “ Hunter Coll. ZIHU 2199 ” [printed label, museum no.], “ Type ” [printed label with red frame], “ 139881 ” [catalogue number GLAHM 139881 ], “ Dytiscus fasciatus Fab. Sandracottus fasciatus Fab. ” [printed label with red frame] ( HMUG). Examined.

Type material of Sandracottus manipurensis .

Holotype not examined ( ZSI). Paratype, 1 male: “ India Assam, Manipur 5000 ft. Phaiphenigaum 30. VIII. 60 F. Schmid ” [handwritten label], “ Z. S. I. Lot No 61 1960 ” [printed label], “ PARATYPE ” [printed label], “ Sandracottus manipurensis n. sp. T. G. Vazirani, det. ” [handwritten and printed label], “ 5147 H 4 A ”, “ Sandracottus hunteri (Crotch) det. M. Brancucci & L. Hendrich 10 ” ( ZSI). Examined.

Additional material.

(473 specimens): • Nepal: 7 exs., “ E-Nepal, Dhankuta-Hills, Thamur Valley, 1150–2000 m, 23. - 25 V. 1983, M. Brancucci leg. ” ( NMB). • India: 2 exs., “ NE India, Assam Bhalukpong, 150 m, 27 ° 02 ' N 92 ° 36 ' E, 26. V. - 3. VI. 2005, Dembicky leg. ” ( NMB); 2 exs., “ NE India, Arunachal Prov., 8 km S Jamiri Sessa vicinity, 27 ° 07 ' N 92 ° 34 ' E, 26. V. - 4. VI. 2005, 350 m, P. Pacholatko leg. ” ( NMB); 1 ex., “ Darjeeling Distr., Alghera, 2900 m, 25. IV. 1982, Ch. J. Rai leg. ” ( NMB); 3 exs., “ Darjeeling Distr., Kalimpong env., III. - IV. 1989, Ch. J. Rai leg. ” ( NMB). • China: 1 ex., “ China Kouy-Tcheou [= Guizhou] Kouy-Tcheou, Reg. de Pin-Fa, 1909 Cavalerie ” ( CHF); 1 ex., “ China Yunnan-See, 1898 Excoiffier ” ( CHF); 1 ex., “ Chengtu 1933 ” [Chengdu], “ Szechwan China DC Graham XI- 27-1700 ft ” ( USNM); 1 ex., “ Mu Dong Szechwan China DC Graham 6-15 - 33 Alt 1100 ft ” ( USNM); 7 exs., “ Spirit Valley ”, “ Nanking Kiangsu Province ”, “ China H. F. Loomis Oct. 25, 1919 ” ( USNM); 2 exs., “ China 1942 Yun Hsien March WL Jellison ” ( USNM); 17 exs., “ Suifu Sz. China ”, “ June 1928 ”, “ DCGraham Collector ” ( USNM); 1 ex., “ Suifu Szchuen China 1923 ”, “ DCGraham Collector ” ( USNM); 1 ex., “ Suifu 1000-2000 ft Szchuen China 23´DCGraham Collector ” ( USNM); 8 exs., “ Suifu Szchuen ”, “ DCGraham Collector ” ( USNM); 1 ex., “ China, Hayan Isl. Wuzhi Shan Mt. [Hainan, Wuzhi Shan Nature Reserve], 1500 m 18 ° 53 ' N 109 ° 43 ' E 20.02. - 10. 04. 2001 local collector ”, “ coll. H. Hebauer ” ( CHH); 2 exs., “ Chine A. David ” ( MNHN); 1 ex., “ Su Tschuen Siao Lou 1897 “ ( MNHN); 1 ex., “ Guanxi, 350 m a. s. l. 10 km S Yangshuo, muddy pools, fields, pasture 3. XI. 1999 leg. J. Šťastný ” ( CJS); 5 exs., “ Pingshiang [Pingxiang, Jiangxi] Süd-China Dr. Kreyenberg, Coll. Kraatz ”, “ Zimmermann det. ” ( DEI); 3 exs., “ Kiukiang [Jiujiang] June 1887 AE Pratt ”, “ Coll. Kraatz Régimbart det. ” ( DEI); 2 exs., “ Kiukiang July 1887 AE Pratt ”, “ Coll. Kraatz Régimbart det. ” ( DEI); 2 exs., “ Kiangsi [Jiangxi] China ”, “ Coll. Kraatz Régimbart det. ”, “ Zimmermann det. ” ( DEI); 1 ex., “ China, coll. Gärtner ” ( DEI); 2 exs., “ China Prov. Hupeh [Hubei] Mts. Wu-schan ” ( DEI); 2 exs., “ Su Tschuen, Siàn-Lou 1897 ” ( MNHN); 1 ex., “ China, Kouang-si, P. Barrière 1909 ex. Coll. Oberthür ” ( MNHN); 33 exs., “ Su Tschuen Chasseurs Indigènes 1903 ” ( MNHN); 20 exs., “ Sian-Lou Chasseurs du P. Dejean 1904 ” ( MNHN); 1 ex., “ China Kouangsi Rég. de Nanning 1931 ” ( MNHN); 1 ex., “ Foochau [Fuzhou, Fujian] April 1886 Leech ” ( MNHN); 4 exs., “ Kiukiang [= Jiujiang] June 1887 A. E Pratt ” ( MNHN); 4 exs., “ Su-Tchuen, Chasseurs Indigènes 1907 ” ( MNHN); 2 exs., “ Su-Tchuen, Siàn-Lou 1897 ” ( MNHN); 1 ex., “ Bas-Yunnan A. Salvat 1904 ” ( MNHN); 2 exs., “ Chine, A. David “ ( MNHN); 2 exs., “ Kiang-Si A. David ” ( MNHN); 3 exs., “ Kouy Tchéou Abbé Largeteau ” ( MNHN); 6 exs., “ Kouy Tchéou Rég. De Pin-Fa Père Cavalerie 1908 ” ( NMB, MNHN); 1 ex., “ Chine Shanghai ” ( MNHN); 1 ex., “ China bor. Ex Musaeo Thorey ” ( MNHN); 4 exs., “ China “ ( NHMUK); 1 ex., “ Danes Island [Changzhou Island, Guangdong] 6756 ” ( NHMUK); 4 exs., “ Canton [Guangzhou, Guangdong], C. W. Howard, B. M. 1922-22 ” ( NHMUK); 5 exs., “ Da-Iaen-saen near Nong-po [village near Ningpo, Zhejang] Walker Coll. 93. - 18. ” ( NHMUK); 3 exs., “ Hong Kong, J. J. Walker ”, “ G. C. Champion Coll. B. M. 1927-409 ” ( NHMUK); 19 ex., “ China, Da-Iaen-saen [village near Ningbo, Zhejang], J. J. W. ”, “ G. C. Champion Coll. B. M. 1927-409 ” ( NHMUK); 16 exs., “ Western China Suiling W. A. Maw. 1909-38 ” ( NHMUK); 1 ex., “ Prov. Hunan, Huitong, 550 m, 9. - 16. VII. 1992, Ji Lanzhou leg. ” ( NMW); 1 ex., “ Yunnansen ”, “ From R. Oberthür ”, “ Coll´n J. D. Sherman Jr. 1926 ” ( USNM); 1 ex., “ Yünnan, Xishuangbanna ca. 10 km NW Menglun, ca. 700–800 m, 7. XI. 1999, leg. Jäch et al. (CWBS 360) ” ( NMW); 1 ex., “ SE-Guangxi, District Yulin, Liuwan Mts. SW Yulin, 350–400 m, 16. XI. 1993, Schillhammer leg. (20) ” ( NMW); 1 ex., “ Kiu Kiang [= Jiujiang] 1889 ”, “ Coll. Kraatz ” ( NMW); 2 exs., “ Nanking, Dr. Jettmar ” ( NMW); 1 ex., “ China, Dr. Stoltz BH ” ( NMW); 1 ex., “ Kiukiang [= Jiujiang] VII. 1887 A. E. Pratt ” ( NMW); 1 ex., “ Chekiang, Chusan, 17. VII. 1931, O. Piel leg. ” ( MNHN); 2 exs., “ China ” ( MNHN); 2 exs., “ Prov. Hupeh ” ( MNHN); 1 ex., “ Kiangsi, Staudinger & Bang Haas ” ( MNHN); 1 ex., “ Szechuan, Chungking, E. Reitter ” ( MNHN); 1 ex., “ Kuong Si ” ( MNHN); 2 exs., “ China Yunnan ” ( MNHN); 1 ex., “ S-China Canton S. W. Howard ”, “ BM 1922-22 ” ( CGW); 3 exs., “ Central China ”, “ Nonfried Coll. ” ( CGW); 5 exs., “ W-Guizhou Prov., Leigongshan, Xijiang, 1200–1900 m, 29. V. - 2. VI. 1997, Bolm leg. ” ( NMB); 2 exs., “ Hainan (217) 6 km W Dongxing, 50 m, 25. I. 1996, Jäch leg. ” ( NMW); 37 exs., “ Chine Tchékiang [Ningpo] Coll. David ” ( MNHN); 1 ex., “ Ou Hou “, “ Gebrüder W. Müller, Vermächt. 1909 ” ( SMTD); 2 exs., “ Kiangsi [Jiangxi] ” ( SMTD); 2 exs., “ Kiautschau ” ( SMTD); 3 exs., “ Shanghai “, “ Coll. C. Felsche Geschenk 1907 ” ( SMTD); 3 exs., “ Kiukiang [Jiujang] June 1887 A. E. Pratt ”, “ Samml. A. Zimmermann ” ( ZSM); 3 exs., “ China, “ Samml. A. Zimmermann ” ( ZSM); 4 exs., “ China, Kiautschou ”, “ Samml. A. Zimmermann ” ( ZSM); 2 exs., “ China, Pingshiang ”, “ Samml. A. Zimmermann ” ( ZSM); 2 exs., “ China, Kiaugsi ”, “ Samml. A. Zimmermann ” ( ZSM); 1 ex., “ China Su-Tchuen chasseurs indigènes 1903 ” ( CLH); 2 exs., “ China, Central Sichuan Baoguo Emet Co, 27. VII. - 3. VIII. 1994, Benes leg. ” ( CLH); 4 exs., “ Sechuan, Pingwu, 32 ° 15 ' N 104 ° 16 ' E, 3. - 9. VI. 1997, E. Kucera leg. ” ( CLH); 1 ex., “ China, Shanghai ”, “ Coll. E. Csiki ”, “ Sandracottus fasciatus v. hunteri Crotch ” ( TDMB). • Myanmar: 6 exs., “ Taunggyi, Burma 2 ' 27 Miss Northup ” ( USNM); 1 ex., “ Myanmar Catcin Cauri Birranta Fea Ag. Nov. 1886 ” ( MNHN); 1 ex., “ Myanmar Sagaing Division, Alaungdaw Katthapa NP, Khaung Din stream, ca. 450 m, 11. V. 2003, Boukal et al. leg. (119) ” ( NMW); 1 ex., “ Myanmar Sagaing Division, Alaungdaw Katthapa NP, Khaung Din stream, ca. 360 m, 7. V. 2003, Boukal et al. leg. (119) ” ( NMW); 4 exs., “ Myanmar, Sagaing Division, Chatthin Wildlife Sanctuary, 23 ° 31.481 N 95 ° 38.804 E, ca. 260 m, 9. X. 1998, Schillhammer leg. (9) ”; 4 exs., “ Myanmar Shan State, NE Mintaingbin Forest Camp [ca. 35 km N Aungban] above (150), pool, 22. VIII. 2004, Shaverdo leg. (152) ” ( NMW); 6 exs., “ Myanmar Shan State, NE Mintaingbin Forest Camp above (150), ca. 1200 m, puddles, 14. - 20. VIII. 2004, Shaverdo leg. ” ( NMW); 6 exs., “ Myanmar Shan State, NE Mintaingbin Forest Camp above (150 A), ca. 1200 m, pond, 14. - 20. VIII. 2004, Shaverdo leg. ” ( NMW); 1 ex., “ Myanmar Shan State, Shwedaung Wildlife Sanctuary, 23 ° 05.129 N 96 ° 13.527 S, 360 m, 19. VII. 2002, M. Hlaing & A. Moa leg. ” ( NMW); 9 exs., “ Myanmar SW Shan-State, 70 km W Taungguyi, 10. - 11. VI. 1997, J. Rejsek leg. ” ( CJS, NMPC); 1 ex., “ SW Shan state, Inle lake Nyaungshwe, 7. - 16. VI. 1997, J. Rejsek ” ( CJS); 1 ex., “ Myanmar Shan State Kalaw env. 1356 m N 20.63200 ” E 96.56197 ” 8. - 20. 6. 2015 Walter Grosser lgt. ” ( CLH); 10 exs., “ Myanmar, Shan-State, near Kalaw, 20 ° 36 ' 48 ´´ N 96 ° 34 ' 46 ´´ E, 1400 m, 15. XI. 2003, M. Hornburg leg. ” ( CLH). • Cambodia: 1 ex., “ Cambodscha Schmidt ” ( MNHN); 5 exs., “ Pnom Penh V. de Salvara ” ( MNHN); 1 ex., “ Cambodia sept. or., Stung Treng 18. - 22. IV. 1998, 13 ° 32 ' N 105 ° 58 ' E, J. Mlíkovský leg. ” ( NMPC). • Laos: 1 ex., “ NE Laos Huaphanne Prov. Mt. Phu Pane, 1220–1900 m Ban Saluei v. env., 18. V. - 2. VI. 2012 20 ° 12 ' N 103 ° 59 ' E, St. Jakl + Lao collector leg. Coll. Hendrich ” ( CLH); 1 ex., “ Khammouan Prov., Nakai env., 500–600 m, 22. V. - 8. VI. 2001, 17 ° 43 ' N 105 ° 09 ' E, E. Jendek & O. Šauša leg. ” ( CHH); 8 exs., “ Laos Attapeu prov., Ban Vang Tat Noi env. 900 m, 15 ° 03-04 ' N / 107 ° 24 ' E, 10. - 25. V. 2011 ”, “ NHMB Basel 2011 Expedition, M. Brancucci, M. Geiser, D. Hauck, Z. Kraus, A. Phantala & E. Vongphachan ” ( CLH, NMB); 2 exs., “ Laos Svannakhet prov., Phou Xang He NBCA, ca. 5 km SW Ban Pa Phaknau, 250–400 m 17 ° 00 ' N / 105 ° 38 ' E, 31. V. - 6. VI. 2011 ”, “ NHMB Basel 2011 Expedition, M. Brancucci, M. Geiser, D. Hauck, Z. Kraus, A. Phantala & E. Vongphachan ” ( NMB); 2 exs., “ Laos Bolikhamsay prov., Nam Kading NPA research training center near Ban Phon Kham, 18 ° 20 ' N / 104 ° 08 ' E, 250 m, 23. - 29. V. 2011 ”, “ NHMB Basel 2011 Expedition, M. Brancucci, M. Geiser, D. Hauck, Z. Kraus, A. Phantala & E. Vongphachan ” ( NMB); 2 exs., “ Xayaboury Prov., Xayaboury env., 19 ° 13 ' N, 101 ° 42 E, 300 m, 27. - 30. VI. 2010, leg. D. Hauck ” ( NMB); 1 ex., “ Laos Umg. Vientiane III. - VI. 1963 ” ( NMB); 20 exs., “ ATTAPEU prov., Annam Highlands Mts., Dong Amphan NBCA, ca. 1160 m, NONG FA [crater lake] env., 15 ° 05.9 ' N, 107 ° 25.6 ' E, 30. iv- 6. V. 2010, J. Hájek leg. ” ( NMPC); 1 ex., “ Pak Lag, 13. VIII. 1918 V. de Salvara leg. ” ( MNHN); 1 ex., “ Luang Prabang I. 1917 V. de Salvara ” ( MNHN); 3 exs., “ Ban Mong 15. XII. 1917 V. de Salvara ” ( MNHN); 2 exs., “ Phongsaly Prov., Phongsaly env., 1500 m, 21 ° 41 ' N 102 ° 06 ' E, 28. V. - 20. VI. 2003, M. Brancucci leg. ” ( NMB); 4 exs., “ Phongsaly Prov. Ban Hatsa, 550 m, 21 ° 44 ' N 102 ° 12 ' E, 9. V. + 17. VI. 2004, M. Brancucci leg. ” ( NMB); 8 exs., “ Phongsaly Prov., Ban Sano Mai, 1150 m, 21 ° 44 ' N 102 ° 12 ' E, 19. V. - 26. V. 2004, M. Brancucci leg. ” ( NMB); 3 exs., “ NE Laos, Houa Phan prov., 20 ° 13 ' N 104 ° 00 ' E, Phou Pane Mt., 1. - 16. VI. 2009, 1350–1500 m, M. Brancucci leg. ” ( NMB); 2 exs., “ Xieng Khouang prov., 19 ° 03 ' N 103 ° 24 ' E, Ban Thaviang env., muddy puddle, 500–600 m, 19. V. 2010, M. Geiser leg. ” ( NMB); 1 ex., “ N-Laos, Louangphrabang Prov., 19 ° 53 ' N 102 ° 09 ' E, Khan river, 300 m, V. Kuban leg. ” ( NMB); 1 ex., “ N-Laos, Louangphrabang Prov., 21 ° 09 ' N 101 ° 19 ' E, Namtha, 900–1200 m, 5. - 31. V. 1997, V. Kuban leg. ” ( NMB); 1 ex., “ Boloikhamxai Prov., 70 km NEE Vientiane, 150 m, 18 ° 16 ' N 103 ° 11 ' E, 27. - 30. IV. 1997, V. Kuban leg. ” ( NMB); 1 ex., “ Vientiane Prov., Lao Pako env. 55 km NE Vientiane, 1. - 4. V. 2004, J. Bezděek leg. ” ( NMPC); 1 ex., “ C-Laos, Bolikhamsai Province, Ban Nape – Kaew Nua Pass, 18. IV- 1. V. 1998, alt. 600 m, 18 ° 22.3 N 105 ° 09.1 E, M. Strba & R. Hergovits leg. ” ( CLH); 1 ex., “ N Laos, Vientiane Prov., Vang Vieng, 300 m, 18 ° 55 ' 23 ' N 102 ° 26 ' 55 ' E, 1. - 15. V. & 1. - 6. VII. 2001, J. Koubac leg. ” ( NMB); 1 ex., “ CE Laos, Boli Khan Xai prov., 18 ° 21 ' N 105 ° 08 ' E, 8 km NE Ban Nape, 600 m, 1. - 18. V. 2001, V. Kuban leg. ” ( NMB); 20 exs., “ Annam Laos ”, “ Sandracottus mixtus ab. crucialis Rég. Dr. Guignot det. 1956 ” ( TDMB). • Thailand: 2 exs., “ Siam Kra Ding 12 III 56 RE Elbel ” ( USNM); 7 exs., “ Siam 17 V 53 RE Elbel ”, “ Pek-chong Sikiu-kerat Nong Min ” ( USNM); 7 exs., “ Thailand 27 VI 54 REElbel ”, “ Sakon Nakhon, Muang Sakon Nakhon, Phu Phan ” ( USNM); 24 exs., “ Nakhon Ratchasima Province, Sakaerat Biosphere Reserve, King Cobra Cave, Thailand Coords: 14 ° 30.536 ’ N, 101 ° 55.921 ’ E, 362 m a. s. l., 27 Oct 2013, W. Atthakor leg., Sakaerat Expedition, lead by Professor Somsak Panha of Chulalongkorn University ” ( CSUT); 6 exs., “ THAILAND Kanchanaburi Prov. Sai Yok N. P. – Pools on road, 23. VII. 1996 leg. P. Mazzoldi “ ( CPM); 17 exs., “ Thailand, Mukdahan LL 2000, Phu Pha Thoep N. P. Small pools on dry stream bed, Mazzoldi P. leg. (23) ” ( CPM); 1 ex., “ Chiang Mai prov. 35 km NW of Muang Ngai, 18 ° 40 ’ N 98 ° 42 ’ E, 10. I. 2006, S. Bečvář S. & R. Fouque leg. ” ( CJS); 2 exs., “ NE Thain Nan district, Ban Pha Khap, 15. - 20. V. 1992, Pacholatko leg. ” ( NMW); 2 exs., “ NW-Thailand, Chiang Mai, Soppong-Pai, 1800 m, 1. - 8. V. 1993, Pacholatko & Dembicky leg. ” ( NMW); 4 exs., “ NW-Thailand, Chiang Mai (Zoo), 9. - 16. V. 1988, at light, Malicky leg. ” ( NMW); 1 ex., “ NW-Thailand, Chiang Mai, 10. - 17. V. 1989, Malicky leg. ” ( NMW); 2 exs., “ Pang, 300 m, 19 ° 55 ' N 99 ° 12 ' E, D. Král leg. ” ( NMB); 4 exs., “ NW-Thailand, Mae Hong Son, Ban Huai Po, 1500 m, 8. - 17. V. 1992, S. Bily leg. ” ( NMB); 16 exs., “ NW-Thailand, Mae Hong Son, Ban Huai Po, 1600–2000 m, 9. - 16. V. 1991, J. Horák leg. ” ( NMB, NMW); 1 ex., “ NW-Thailand, Mae Hong Son, Ban Huai Po, 1700 m, 24. - 30. VI. 1993, Schneider leg. ” ( NMW); 1 ex., “ NW-Thailand, Mae Hong Son, Ban Huai Po, 1600–2000 m, 9. - 16. V. 1991, Pacholatko leg. ” ( NMB); 1 ex., “ Lansang n. g., Thanon Thong Chai, 500 m, 18. - 24. IV. 1991, 16 ° 48 ' N 98 ° 57 ' E, D. Kral & V. Kuban leg. ” ( NMB); 1 ex., “ Thanon Thong Chai, Chiangdao, 19 ° 24 ' N 98 ° 55 ' E, 600 m, 10. - 16. V. 1991, D. Kral & V. Kuban leg. ” ( NMB); 1 ex., “ Lansang NP, 18. - 24. IV. 1991, 500 m, 16 ° 48 ' N 98 ° 57 ' E, D. Král leg. ” ( NMPC); 1 ex., “ Siam ”, “ Gebrüder W. Müller Vermächt. 1909 ” ( SMTD). • Malaysia: 1 ex., “ West Malaysia, Kuala Lipis, small forest pool in rubber plantation, 15. IV. 1997, Balke & Hendrich leg. ” ( CLH); 3 exs., “ Kedah Peak, 3200 feet, XII. 1915 ”, “ ex. F. M. S. Museum B. M. 1955-354 ” ( NHMUK); 2 exs., “ West Malaysia, Penang, Botanical Garden, 27. I. 1992, M. Jäch leg. ” (9) ( NMW). • Vietnam: 1 ex., “ Vietnam 19. 05. 2007 Quang Tri Da Krong NP near headquarter lux Csorba leg. ” ( CHF); 2 exs., “ Vietnam N, Quang Binh prov. 1 km N of Cha Lo, 400 m Vitenam-Laos border area 17 ° 41 ' 22 ´´ N 105 ° 45 ' 45 ´´ E, L. Dembicky leg., 11. - 24. iv. 2010 ” ( NMB); 4 exs., “ Pa-kha 4914 ”, “ Indochine Coll. Dussault ” ( NMB); 5 exs., “ N Vietnam, Sa Pa, 1530 m, 25. - 9. VI. 1991, J. Strnad leg. ” ( NMB); 1 ex., “ S Vietnam, Saigon, jardin botanique [botanical garden], octobre 1870, A. Krempf leg. ” ( MNHN); 3 exs., “ Cuc Phuong (170 m) Ninh Bin Prov. (N-Vietnam) 10. vii. 1997, S. Nomura leg. ” ( NMST); 1 ex., “ Tonkin Occ. Rég. De Hoa Binh R. P. A de Cooman 1919 ” ( MNHN); 1 ex., “ Sikkim, Kurseong, R. P. Bretaudeau 1894 ” ( MNHN); 1 ex., “ Chasseurs indigènes de ta-Tsién-Loù R. P. Dejean, 1901 ” ( MNHN); 4 exs., “ Tonkin Backan P. Lemée 1907-08 ” ( MNHN); 8 exs., “ Tonkin Backan P. Lemée, 1908 ” ( MNHN); 1 ex., “ Tonkin Occ., Env. de Hoa Binh, R. P. A. de Coomann, 1919 ” ( MNHN); 5 exs., “ S-Vietnam, 12 km N Dalat, Lang Bian, 28. - 30. IV. 1994, Pacholatko & Dembicky leg. ” ( NMW); 1 ex., “ S Vietnam, 16 km N Dalat-Ankroat, 1400 m, 12 ° 05 N 108 ° 24 E, 15. IV. 1995, Pacholatko & Dembicky leg. ” ( NMW); 1 ex., “ Tonkin, Chaba, 27. VII. 1917, Jeanvoine leg. ” ( MNHN); 8 exs., “ Hoa Binh Tonkin A. deCooman leg. ” ( MNHN); 10 exs., “ Lac Tho Tonkin, A. deCooman leg. ” ( MNHN); 1 ex., “ Hanoi 29. III. 1917 V. de Salvara ” ( MNHN); 1 ex., “ Hoa Binh I. 1917 V. deSalvara ” ( MNHN); 1 ex., “ N-Vietnam, Shonla [Son La], 9. X. 1991, Murzin leg. ” ( NMW); 1 ex., “ N-Vietnam, Ma Da, 27. XII. 1990, Murzin leg. ” ( NMW); 1 ex., “ Hoang Lien Son Prov., Sa Pa, 11. - 19. VI. 1990, M. Dvorák leg. ” ( NMPC); 2 exs., “ S Vietnam, Lao Cai Prov., Sa Pa district, 22 ° 20 ' 48.1 ´´ N, 103 ° 47 ' 45.2 ´´ E, 1690 m, at light, 24. - 25. VIII. 1998, leg. A. Kun ” ( TDMB); 1 ex., “ S Vietnam, Sa-Pa, Hoang Lien Son, 11. - 19. VI. 1990, Mir. Dvořák leg. ” ( CJS); 1 ex., “ N-Vietnam, Tam Dao, V. - VI. 1990, Pich Richard leg. ” ( NMB); 1 ex., “ N-Vietnam, Tam Dao, 20. - 28. VI. 1990, S. Brantlova leg. ” ( NMB); 1 ex., “ Tam Dao, 8. V. 1990, M. Dudycha leg. ” ( CJS). • Indonesia: 1 ex., “ Java ” ( NMB); 1 ex., “ Java Cote Sud Salatri ” ( MNHN); 1 ex., “ Java ”, “ Zimmermann det. ” ( DEI); 9 exs., “ Java ”, “ Prov. Pasuruan, Kalipari, 300–500 met., W. Doherty, 1891 ” ( MNHN); 3 exs., “ Java occidental, Mons. Tjikorai, 4000 1852 ”, “ H. Fruhstorfer ” ( MNHN); 2 exs., “ Java occ. Toegoë 1902 ” ( MNHN); 1 ex., “ Jav. Occ. G. Lamboreth, 1902 ” ( MNHN); 1 ex., “ Java Occ., Preanger [Parahyangan or Priangan] Tac Prau Ex. Musaeo van Lansberge ” ( MNHN); 1 ex., “ Java Mts Kawie J. B. Lediou 1898 ” ( MNHN); 2 exs., “ Java Ex Museo Thorey ” ( MNHN); 1 ex., “ Java Occ, G. Lambreth, 1908 “ ( MNHN); 2 exs., “ Java Schaum ” ( MNHN); 1 ex., “ Java Coll. Le Moult ” ( MNHN); 6 exs., “ E Java 50 km S Surabaya, Tretes, Kekek Bodo WF, 20. IX. 1995, Schillhammer leg. (1) ” ( NMW); 1 ex., “ Java Japara I. 1933 ” ( MNHN); 1 ex., “ Karanggandoel Banjoemas K. Benner 1922 Muzeum Buitenzorg ” ( DEI); 1 ex., “ Java ” ( NMW); 1 ex., “ E Java, Bajoekidoel b. Banjoawangi 1930 Lucht leg. ” ( MNHN); 1 ex., “ Java Coll. Boucard ” ( MNHN); 2 exs., “ NE Java Baluran NP 600 m 16. - 19. IV. 1996 R. Zajicek leg. ” ( NMB); 2 exs., “ Java Gebrüder W. Müller, Vermächt. 1909 “ ( SMTD); 1 ex., “ East Java Pare 10. III. 1949 W. J. M. Vestjens leg. ” ( ANIC); 1 ex., “ Bali, Bedugul 3 km NE Candi Kuning, 1320 m, 11. VII. 1991, Balke & Hendrich leg. (BA 8) ” ( CLH); 1 ex., “ Holländisch Indien ”, “ Zimmermann det. ” ( DEI); 1 ex., “ Sumatra ”, “ Sandracottus hunteri Zimmermann det. ” ( ZSM); 1 ex., “ Sumatra, Balighe, X. 90 - III. 91, E. Modiglioni ” ( MNHN); 1 ex., “ Sumatra, Fort de Kock II. 1925, native collector ” ( MNHN); 1 ex., “ Sumatra 1873 ” ( MNHN); 1 ex., “ Sumatra Medan ” ( MNHN); 4 exs., “ Sumatra, Dolog Merangir, 8. V. - 10. VI. 1983, E. W. Diehl leg. ” ( NMB); 1 ex., “ Sumatra, Dolog Merangir, 10. - 12. V. 1981, E. W. Diehl leg. ” ( NMB); 3 exs., “ NE Sumatra, Medan, Dr. Marx leg. ” ( ZSM); 4 exs., “ Sumatra O. K Medan 19. 9. 1921 J. B. Corporaal ”, “ Museum Paris 1902 J. B. Corporaal ” ( NMB); 3 exs., “ Sumatra Medan J. B. Corporaal ”, “ Coll. A. Zimmermann ” ( ZSM); 1 ex., “ Sumatra Palembang ” ( ZSM); 1 ex., “ Sumatra, coll. Martin ” ( ZSM); 3 exs., “ Sumatra Padang Fandjang, 800 m, 1. trim. 1896, Kannegieter “ ( ZSM); 2 exs., “ Höllandisch Indien ” [Indonesia] ( ZSM); 4 exs., “ Sumatra Palembang, Knappert leg. ” ( NMW); 1 ex., “ Sumatra Monte Battak [Bukit Batak] ex. coll. Fruhstorfer ” ( MNHN).

Remarks.

Specimens with an absent subapical transverse band were described as var. crucialis Régimbart, 1899 . Vazirani (1969) described his S. manipurensis from Assam based on specimens with more extensive black markings of the elytra. When studying the only male paratype of S. manipurensis available to us, we were not able to detect any morphological differences in the structure of the median lobes of S. manipurensis and S. hunteri . The colour pattern of the elytra of the paratype can also be found within the range of variation in S. hunteri . We therefore consider S. manipurensis synonymous with S. hunteri .

Redescription.

Body broad oval, shiny, testaceous with black markings. Ventral side completely dark brown to black, legs ferrugineus brown to testaceous.

Head testaceous with posterior part and eye margin black, shiny (Fig. 7 ). Surface sculpture consisting of very small dense punctures and of larger, much sparser ones, particularly numerous on frons. Clypeal grooves, punctures alongside eyes and transverse depression beside eyes distinctly impressed, punctures large and coalescent. Antennae testaceous; antennomeres slender, antennomere V 3.5 × as long as broad.

Pronotum testaceous with median black marking reaching from posterior to anterior margins (Figs 7 , 53 ); broad on posterior, distinctly narrower on anterior margin. Surface shagreened with dense punctation; punctures medium-sized. Anterior and lateral puncture lines with dense coalescent punctures, becoming sparse towards middle, and lacking in middle of anterior margin. Posterior puncture line distinctly visible only at sides, towards middle punctures become superficial and transformed into long wrinkles.

Elytra testaceous to ferrugineus brown with black markings consisting of three transverse bands: broad antemedian, subapical, and apical; suture with narrow black frame, and puncture lines marked by black spots (Figs 9 , 53 ). Epipleura testaceous to ferrugineus brown. Surface very slightly and superficially shagreened and covered with double punctation; denser smaller punctures and much sparser larger ones. Puncture lines with isolated groups of five or six medium-sized punctures; discal row almost complete and strongly impressed. Sutural puncture line marked only by few punctures along suture.

Ventral side dark brown. Legs, particularly fore and mid legs testaceous, hind legs ferrugineus brown to dark brown (Figs 9 , 53 ). Prosternal process short and broad, 1.7 × longer than broad, rounded apically, and finely but distinctly sculptured. Posterior border triangular, narrowly ended. Whole surface very superficially shagreened and finely punctured. Metatibia with numerous and dense small punctures on basal half. Ventrites II – VI shagreened, longitudinally wrinkled on lateral parts, densely covered with very small punctures and larger sparser ones. Posterior margins rounded, bordered with some large and coalescent punctures along margin.

Measurements: TL = 12.5–15.0 mm, TL-h = 11.4–14.0 mm, TW = 8.3–8.8 mm.

♂. Protarsomeres I – III strongly enlarged with three larger suckers and numerous smaller one. Mesotarsomeres I – III with two rows of small suckers. Median lobe of aedeagus, in ventral view, broad, nearly parallel-sided on whole length, lobes narrowly rounded at apex (Fig. 18 a ). Parameres slightly longer than median lobe, broad and pointed at apex (Fig. 18 b ).

♀. Similar to male, tarsi not enlarged. Microsculpture of ventrite VI as in male.

Differential diagnosis.

The species is clearly distinguished from most other Indomalayan species of the genus by its characteristic colouration. From the dorsal colouration S. hunteri is near to S. chevrolati from the Lesser Sunda Islands which is generally smaller and more elongated. Furthermore, both species can be separated by the shapes of their median lobes.

Distribution.

The most widespread and common species of the genus. India: Andhra Pradesh, Nagaland, Tamil Nadu, Uttarakhand, Uttar Pradesh, West Bengal, and Andaman and Nicobar Islands ( Ghosh and Nilsson 2012; Ghosh and Gupta 2012); Nepal, China ( Jäch et al. 2012), South Korea (Jeju-do Island) ( Park et al. 2009; Lee et al. 2014), Japan (old record from Higo Province, Sharp 1884), Myanmar, Cambodia, Laos, Vietnam, West Malaysia, Indonesia (Sumatra, Java, Bali) (Fig. 30 ). The recent Indian records from the states Kerala, Karnataka, Maharashtra, and Tamil Nadu seems doubtful as the photograph in Deb and Subramanian (2023) clearly shows S. festivus and its median lobe, not S. hunteri . Specimens were collected from near sea level to 2900 m a. s. l.

Habitat.

Sandracottus hunteri inhabits a broad variety of pools, puddles, ditches, and swamps of different sizes, shaded and exposed ( Alarie et al. 2023). In Laos numerous specimens were collected by local collectors in pools in the littoral zone of a lake (Figs 37 , 38 ). In Thailand, near Salawin NP, S. hunteri was found in open and muddy puddles made by elephants (Fig. 40 ) and near the King Cobra Cave at Sakaerat Biosphere Reserve, in a large rest pool of a rocky stream bed enriched with decaying leaves and twigs (Fig. 41 ). In Myanmar S. hunteri was collected from roadside pools on a muddy limestone bed with decomposing leaves ( Alarie et al. 2023). Sandracottus hunteri also occurs in cultivated areas and is not restricted to forested sites. On Bali the species was collected several times in smaller pools of intermittent streams with decomposing leaves ( Suprayitno et al. 2017). The species was attracted to light. All instar larvae were recently described by Alarie et al. (2023).

Figures

Figure 30. Distribution of Sandracottus hunteri, the most widespread species of the genus.

Figures 4–7. Habitus of 4 Sandracottus dejeanii (South India) 5 S. festivus (Sri Lanka) 6 S. femoralis (Papua New Guinea, Rigo) 7 S. hunteri (Thailand).

Figures 16–19. Median lobe of aedeagus in ventral view (a), and right paramere in lateral view (b): 16 Sandracottus festivus 17 S. femoralis 18 S. hunteri and 19 S. insignis.

Figures 37–42. Habitats of Sandracottus 37, 38 Laos, Attapeu prov., Annam Highlands Mts., Dong Amphan NBCA, Nong Fa [puddles near lake shore] and lake, habitat of S. hunteri 39 Laos, Katamtok, rest pools along a rocky river bed, habitat of S. maculatus (photos: J. Hájek) 40 Thailand, near Salawin NP, open and muddy puddle made by elephants (photo: J. Šťastný), and 41 Sakaerat Biosphere Reserve, rocky pool in an otherwise dry stream bed, habitats of S. hunteri and S. maculatus (photo: W. Atthakor) 42 Philippines, Luzon, Pampanga Prov., Bano San Juan, Mt. Arayat NP, spring creek, habitat of S. insignis (photo: H. Freitag).

Figures 50–53. Habitus of 50 Sandracottus dejeanii (India, Himachal Pradesh, Kangra Valley, Dharamsala) 51 S. festivus (Sri Lanka, Ratnapura, Sincharaja 52 S. femoralis (Indonesia, West Papua, Kapupaten Paniai, Nabire) 53 S. hunteri (China, Central Sichuan).

Figures 8–11. Habitus of 8 Sandracottus insignis (Luzon, Philippines) 9 S. jaechi (paratype from Nuwara Eliya, Sri Lanka) 10 S. maculatus (Thailand) 11 S. rotundus (Sulawesi, Togian Islands).