Tijdschrift voor Entomologie 160 (2017) 89–138
An initial survey of aquatic and semi-aquatic Heteroptera (Insecta) from the Cardamom Mountains and adjacent uplands of southwestern Cambodia, with descriptions of four new species Dan A. Polhemus
Previous collections of aquatic Heteroptera from Cambodia have been limited, and the biota of the country has remained essentially undocumented until the past several years. Recent surveys of aquatic Heteroptera in the Cardamom Mountains and adjacent Kirirom and Bokor plateaus of southwestern Cambodia, coupled with previous literature records, demonstrate that 11 families, 35 genera, and 68 species of water bugs occur in this area. These collections include 13 genus records and 37 species records newly listed for the country of Cambodia. The following four new species are described based on these recent surveys: Amemboa cambodiana n. sp. (Gerridae); Microvelia penglyi n. sp., Microvelia setifera n. sp. and Microvelia bokor n. sp. (all Veliidae). Based on an updated checklist provided herein, the aquatic Heteroptera biota of Cambodia as currently known consists of 78 species, and has an endemism rate of 7.7%, although these numbers should be considered provisional pending further sampling. Keywords: Heteroptera; Cambodia; water bugs; new species; new records Dan A. Polhemus, Department of Natural Sciences, Bishop Museum, 1525 Bernice Street, Honolulu, HI 96817 USA. [email protected]
Introduction of collections or species records from the country in Aquatic and semi-aquatic Heteroptera, commonly the period preceding World War II. Following that known as water bugs, are a group of worldwide dis- war, the country’s traumatic social and political his- tribution with a well-developed base of taxonomy. As tory left it isolated until the 1990s, and therefore such, they have proven to be a useful entomological unvisited by collectors. Despite a renewed interest component for biodiversity surveys in many parts of in the survey of aquatic Heteroptera in neighbor- the world. This paper provides the results of targeted ing countries, such as Thailand and Vietnam, from collections of this group in the Cardamom Moun- the mid-1980s onward, Cambodia continued to re- tains area of southwestern Cambodia. The paucity main overlooked and unvisited, in part because of of previous work is illustrated by the fact that of the the hazards involved with the numerous land mines 78 species listed in this paper, 37 represent new and that had been planted throughout the country. Only previously unpublished records for the country. in 2017 was a first preliminary checklist of aquatic The aquatic Heteroptera biota of Cambodia re- Heteroptera published for the country, containing mained essentially un-sampled until the last several records of 41 species, based primarily on collec- years. In contrast to other French colonial possessions tions from nine widely scattered localities (Zettel in Indochina, there seems to be no documentation et al. 2017). By contrast, the current report, although
Tijdschrift voor Entomologie 160: 89–138, Appendices 1–2, Figs 1–58. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 15 December 2017. DOI 10.1163/22119434-00002068 Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Fig. 1. View to the southeast from a high point on the Kirirom Plateau at 715 m elevation, showing typical topog- raphy and vegetation of the southwestern Cambodian highlands. The forest in the foreground on the left side of the valley is dominated by Tenassarim pine (Pinus merkusii), while the forest on the right side of the valley is a wetter semi-deciduous broadleaf forest. The blue ridge in the distance is the Elephant Mountains, terminating in the Bokor Plateau visible in the left-central portion of the uplift, and supporting montane evergreen broadleaf rainforest. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Fig. 2. Wet lowland evergreen broadleaf forest near Thmar Bang village at about 400 m elevation, on the southeast- ern flank of the Cardamom Mountains facing the Gulf of Thailand. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Figs 3–6. Exemplar forest formations of the southwestern Cambodian highlands. — 3, Wet montane broadleaf rainforest, Bokor Plateau summit, 1000 m; 4, middle elevation pine forest dominated by Pinus merkusii, Kirirom Plateau, 700 m; 5, semi-deciduous broadleaf forest, Cardamom Mountain foothills at Rolak village, 150 m; 6, dry deciduous broadleaf forest dominated by Dipterocarpus obtusifolius, northeastern base of Kirirom Plateau, 100 m. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Figs 7–10. Exemplar freshwater habitats of southwestern Cambodia. — 7, Stream in bed of sandstone plates on the road to Thmar Bang village, 355 m elev., Koh Kong Prov., CL 6017; 8, stream in semi-deciduous broadleaf forest at Rolak village, along the eastern edge of the Cardamom Mountains, 145 m elev., Kampong Speu Prov., CL 6041; 9, lowland stream bordered by bamboo east of Aural, flowing in a bed of sand and granitic cobbles derived from the Phnom Aural massif, 75 m elev., Kampong Speu Prov., CL 6044; 10, upland pond surrounded by montane rain forest on the Bokor Plateau, 985 m elev., Kampot Prov., CL 6036. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Fig. 11. Map of aquatic Heteroptera collecting sites in southwestern Cambodia, plotted against vegetation classes. Dark green areas indicate semi-deciduous or evergreen broadleaf forests; pale green areas indicate deciduous diptero- carp forest; tan areas indicate cultivated lowlands; pale blue areas indicate wetlands.
Site 2 Cambodia, Koh Kong Prov., Cardamom 11°36'13''N, 103°29'34''E, water temp. Mountains, shaded forest streamlet on road 25.5°C, 31 August 2015, 10:00–10:15 hrs, to Thmar Bang village, 420 m [1375 ft], CL 6018. Large, swift mountain river with 11°36'14''N, 103°15'38''E, water temp. banks shaded by low elevation evergreen 26°C, 30 August 2015, 16:45–17:30 hrs, rainforest. CL 6016. Rocky first-order stream densely shaded by dwarf evergreen rainforest forest Site 5 Cambodia, Koh Kong Prov., Cardamom on sandstone. Mountains, ponds and ditches at Thmar Daun Peov village, 192 m [630 ft], Site 3 Cambodia, Koh Kong Prov., Cardamom 11°37'43''N, 103°31'46''E, water temp. Mountains, cascading stream in sandstone 28°C, 31 August 2015, 11:45–14:00 hrs, CL bed on road to Thmar Bang village, 355 m 6019. Artificial ponds and ditches set amid [1165 ft], 11°35'26''N, 103°13'50''E, water village fields, mostly unshaded. temp. 26°C, 30 August 2015, 17:45–18:30 hrs, CL 6017. Rocky first-order upland Site 6 Cambodia, Koh Kong Prov., Cardamom stream dropping through bed of sandstone Mountains, Toeuk Noeng River, 9 km SSE of plates, partially shaded by dwarf evergreen Thmar Bang village crossroads, 165 m [550 rainforest forest on sandstone (Fig. 7). ft], 11°37'33''N, 103°28'51''E, water temp. 26°C, 31 August 2015, 14:45–16:00 hrs, Site 4 Cambodia, Koh Kong Prov., Cardamom CL 6020. Moderately large, swiftly flowing Mountains, Arang River, 152 m [500 ft], river with alternating deep pools and rapids Downloaded from Brill.com10/07/2021 06:11:13PM via free access
over exposures of sandstone bedrock, banks forest, ~3 km NW of Chambok village, shaded by montane evergreen rainforest on 95 m [315 ft], 12°24'19''N, 104°06'01''E, sandstone. water temp. 28.5°C, 21 May 2016, 09:30– 10:30 hrs, CL 6033. Small stream in sandy Site 7 Cambodia, Koh Kong Prov., Cardamom bed with alternating shallow riffles and Mountains, stream ~31 km NE of Koh deeper pools, partially shaded by semi- Kong on road to Pursat, 310 m [1020 ft], decidous mesic forest, bamboo and second- 11°41'43''N, 103°06'57''E, water temp. ary thorn scrub. 26°C, 1 September 2015, 10:00–10:40 hrs, CL 6021. Smoothly flowing, moderate sized Site 13 Cambodia, Kampong Speu Prov., Sva Slab stream with banks shaded by montane ever- river at rapids in igneous bedrock, ~4.5 km green rainforest. NW of Chambok village, 95 m [310 ft], 12°24'45''N, 104°05'025''E, water temp. Site 8 Cambodia, Koh Kong Prov., Cardamom 30.5°C, 21 May 2016, 11:00–12:15 hrs, Mountains, stream in sandstone bed cross- CL 6034. Large premontane river with deep ing pine barren on road from Koh Kong pools, traversing sill of diabase bedrock with to Pursat, 405 m [1330 ft], 11°49'29''N, scattered water-filled potholes, banks par- 103°04'14''E, water temp. 25°C, 1 Septem- tially shaded by semi-decidous mesic forest, ber 2015, 11:50–12:20 hrs, CL 6022. Swiftly bamboo and secondary thorn scrub. flowing, moderate-sized stream emerging from montane evergreen broadleaf forest and Site 14 Cambodia, Kampot Prov., Elephant Moun- traversing open, grassy pine barren with scat- tains, Bokor Plateau, rocky stream in wet tered Pinus merkusii. upland forest, ~20 km from Hwy 3 on road to Bokor resort, 965 m [3165 ft], Site 9 Cambodia, Koh Kong Prov., Cardamom 10°37'33''N, 104°04'10''E, water temp. Mountains, ponded stream on road from 22°C, 22 May 2016, 08:00–09:00 hrs, CL Koh Kong to Pursat, 540 m [1770 ft], 6035. Small stream heavily shaded by dense 12°00'58''N, 103°10'47''E, water temp. evergreen montane rainforest. 24.5°C, 1 September 2015, 13:45–14:30 hrs, CL 6023. Stream emerging from montane Site 15 Cambodia, Kampot Prov., Bokor Plateau, evergreen rainforest and forming large, un- pond in wet upland forest, ~22 km from shaded pond on upstream side of road, with Hwy 3 on road to Bokor resort, 985 m swift outlet through road culvert. [3240 ft], 10°37'42''N, 104°03'14''E, wa- ter temp. 22°C, 22 May 2016, 09:30–10:15 Site 10 Cambodia, Pursat Prov., Cardamom Moun- hrs, CL 6036. Deep pond with grassy mar- tains, O Sarb Morth stream (= Sap Mat gins, set amid dense evergreen montane stream), N of Vealveang village on road rainforest (Fig. 10). to Pursat, 230 m [760 ft], 12°18'31''N, 103°06'36''E, water temp. 27°C, 2 Sep- Site 16 Cambodia, Kampot Prov., Elephant Moun- tember 2015, 08:00–09:30 hrs, CL 6024. tains, Bokor Plateau, stream in sandstone Shallow, moderately swift stream in gravel bed, ~15 km from Hwy 3 on road to Bokor bed with spring-fed side channel, partially resort, 815 m [2675 ft], 10°37'48''N, shaded by disturbed evergreen rainforest and 104°05'11''E, water temp. 23°C, 22 May secondary regrowth. 2016, 11:00–12:15 hrs, CL 6037. Steeply dropping stream in bed of horizontal sand- Site 11 Cambodia, Kampong Chhnang Prov., Bori stone strata, with alternating pools and small Bo River at Hwy 4 bridge, 20 m [70 ft], trickling cascades, partially shaded by ever- 12°23'04''N, 104°29'11''E, water temp. green montane rainforest. 29°C, 2 September 2015, 15:00–15:45 hrs, CL 6025. Broad, shallow, unshaded Site 17 Cambodia, Sihanoukville Prov., Carda- river in sandy bed, bordered by secondary mom Mountains, river in sandstone bed at vegetation. Thmor Roung recreation area, 65 m [210 ft], 11°11'05''N, 103°56'00''E, water temp. Site 12 Cambodia, Kampong Speu Prov., small per- 28.5°C, 24 May 2016, 10:00–11:30 hrs, CL ennial stream in disturbed semi-deciduous 6038. Large premontane river flowing over
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sandstone bedrock and gravel, with alternat- and crossroads, 75 m [255 ft], 11°38'57''N, ing deep pools and shallow rapids, bordered 104°11'46''E, water temp. 30°C, 29 May by disturbed low elevation evergreen rain- 2016, 11:00–11:30 hrs, CL 6044. Moder- forest on sandstone ately flowing stream in gravel bed with alter- nating deep pools and shallow riffles, shaded Site 18 Cambodia, Sihanoukville Prov., Cardamom along its margins by bamboo and secondary Mountains, rocky stream crossing Hwy 4, thorn scrub (Fig. 9). ~6 mi. SW of pass at Pich Nil, 45 m [155 ft], 11°11'07''N, 104°01'52''E, water temp. Site 24 Cambodia, Kampong Speu Prov., outflow 28.5°C, 24 May 2016, 12:00–13:00 hrs, CL stream from diked wetlands on Hwy 4, 10 6039. Moderately large, swift stream in bed km E of Krong Chbar Mon, 35 m [115 ft], of boulders, rocks and gravel, bordered by 11°28'40''N, 104°35'56''E, water temp. disturbed secondary vegetation. 31°C, 29 May 2016, 12:45–13:30 hrs, CL 6045. Moderately swift stream in unshaded Site 19 Cambodia, Kampong Speu Prov., hot artificial channel draining extensive diked springs and outflow creek 9.5 km W of wetlands. Aoral District village and crossroads, 105 m [340 ft], 11°43'26''N, 104°03'19''E, water Site 25 Cambodia, Kampong Speu Prov., Kirirom temp. 45°C (spring head), 37°C (outflow Plateau, Tachot Stream, crossing at first creek), 28 May 2016, 07:15–07:40 hrs, CL bridge on road up to summit, 315 m [1035 6040. Unshaded thermal spring outflows ft], 11°32'42''N, 104°55'23''E, water temp. on valley floor surrounded by open grassy 27°C, 4 June 2016, 08:45–09:20 hrs, CL pastures. Note: no aquatic Heteropera were 6046. Small, slowly flowing stream with taken at this unusual site. deep pools and shallow riffles, surrounded by recently cleared gardens and second growth. Site 20 Cambodia, Kampong Speu Prov., Carda- mom Mountains, trib. to Thom River, 1 km Site 26 Cambodia, Kampong Speu Prov., Kirirom E of Rolak Kong Cheung village at forest Plateau, Tasek Stream, crossing at second ranger station, 145 m [474 ft], 11°43'30''N, bridge on road up to summit, 640 m [2095 103°50'54''E, water temp. 28.5°C, 28 ft], 11°19'23''N, 104°04'59''E, water temp. May 2016, 09:00–1300 hrs, 29 May 2016, 25°C, 4 June 2016, 9:30–10:30 hrs, CL 08:00–09:00 hrs, CL 6041. Moderate sized 6047. Moderately swift stream in gravel premontane foreland stream in bed of rocks bed, heavily shaded by broadleaf evergreen and gravel, partially shaded by semi-decidu- gallery forest corridor within upland pine ous mesic forest (Fig. 8). forest dominated by Pinus merkusii. Site 21 Cambodia, Kampong Speu Prov., Carda- Site 27 Cambodia, Kampong Speu Prov., Kirirom mom Mountains, Thom River, 2 km W of Plateau, roadside reservoir in pine forest, 690 Rolak Kong Cheung village, 185 m [610 m [2260 ft], 11°20'01''N, 104°03'08''E, ft], 11°43'15''N, 103°49'21''E, water temp. water temp. 32°C, 4 June 2016, 10:45– 28.5°C, 28 May 2016, 14:00–15:00 hrs, CL 11:30 hrs, CL 6048. Open reservoir with 6042. Moderately large, swift premontane grassy margins surrounded by upland pine river with extensive overflow channels, heav- forest dominated by Pinus merkusii. ily shaded by evergreen gallery forest. All species listed in the following taxonomic section Site 22 Cambodia, Kampong Speu Prov., rocky represent new records for the country of Cambodia stream 7 km E of Rolak Kong Cheung village, unless otherwise noted. 135 m [440 ft], 11°42'43''N, 103°53'55''E, The following abbreviations are used for specimen water temp. 28.5°C, 29 May 2016, 09:00– depositories: 09:45 hrs, CL 6043. Slowly flowing stream in gravel bed with alternating deep pools and shallow riffles, shaded along its margins by BPBM Bernice P. Bishop Museum, Honolulu, bamboo and secondary thorn scrub. Hawaii, USA USNM United States National Museum of Site 23 Cambodia, Kampong Speu Prov., stream in Natural History, Smithsonian Institution, gravel bed 9 km E of Aoral District village Washington, DC, USA Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Taxonomy D.A. Polhemus (BPBM); 6 wingless males, 6 wing- In the species treatments below, only abbreviated less females, Cardamom Mountains, Toeuk Noeng locality data is provided. For complete data regard- River, 9 km SSE of Thmar Bang village crossroads, ing any particular record, the reader is referred to 165 m [550 ft], 11°37'33''N, 103°28'51''E, water the preceding list of sampling localities. Complete temp. 26°C, 31 August 2015, 14:45–16:00 hrs, CL locality data for comparative Vietnamese localities 6020, D.A. Polhemus (BPBM). referenced in the text may be found in Appendix 2. Description Infraorder Gerromorpha Apterous male (Fig. 14). Body length 3.60 mm; max- Family Gerridae imum width (across thorax) 1.25 mm. Subfamily Cylindrostethinae Matsuda Color. Black Russet brown with limited dark brown to black markings on pronotum, mesonotum, dorsal Cylindrostethus costalis Schmidt abdomen, pleurae and venter (Fig. 14), and patches Cylindrostethus costalis Schmidt, 1915: 364. of silvery setae present on acetabulae and pleurae. Head russet brown with a faintly indicated V-shaped Material examined. Cambodia, Kampong Speu Prov.: black mark posteromedially; eyes dark red; tylus 1 winged male, 4 winged females, nr. Chambok, CL 6033; and antennal tubercles dark, bearing patches of sil- 2 winged males, 3 winged females, nr. Rolak, CL 6041; very setae; antennae with segment I medium brown, 1 winged male, 1 winged female, nr. Rolak, CL 6042; segments II–IV darker brown; rostrum yellowish- 1 winged female, nr. Rolak, CL 6043; 2 winged females, brown, with a ventromedial line and apex black. Pro- nr. Aoral, CL 6044; 1 winged male, Kirirom, CL 6047. notum russet brown, with a pair (1 + 1) of irregularly Ecological notes. This large, black-colored gerrid elongate dark patches to either side of longitudinal species was common on the lowland streams of midline, these patches initiating at anterior prono- Kampong Speu Province. It seems to be an adapt- tal margin, tapering posteriorly, not quite attaining able generalist which can tolerate a modest degree of posterior pronotal margin (Fig. 14); propleura with habitat alteration and degraded water quality. narrow, weakly indicated longitudinal dark stripe centrally, bearing small patches of silvery setae be- Subfamily Eotrechinae Matsuda low eyes and on fore acetabula. Mesonotum russet brown, with a pair (1 + 1) of irregularly elongate Amemboa cambodiana n. sp. dark patches to either side of longitudinal midline, Figs 14–16, 19–24 these patches extending posteriorly for only half the mesonotal length (Fig. 14); mesopleuron with a Type material. Holotype, wingless male, Cambodia, strongly developed longitudinal stripe running from Kampot Prov., Elephant Mountains, Bokor Plateau, anterior margin to near metanotal suture, inner por- stream in sandstone bed, ~15 km from Hwy 3 on tion of this dark stripe bearing band of silvery se- road to Bokor resort, 815 m [2675 ft], 10°37'48''N, tae; mesoacetabula dark, bearing prominent patch 104°05'11''E, water temp. 23°C, 22 May 2016, of silvery setae. Metanotum broadly dark centrally 11:00–12:15 hrs, CL 6037, D.A. Polhemus and across anterior margin, russet brown laterally, (USNM). Paratypes: Cambodia, Kampot Prov., 9 bearing scattered very short, recumbent gold setae winged males, 2 winged females, 10 wingless males, on anterior half, these setae also extending onto ad- 10 wingless females, same data as holotype (USNM, jacent inner angles of metapleuron. Legs medium BPBM); 6 winged males, 6 winged females, 2 wing- brown, tarsi black. Abdominal tergites dark brown, less females, Elephant Mountains, Bokor Plateau, with central portions of tergites VI–VIII russet rocky stream in wet upland forest, ~20 km from brown, connexiva dark russet brown; both abdomi- Hwy 3 on road to Bokor resort, 965 m [3165 ft], nal tergites and connexiva bearing scattered very 10°37'33''N, 104°04'10''E, water temp. 22°C, 22 short, recumbent gold setae. Ventral surface yellow- May 2016, CL 6035, D.A. Polhemus (BPBM); Kam- ish-brown, posteromedial portion of mesosternum pong Chhnang Prov.: 10 wingless males, 6 wingless and longitudinal midline of abdomen suffused with females, Bori Bo River at Hwy 4 bridge, 20 m [70 darker brown. ft], 12°23'04''N, 104°29'11''E, water temp. 29°C, 2 Structural characteristics. Head — Width/length = September 2015, 15:00–15:45 hrs, CL 6025, D.A. 0.85/0.70, eye width 0.25, interocular width 0.50; Polhemus (BPBM). Koh Kong Prov.: 1 wingless lengths of antennal segments I–IV = 0.75, 0.70, male, Cardamom Mountains, cascading stream in 0.80, 1.05 respectively; rostrum length 1.45, extend- sandstone bed on road to Thmar Bang village, 355 m ing on mesosternum. [1165 ft], 11°35'26''N, 103°13'50''E, water temp. Pronotum — Width/length = 0.90/0.50; mesono- 26°C, 30 August 2015, 17:45–18:30 hrs, CL 6017, tum length 1.15; metanotum length 0.25. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Figs 12–16. Amemboa species, dorsal habitus, appendages omitted. Figs 12, 13. A. burmensis, paratype specimens from Burma, Shingbwiyang. 12. Apterous male. 13. Apterous female. Figs 14–16. A. cambodiana, n. sp., specimens from Cambodia, Bokor Plateau, CL 6037. 14. Apterous male. 15. Apterous female.Downloaded 16. Macropterous from Brill.com10/07/2021 male. 06:11:13PM via free access
Figs 17–19. Amemboa species, male right fore femur, dorsal view. 17. A. kumari, paratype specimen from India, Karnataka, Mudigere. 18. A. burmensis, paratype specimen from Burma, Shingbwiyang. 19. A. cambodiana, n. sp., specimen from Cambodia, Kampot Prov., Bokor Plateau, CL 6037. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Figs 20–24. Amemboa cambodiana n. sp., male genitalia, specimen from Cambodia, Kampot Prov., Bokor Plateau, CL 6037. 20. Proctiger, dorsal view. 21. Pygophore, ventral view. 22. Pygophore, left ventrolateral oblique view, showing form of posterior projection. 23. Endosoma, lateral view. 24. Endosoma, dorsal view.
Legs — Fore femur slightly incrassate, bearing 2 middle femur/tibia/tarsal I/tarsal II = 3.10/2.30/ tufts of stiff black setae on distal half, these tufts of 0.90/0.45; hind femur/tibia/tarsal I/tarsal II = roughly equal size, separated by a gap about 2× the 3.20/1.20/0.50/0.35. width of each tuft (Fig. 19); fore tibia very slightly Male genitalia — Pygophore roughly triangular, sinuate on basal half, bearing a fringe of short, dense bearing a small, bulb-like apical process (Fig. 20). black setae along inner margin on basal two-thirds Proctiger with lateral process elongate, slightly ex- (Fig. 19); lengths of leg segments as follows: fore ceeding posterior margin of abdominal tergite VIII, femur/tibia/tarsal I/tarsal II/ = 1.35/1.30/0.20/0.30; with distal sections gently curving and slightly Downloaded from Brill.com10/07/2021 06:11:13PM via free access
A. cambodiana taken syntopically with any of them, bed of a stream that descended from the flanks of this being its co-occurrence with A. cristata on sand- the Bokor Plateau. It was also seen in lesser num- bars at the Bori Bo River in Kampong Chhnang bers on sandstone-bedded streams in the Cardamom Province. Mountains.
Amemboa cristata J. Polhemus & Andersen Subfamily Gerrinae Leach Amemboa cristata J. Polhemus & Andersen, 1984: 95. Limnogonus fossarum fossarum (Fabricius) Cimex fossarum Fabricius, 1775: 727. Material examined. Cambodia, Kampong Chhnang Gerris fossarum: Fabricius, 1794: 188. Prov.: 1 winged female, 1 wingless female, Bori Bo, CL Hydrometra fossarum: Fabricius, 1803: 258. 6025. Gerris discolor Stål, 1859: 265; syn. by Lundblad, Ecological notes. This species, which can be recog- 1933: 377. nized by the acuminate tufts of dark hairs at the pos- Limnogonus fossarum: Stål, 1868: 133. terior margin of the female abdomen, was taken as a Limnogonus discolor: Stål, 1868: 133. few scattered individuals on sandbars along the low- Tenagogonus nymphae Esaki, 1925: 58; syn. by Esaki, land Bori Bo River which drains the Mekong River 1926: 181. side of the Cardamom uplift. Tenagogonus okinawanus Esaki, 1925: 59; syn. by Andersen, 1975: 30. Amemboa javanica Lundblad Limnogonus okinawanus: Lundblad, 1933: 371. Amemboa javanica Lundblad, 1933: 405. Limnogonus fossarum fossarum: Andersen, 1975: 30.
Material examined. Cambodia, Kampong Speu Prov.: 2 Material examined. Cambodia, Kampong Speu Prov.: 2 winged males, 1 wingless male, 1 wingless female, nr. Ro- wingless females, Kirirom, CL 6046. Koh Kong Prov.: 1 lak, CL 6041. winged male, nr. Thmar Bang, CL 6016. Ecological notes. Amemboa javanica was found as Discussion. The nominate subspecies of L. fossarum scattered individuals along the margins of pools which occurs in Cambodia is widely distributed along a rocky stream at Rolak village (Fig. 8), in the from India and Ceylon eastward through Indochina premontane foreland near the northeastern margin to China, Japan, the Philippines, the Malay Penin- of the Cardamom Mountains. sula, Sumatra and Borneo (Andersen 1975).
Amemboa speciosa J. Polhemus & Andersen Limnogonus hungerfordi Andersen Amemboa speciosa J. Polhemus & Andersen, 1984: 94. Limonogonus hungerfordi Andersen, 1975: 46.
Material examined. Cambodia, Koh Kong Prov.: 1 Material examined. Cambodia, Pursat Prov.: 1 wingless winged male, Koh Kong–Pursat, CL 6023. male, 1 wingless female, nr. Vealveang, CL 6024. Discussion. As noted by Andersen (1975), this spe- Ecological notes. A pair of specimens representing cies is externally all but indistinguishable from this species was splashed from a half-submerged log (Montrousier), with definitive identifi- lying in the middle of a deep pool on a quiet, shaded, L. luctuosus cation reliant upon dissection of the male genitalia spring-fed tributary backwater to the O Sarb Morth and examination of the posterior margin of the male stream just north of Vealveang. dorsal vesical plate. The male genitalia of the Cam- bodian specimen in hand show that the shape of this Onychotrechus esakii Andersen plate falls within the range of variation illustrated for Onychotrechus esakii Andersen, 1980: 127. L. hungerfordi by Andersen (1975, Figs 120–122). As noted by Cheng et al. (2001), although widely Material examined. Cambodia, Kampot Prov.: 5 wingless distributed across a range encompassing northeast- males, 4 wingless females, Bokor Plateau, CL 6037. Koh ern Australia, New Guinea, the Malay Archipelago, Kong Prov.: 1 wingless male, nr. Thmar Bang, CL 6017; 4 the Malay Peninsula, the Philippines and Taiwan wingless males, Touek Noeng, CL 6020. (Andersen 1975), this is a rare species in continen- Ecological notes. This species is the most widespread tal Southeast Asia, and the current record is the first Onychotrechus in Southeast Asia (Andersen 1980), for Cambodia, and apparently for Indochina as a and was abundant on wet sandstone plates in the whole.
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Limnogonus nitidus (Mayr) CL 6048. Kampot Prov.: 1 winged male, 1 winged female, Hydrometra nitida Mayr, 1865: 443. 2 wingless males, 1 wingless female, Bokor Plateau, CL Gerris nitida: Distant, 1904: 178. 6036. Limnogonus nitidus: Kirkaldy, 1908: 21. Ecological notes. This is a widespread species on len- tic habitats throughout Southeast Asia. Material examined. Cambodia, Sihanoukville Prov.: 1 winged male, nr. Pich Nil, CL 6039. Pursat Prov.: 1 winged male, nr. Vealveang, CL 6024. Subfamily Halobatinae Bianchi Discussion. This species is widespread, with range ex- tending from the Maldive Islands through Ceylon, Metrocoris nigrofascioides Chen & Nieser India, Indochina, the Malay Peninsula, Sumatra, Metrocoris nigrofascioides Chen & Nieser, 1993: 16. Java and Borneo (Andersen 1975). Material examined. Cambodia, Kampong Speu Prov.: 2 Limnometra matsudai (Miyamoto) wingless females, Sva Slab, CL 6034. Koh Kong Prov.: 2 Tenagogonus (Limnometra) matsudai Miyamoto, 1967: winged females, 4 wingless males, 1 wingless female, nr. 223. Thmar Bang, CL 6016; 3 winged males, 1 winged female, Limnometra matsudai: Andersen, 1995: 118. nr. Thmar Bang, CL 6017; 2 wingless males, 2 wingless fe- males, Touek Noeng, CL 6020; 1 wingless male, 4 wingless Material examined. Cambodia, Kampot Prov.: 1 wingless females, Koh Kong–Pursat, CL 6023. female, Bokor Plateau, CL 6035; 2 winged males, 4 winged Ecological notes. This species occurs as scattered pop- females, 1 wingless male, 4 wingless females, Bokor Plateau, ulations at intermediate elevations in the Cardamom CL 6037. Koh Kong Prov.: 2 wingless males, 5 wingless fe- Mountains (Tran & Polhemus 2017). males, nr. Thmar Bang, CL 6016; 2 wingless males, Toeuk Noeng, CL 6020. Pursat Prov.: 1 winged female, 1 wing- Metrocoris tenuicornis Esaki less female, nr. Vealveang, CL 6024. Sihanoukville Prov.: 1 Metrocoris tenuicornis Esaki 1926: 125. winged male, 2 wingless females, Thmar Roung, CL 6038. Ecological notes. This is a common species along first- and second-order upland streams draining the Material examined. Cambodia, Koh Kong Prov.: 2 Cardamom Mountains, Kirirom Plateau, and Bokor wingless males, 3 wingless females, Koh Kong–Pursat, Plateau. In these uplifts it was often one of the only CL 6021. species encountered on small headwater streamlets at Ecological notes. This species is widely distributed elevations above 500 m. on lowland forest streams throughout Southeast Asia, but so far encountered in Cambodia only on Limnometra octopunctata Hungerford a single stream in the mountains behind Koh Kong. Limnometra octopunctata Hungerford, 1955: 67. Geographic variation in paramere shape suggests that more than a single species may be involved Material examined. Cambodia, Sihanoukville Prov.: 1 across its wide range in Indochina (Tran & Polhemus winged female, Thmar Roung, CL 6038. 2017). Ecological notes. Limnometra octopunctata appears to be uncommon and localized in southwestern Cam- Ventidius (Ventidius) distanti Paiva bodia, being taken only at a single locality at Thmar Ventidius distanti Paiva, 1918: 25. Roung where a sandstone-bedded river exits the Ventidius modulatus Lundblad, 1933: 339; syn. by mountain foothills onto the coastal plain. Chen, Nieser & Zettel, 2005: 404. Ventidius chinai Hungerford & Matsuda, 1960: 331; Neogerris parvula (Stål) syn. by Chen & Zettel, 1998: 170. Gerris parvula Stål, 1859: 265. Ventidius pubescens Cheng, 1965: 160; syn. by Chen & Limnogonus parvulus: Stål, 1868: 133. Zettel, 1998: 170. Gerris tristan Kirkaldy, 1899: 88; syn. by Lundblad, 1933: 385. Material examined. Cambodia, Kampong Chhnang Gerris ysolt Breddin, 1905: 130; syn. by Lundblad, Prov.: 3 winged females, 3 wingless males, 6 wingless fe- 1933: 385. males, Bori Bo, CL 6025. Kampong Speu Prov.: 2 winged Neogerris parvulus: Andersen, 1975: 86. females, nr. Rolak, CL 6043; 3 winged females, nr. Aoral, CL 6044; 6 wingless males, 15 wingless females, nr. Krong Material examined. Cambodia, Kampong Speu Prov.: 1 Chbar Mon, CL 6045. Koh Kong Prov.: 1 winged male, winged female, Sva Slab, CL 6034; 1 winged female, nr. 2 winged females, 1 wingless male, Thmar Daun Peov, Krong Chbar Mon, CL 6045; 1 wingless female, Kirirom, CL 6019; 9 wingless males, 12 wingless females, Koh
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Kong–Pursat, CL 6021. Pursat Prov.: 1 wingless male, 1 were taken on upland ponds (Fig. 4), and on calm wingless female, nr. Vealveang, CL 6024. Sihanoukville pools or backwaters of streams in the premontane Prov., 3 winged males, 10 wingless females, nr. Pich Nil, foreland of the Cardamom Mountains and the Kiri- CL 6039. rom Plateau. Discussion. This species was originally described from the He-Ho River gorge in Yawngshwe State, Lathriobates johorensis (J. Polhemus & D. Polhemus) Burma. When preparing their revision of Ventidius, Cryptobates johorensis J. Polhemus & D. Polhemus, Chen & Zettel (1998) did not examine the types of 1995: 102. this species, but noted that it had close similarities to Lathriobates johorensis: J. Polhemus, 2004: 212. the widespread V. modulatus Lundblad. It was sub- sequently determined that V. modulatus is in fact a junior synonym of V. distanti (Chen et al. 2005). Material examined. Cambodia, Kampong Speu Prov.: 1 Ecological notes. In Cambodia, this species is com- winged female, 2 wingless females, nr. Chambok, CL 6033. mon and widespread, being found on the smoothly Sihanoukville Prov.: 1 winged female, Thmar Roung, CL flowing streams of the premontane foreland below 6038; 1 winged male, 2 winged females, 1 wingless female, the Cardamom Mountains, sometimes in large nr. Pich Nil, CL 6039. schools, and also ranging upward into the mountains Ecological notes. This species occasionally co-occurs to at least 640 m elevation. with Gnomobates kuiterti, but is larger in size and more greyish in overall coloration. It was originally Ventidius (Ventidioides) karen Lansbury described from Johor, in the southern Malay Penin- Ventidius (Ventidiopsis) karen Lansbury, 1988: 61. sula, with all previous records coming from localities south of the Isthmus of Kra; as such, the new Cam- bodian records represent a northern range extension Material examined. Cambodia, Kampong Speu Prov.: into Indochina. 1 winged male, Kirirom, CL 6047. Sihanoukville Prov.: 2 winged females, 2 wingless males, Thmar Roung, Naboandelus signatus Distant CL 6038. Naboandelus signatus Distant, 1910: 152. Discussion. The dorsal coloration and male genitalic structures of the Cambodian material match the re- description in Chen & Zettel (1998). This species Material examined. Cambodia, Kampong Chhnang appears to be of relatively localized occurrence in Prov.: 2 winged males, 2 wingless males, 2 wingless females, southwestern Cambodia. Bori Bo, CL 6025. Ecological notes. A widespread species ranging from Ceylon through India and Thailand to Cambodia, Subfamily Trepobatinae Matsuda N. signatus was found on open waters along the mar- gins of the shallow, sandy Bori Bo River which drains Gnomobates kuiterti (Hungerford & Matsuda) to the Tonle Sap. Cryptobates kuiterti Hungerford & Matsuda, 1958: 246. Gnomobates kuiterti: J. Polhemus & D. Polhemus, Subfamily Ptilomerinae Bianchi 1995: 108. Ptilomera hylactor Breddin Ptilomera hylactor Breddin, 1903: 148. Material examined. Cambodia, Kampong Speu Prov.: 1 winged male, nr. Chambok, CL 6033; 3 wingless females, nr. Rolak, CL 6043. Kampot Prov: 1 wingless male, 5 Material examined. Cambodia, Pursat Prov.: 2 wingless wingless females, Bokor Plateau, CL 6036. S ihanoukville males, 5 wingless females, nr. Vealveang, CL 6024. Prov.: 2 winged females, 1 wingless male, 1 wingless fe- Ecological notes. This species was encountered in male, nr. Pich Nil, CL 6039. southwestern Cambodia only on a single stream Discussion. With a body length of less than 3.0 mm, immediately outside Vealveang. Ptilomera hylactor this is the smallest species of Trepobatinae occurring is generally a species of larger, more open, gravel- in Cambodia. There is some intraspecific variation in bottomed rivers in comparison to P. tigrina, which color, with specimens from higher elevation on the prefers small to moderate-sized rocky upland streams. Bokor Plateau somewhat darker than those from the lowlands of Kampong Speu Province. Ptilomera tigrina Uhler Ecological notes. This species was described from ma- Ptilomera tigrina Uhler, 1860: 230. terial taken in Burma, and previously known from Ptilomera harpyia Schmidt,1926: 65; syn. by J. Pol- India and Thailand as well. In Cambodia individuals hemus, 1992: 439.
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Material examined. Cambodia, Kampong Speu Prov.: from New Guinea, Borneo and Sumatra (Polhemus 1 wingless female, Kirirom, CL 6046. Koh Kong Prov.: 1 & Karunaratne 1993), but the Cambodia records are winged female, 2 wingless males, 2 wingless females, nr. the first for mainland Southeast Asia. Thmar Bang, CL 6017; 1 wingless male, 1 wingless female, Arang River, CL 6018; 1 wingless male, 2 wingless females, Rhagadotarsus kraepelini Breddin Touek Noeng, CL 6020; 1 wingless male, 1 wingless fe- Rhagadotarsus kraepelini Breddin, 1905: 137. male, Koh Kong–Pursat, CL 6023. Sihanoukville Prov.: 2 wingless males, 2 wingless females, nr. Pich Nil, CL Material examined. Cambodia, Kampong Speu Prov.: 5 6039. wingless males, 6 wingless females, nr. Rolak, CL 6043. Ecological notes. This species was previously recorded from Cambodia by D. Polhemus (2001). Ptilomera ti grina is confined to rocky hill streams and iscommon Discussion. This species has a very wide range, from in the interior Cardamom Mountains. It essentially Ceylon and India eastward through Southeast Asia replaces Cylindrostethus costalis as the large-sized, to Taiwan, Palau and New Guinea (Polhemus & stream-dwelling stream gerrid above 100 m eleva- Karunaratne 1993). Its presence in Cambodia was tion, and is never seen on the alluvial streams of the therefore to be expected. Cambodian Plain frequented by that latter species. The zone of sympatry for the two genera on the flanks Family Hebridae Amyot & Serville of the Kirirom Plateau lay in a zone between 45 and 315 m. The overall distribution of this species in Hebrus nieseri Zettel Southeast Asia was summarized and mapped by D. Hebrus nieseri Zettel, 2004: 544. Polhemus (2001), and further data for southern Thai- land were provided by Raruanysong et al. (2014). Material examined. Cambodia, Pursat Prov.: 1 winged male, nr. Vealveang, CL 6024. Rheumatogonus vietnamensis Zettel & Chen Discussion. The male genitalic structures and small, Rheumatogonus vietnamensis Zettel & Chen, 1996: triangular lateral tubercles on the frons match the il- 171. lustrations in Zettel (2004). The species was origi- nally described from specimens taken in northern Material examined. Cambodia, Sihanoukville Prov.: 3 Thailand, so the Cambodian record is a considerable winged males, 9 wingless males, 5 wingless females, nr. range extension. Pich Nil, CL 6039. Ecological notes. This species was originally de- scribed from Vietnam (Zettel & Chen 1996) and Hebrus cf. polysetosus Zettel subsequently recorded from southern Thailand by Hebrus polysetosus Zettel, 2004: 541. Raruanysong et al. (2014), and from several locali- ties in Cambodia by Zettel et al. (2017). During the Material examined. Cambodia, Kampong Speu Prov.: current surveys it was encountered only at a single 1 winged female, nr. Rolak, CL 6041. site, on a large, rocky stream below the Kirirom hy- Discussion. The one female specimen at hand from a droelectric plant. stream at Rolak village (Fig. 5) conforms to Zettel’s (2004) description of this species in regard to size (length 2.0 mm), blackish coloration with yellow Subfamily Rhagadotarsinae Lundblad legs, the presence of elongate setae on the head, and the shape of the scutellum. A definitive identifica- Rhagadotarsus borneensis J. Polhemus & tion, however, cannot be made in the absence of a Karunaratne male specimen, so the assignment of this specimen to Rhagadotarsus borneensis J. Polhemus & Karuna H. longisetosus must be considered provisional until ratne, 1993: 106. the Rolak specimen can be compared to females in the type series. This species was originally described Material examined. Cambodia, Kampong Speu Prov.: ; from specimens taken in Thailand and southern 1 winged female, nr. Rolak, CL 6041; 1 winged male, 1 Laos, so has a broad range within Indochina. winged female, nr. Rolak, CL 6042. Pursat Prov.: 1 winged male, 2 wingless males, 5 wingless females, nr. Vealveang, CL 6024. Family Hydrometridae Billberg Discussion. The series listed above key to this species on the basis of hind femoral length, and the pres- Hydrometra annamana Hungerford & Evans ence of long setae on antennal segments III and IV Hydrometra annamana Hungerford & Evans, 1934: in the female. This species was previously recorded 68.
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Material examined. Cambodia, Koh Kong Prov.: 1 wing- Hydrometra sumatrana Ruhoff, 1964: 32; syn. by J. less male, Koh Kong–Pursat, CL 6022. Polhemus & Riesen, 1986: 284. Discussion. This is a widespread east Asian species, with a range encompassing Japan, Formosa, China Material examined. Cambodia, Kampong Speu Prov.: 1 and Indochina. Hydrometra annamana was previous- winged male, 1 winged female, Sva Slab, CL 6034. Kam- ly recorded from Thailand and Vietnam (Polhemus pot Prov.: 1 winged male, 1 winged female, Bokor Plateau, & Polhemus 1995), and its presence in Cambodia is CL 6036. thus logical. Discussion. This species is broadly distributed in in- sular Southeast Asia, ranging from northern Australia Hydrometra gilloglyi J. Polhemus & D. Polhemus through the Malay Archipelago and the Philippines Hydrometra gilloglyi J. Polhemus & D. Polhemus, to Indochina and Hainan Island (Polhemus & 1995: 20. Polhemus 1995; Tran et al. 2010).
Material examined. Cambodia, Kampong Speu Prov.: 2 Family Mesoveliidae Douglas & Scott winged males, nr. Chambok, CL 6033. Koh Kong Prov.: 1 wingless male, O An Dart, CL 6015. Pursat Prov.: Mesovelia horvathi Lundblad 4 winged females, 4 wingless males, nr. Vealveang, CL Mesovelia horvathi Lundblad, 1933: 190. 6024. Discussion. Hydrometra gilloglyi as currently inter- preted (Tran et al. 2010) has been previously record- Material examined. Cambodia, Kampong Chhnang ed from Vietnam, Thailand, northern Peninsular Prov.: 2 winged males, 1 winged female, 4 wingless fe- Malaysia, and Hainan Island. The new Cambodian males, Bori Bo, CL 6025. Kampong Speu Prov.: 2 winged records fill an obvious gap in the species’ geographic males, 1 wingless female, Sva Slab, CL 6034; 1 wingless range. female, nr. Krong Chbar Mon, CL 6045; 2 winged fe- males, 1 wingless female, Kirirom, CL 6046. Koh Kong Hydrometra jackzewskii Lundblad Prov.: 1 wingless male, O An Dart, CL 6015: 1 wingless Hydrometra jackzewskii Lundblad, 1933: 433. male, Thmar Don Peov, CL 6019. Pursat Prov.: 1 winged female, 1 wingless male, 1 wingless female, nr. Vealveang, CL 6024. Material examined. Cambodia, Kampong Speu Prov.: 2 Discussion. Mesovelia horvathi is a very widespread males, Kirirom, CL 6047. species, ranging from New Guinea and northern Discussion. This species was previously recorded Australia westward through the Malay Archipelago from Java, Sumatra and Burma (Polhemus & Pol- and the Philippines to Japan and Southeast Asia. Re- hemus 1995), and its discovery in Cambodia serves cords for Thailand and Vietnam were provided by to bridge this previously fragmented distributional Polhemus & Polhemus (2000), but the species was pattern. not previously known from Cambodia.
Hydrometra longicapitis Torre Bueno Mesovelia vittigera Horvath Hydrometra longicapitis Torre Bueno, 1927: 31. Mesovelia vittigera Horvath, 1895: 160.
Material examined. Cambodia, Kampong Speu Prov.: 2 Material examined. Cambodia, Pursat Prov.: 1 wingless females, Kirirom, CL 6047. Pursat Prov.: 1 winged female, male, nr. Vealveang, CL 6024. 2 brachypterous females, nr. Vealveang, CL 6024. Discussion. This is the most widespread Mesovelia Discussion. This species was previously known species in the Eastern Hemisphere, occurring from from Sumatra, Peninsular Malaysia, Thailand and Africa through Asia and eastward into the Pacific is- Borneo (Polhemus & Polhemus 1995), so its pres- lands as far as Guam. Despite this, it was not com- ence in Cambodia was to be expected. As with monly encountered during the current surveys, with Rhagovelia tebakang (see subsequent discussion M. horvathi being far more predominent on standing under that species), it has a range that spans the water habitats in the lowlands of Cambodia. South China Sea, indicating a disjunct Sundaland distribution. Family Veliidae Amyot & Serville Hydrometra orientalis Lundblad Subfamily Haloveliinae Esaki Hydrometra orientalis Lundblad, 1933: 430. Hydrometra insularis Hungerford & Evans, 1934: 76; Strongylovelia albopicta Zettel & Tran syn. by J. Polhemus & Riesen, 1986: 284. Strongylovleia albopicta Zettel & Tran, 2004: 80. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Material examined. Cambodia, Kampong Speu Prov.: larger body sizes, symmetrical male parameres, ab- 1 winged male, 2 wingless males, 11 wingless females, nr. sence of lateral projections on the male proctiger, ab- Rolak, CL 6041. Koh Kong Prov.: 1 wingless male, nr. sence of punctations on the dorsum of the head, and Thmar Bang, CL 6017. Pursat Prov.: 3 wingless males, 4 open female connexival structures which do not con- wingless females, nr. Vealveang, CL 6024. verge over abdominal tergites V–VII. In the key of Ecological notes. This very small species, conspicu- Torre-Bueno (1925) to Indian Microvelia, these new ously marked with black and white, is locally abun- species run to various taxa which, although exhibit- dant along the margins of quiet stream pools. It was ing certain similarities, also have obvious differences, originally described from Vietnam, and until now with these being discussed under each individual known only from that country. species treatment. The genus Microvelia as currently interpreted is undoubtedly para- or polyphyletic, but a reso- Subfamily Microveliinae China & Usinger lution of this problem is beyond the scope of the Genus Microvelia Westwood present paper. The widespread M. douglasi and re- cently described M. falcata are members of the sub- genus Picaultia (Andersen & Weir 2003), whereas Discussion. As noted by Andersen et al. (2002a), the all of the other Microvelia species so far taken in Microvelia fauna of Southeast Asia has not been re- Cambodia are assignable at present to the nominate vised since the work of Lundblad (1933), and many subgenus. undescribed species are known to occur in the re- gion. Three new species of Microvelia are described below, based on recent Cambodian collections. In Key to males of the species of addition, two other species are at hand, represented Microvelia by one or two females only, that also likely represent known from Cambodia new species, but their description is deferred pending 1. Grasping combs present on both fore and the collection of male specimens. The widespread M. middle tibiae; genitalia asymmetrical, with douglasi Scott was also encountered during the cur- right paramere much larger than left �������������� rent surveys, but by contrast the similarly widespread ...... 2 (subgenus Picaultia) M. leveillei (Letheirry), widely referenced in the pre- – Grasping comb present on fore tibia only; vious literature under the now-synonymized name male genitalia symmetrical, with both param- M. diluta Distant, was not. In addition, the very re- eres of the same size and shape ���������������������� cently described M. falcata, an apparent Cambodian ��������������������������������� 3 (subgenus Microvelia) endemic, was taken at a single Cardamom Mountain 2. Male right paramere slender and tapering, foothill locality. acuminate, with apex coming to a sharp, acute The three new species described here appear to angle; body length equal to or exceeding 1.80 belong to a closely related assemblage of species mm ��������� Microvelia (Picaultia) douglasi Scott with blackish to brown ground coloration, extensive – Male right paramere broader, gently curv- patches of silvery hairs on the abdominal dorsum, ing and of relatively even width throughout, and slender, elongate, sickle-shaped male parameres. with apex blunt; body length equal to or The male genitalic structures of all these new species less than 1.65 mm ������������������������� Microvelia are not congruent with any illustrated by Lundblad (Picaultia) falcata Zettel, Phauk, Kheam & Freitag (1933). 3. Body length less than 1.9 mm; fore tibia ex- Andersen et al. (2002a) provided a key to the panded and flattened on distal half (Fig. 38); Microvelia species occurring in Singapore and Pen- abdominal tergites III–V with extensive patch- insular Malaysia, and Torre-Bueno (1925) presented es of recumbent silvery setae (Fig. 37). . . . a key to the species found in India and Ceylon. Use ���������������� Microvelia (Microvelia) bokor n. sp. of these resources, in combination with Lundblad’s – Body length exceeding 2.2 mm; fore tibia (1933) work and examination of comparative mate- not expanded and flattened on distal half; rial at the Smithsonian Institution, has provided ad- abdominal tergites IV and V lacking patch- ditional confidence in the validity of the new species es of recumbent silvery setae (Figs 25, 26, described here. In the key of Andersen et al. (2002a), 32, 33)...... 4 all three of the new species described herein will run 4. Body set with numerous long, erect setae to the couplet containing M. petraeus Andersen, (Figs 32, 33); genital segment projecting Yang & Zettel and M. plumbea Lundblad, the lat- posteriorly from end of abdomen (Fig. 32); ter species now placed in the subgenus Pacificovelia abdominal ventrite VII (final segment before (Andersen & Weir 2003). The new Cambodian spe- genital capsule) not depressed or concave; cies are easily separated from these two taxa by their paramere with apex rounded (Figs 35, 36);
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tylus shining blackish, strongly contrast- Ecological notes. The specimens were taken from the ing with dull brown coloration of adjacent margins of a quiet, shaded stream pool. frons ����� Microvelia (Microvelia) setifera n. sp. – Body generally lacking long, erect setae Microvelia (Microvelia) penglyi n. sp. (Figs 25, 26); genital segment retracted into Figs 25–31 end of abdomen (Figs 25, 27); abdominal ven- trite VII (final segment before genital capsule) Type material. Holotype, micropterous male, Cam- broadly depressed and concave across its entire bodia, Pursat Prov., Cardamom Mountains, O Sarb width (Fig. 27); paramere with apex pointed Morth stream (= Sap Mat stream), N of Vealveang vil- (Figs 29, 30); tylus dark blackish brown, not lage on road to Pursat, 230 m [760 ft], 12°18'31''N, strongly contrasting with similar coloration on 103°06'36''E, water temp. 27°C, 2 September 2015, frons ������ Microvelia (Microvelia) penglyi n. sp. 08:00–09:30 hrs, CL 6024, D.A. Polhemus (USNM). Paratypes: Cambodia, Kampong Speu Prov.: 2 wing- Microvelia (Picaultia) douglasi Scott less males, Kirirom Plateau, Tasek Stream, cross- Microvelia douglasi Scott, 1874: 448. ing at second bridge on road up to summit, 640 m [2095 ft], 11°19'23''N, 104°04'59''E, water temp. Material examined. Cambodia, Kampong Speu Prov.: 25°C, 4 June 2016, 9:30–10:30 hrs, CL 6047, D.A. 2 winged males, 1 winged female, Sva Slab, CL 6034. Polhemus (BPBM). Koh Kong Prov.: 1 wingless fe- Sihanoukville Prov.: 2 winged males, Thmar Roung, CL male, Cardamom Mountains, O An Dart River, at 6038. highway bridge ~73 km SE of Koh Kong, 27 m [90 Discussion. The genus Picaultia was described by ft], 11°16'38''N, 103°19'47''E, water temp. 26°C, Distant (1913) based on the type species Microv 30 August 2015, 11:45–13:00 hrs, CL 6015, D.A. elia pronotalis from the Seychelles islands. Andersen Polhemus (BPBM); 1 wingless female, Cardamom & Weir (2003) reduced Picaultia to a subgenus of Mountains, Toeuk Noeng River, 9 km SSE of Thmar Microvelia, and placed M. douglasi within it, along Bang village crossroads, 165 m [550 ft], 11°37'33''N, with 18 other species of Microvelia occurring from 103°28'51''E, water temp. 26°C, 31 August 2015, Europe and Africa eastward through Asia to the 14:45–16:00 hrs, CL 6020, D.A. Polhemus (BPBM). Solomon Islands. All of these species possess grasp- Pursat Prov.: 1 wingless male, 12 wingless females, ing combs on both the fore and middle tibiae in the same data as holotype (BPBM, USNM). males, and have highly asymmetrical male genitalia, with the right paramere much larger in size than Description the left. Microvelia douglasi was originally described from Micropterous male (Fig. 25). Length 2.20–2.30 Japan, and as interpreted by Lundblad (1933), who mm ( x ̅ = 2.25, n = 3); width 0.75–0.80 mm undertook a detailed morphological analysis of the ( x ̅ = 0.78, n = 3). species, has a range extending from India eastward Color. Ground color dark blackish-brown, marked through Indochina to Indonesia, the Philippines, with orange–brown on anterior pronotum and outer New Guinea, Australia, Japan, the Mariana Islands portions of connexiva orange–brown; patches of and Samoa. As such, its presence in Cambodia is not short, shining, silvery pubescence present laterally on surprising. pronotum, metanotum, and abdominal tergites II, Ecological notes. Specimens were taken from the III, VI and VII, and connexival sutures. Head dark margins of quiet stream pools. blackish-brown; eyes dark red; antennae uniformly medium brown; rostrum dark yellow, dark brown along medial line, piceous distally. Pronotum dark Microvelia (Picaultia) falcata Zettel, Phauk, blackish-brown, bearing an elongate transverse or- Kheam & Freitag ange–brown patch behind anterior margin, width of Microvelia (Picaultia) falcata Zettel, Phauk, Kheam this patch subequal to width of vertex, patch faintly & Freitag, 2017: 27. divided medially by narrow, diffuse dark brown lon- gitudinal line. Metanotum and abdominal tergites Material examined. Cambodia, Kampong Speu Prov.: 2 dark blackish-brown, connexiva orange–brown on winged males, 1 winged female, nr. Rolak, CL 6041. outer halves and ventrally. Legs with femora yellow- Discussion. This species was recently described from ish brown with distal apices darker brown; tibiae specimens taken in Siem Reap Province. The record dark brown, each with yellowish brown annulation from Rolak village now extends the known range to centrally; tarsal segment I and base of tarsal segment the foothills of the Cardamom Mountains in Kam- II yellowish-brown, remainder of tarsal segment II pong Speu Province. This species appears to be en- dark brown. Ventral surface uniformly dull, dark demic to Cambodia. blackish-brown, except genital segments brown. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Figs 25–26. Microvelia penglyi n. sp., dorsal habitus; specimens from Cambodia, Pursat Prov., Vealveang, CL 6024. 25. Micropterous male. 26. Micropterous female.
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Fig. 27. Microvelia penglyi n. sp., male terminal abdominal ventrite VII, ventral view; specimen from Cambodia, Pursat Prov., Vealveang, CL 6024.
Structural characters. Head — Of moderate length, erect brown setae present along anterior margins of declivant anteriorly, with weak impressed me- all antennal segments, length of these setae on seg- dian line; length 0.35, width 0.55; width of eye/ ment I slightly exceeding width of segment, lengths interocular space, 0.10/0.35. Antennal formula I : of setae on segments II–IV at least 2× the widths of II : III : IV; 0.25 : 0.30 : 0.35 : 0.45. these segments. Pronotum — Long, covering mesonotum, poste- Legs — Fore femur with ventral margin straight, rior lobe coarsely punctate, length/width, 0.30/0.70. not modified, bearing numerous short, erect pale se- Metanotum coarsely punctate, length 0.25. Abdom- tae along entire length; fore tibiae with inner margin inal tergites coarsely punctate, dull, without shining straight, distal grasping comb very short, present as areas; lengths of tergites II–VII, respectively: 0.20: a small, dark, curving spur ventrally at apex, ante- 0.15 : 0.15 : 0.20 : 0.25 : 0.30. rior margin of fore tibia with scattered long, erect, Dorsal surface — Entire dorsum and lateroter- pale setae; middle femur unmodified, bearing a gites covered with fine appressed pale pubescence, fringe of short, erect pale setae along ventral margin intermixed with scattered, short, semi-erect dark intermixed with a few longer setae distally; middle setae; patches of moderately long, recumbent, pos- tibia with posterior margin bearing a row of evenly teriorly-directed silvery setae present along posterior spaced, long, pilose, pale setae of progressively de- margin of transverse yellowish-brown patch on an- creasing length, longest and most basal of these setae terior pronotum, along lateral margins of mesono- with length exceeding 2× width of tibia; hind femur tum, along posterior margins of abdominal tergites unmodified, with numerous short, erect pale setae II and III, sparingly as a narrow transverse band along ventral margin; claws long, slender, gently across central section of abdominal tergite III, as a curving. Proportions of legs as follows: Femur, tibia, large and prominent patch covering most of central tarsal 1, tarsal 2 of fore leg, 0.60 : 0.55 : 0.30; of mid- section of abdominal tergite VI, as small and diffuse dle leg, 0.70 : 0.70 : 0.20 : 0.25; of hind leg, 0.75 : patch centrally on abdominal tergite VII, and along 0.80 : 0.15 : 0.20. all connexival sutures; wing pads present as small, Ventral surface — Pro-, meso-, and metasterna subtriangular transverse flaps arising beneath pos- broadly sulcate along longitudinal midlines; ab- terolateral margins of pronotum; legs and antennae dominal ventrite I oriented vertically, with small thickly clothed with short pale setae, scattered long tumescence centrally. Abdominal venter set with
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Figs 28–31. Microvelia penglyi n. sp., male genitalic structures; specimens from Cambodia, Kampong Speu Prov., Kirirom Plateau, CL 6047. 28. Proctiger. 29. Paramere, lateral view. 30. Paramere, alternate oblique view. 31. Posterior margin of pygophore, showing angulate lateral lobes. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Figs 32–33. Microvelia setifera n. sp., dorsal habitus; specimens from Cambodia, O An Dart River, CL 6015. 32. Micropterous male. 33. Micropterous female. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Figs 34–36. Microvelia setifera n. sp., male genitalic structures; specimen from Cambodia, Koh Kong Prov., O An Dart River, CL 6015. 34. Proctiger. 35. Paramere, oblique view. 36. Paramere, alternate lateral view.
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2 wingless males, 1 wingless female, Cardamom along inner margins of connexiva adjoining lateral Mountains, O Sarb Morth stream (= Sap Mat margins of abdominal tergites II and III, and also stream), N of Vealveang village on road to Pursat, tergites VI and VII; wing pads present as small, sub- 230 m [760 ft], 12°18'31''N, 103°06'36''E, water quadrate transverse flaps arising beneath posterolat- temp. 27°C, 2 September 2015, 08:00–09:30 hrs, eral margins of pronotum; legs and antennae thickly CL 6024, D.A. Polhemus (BPBM). clothed with short pale setae, scattered long erect brown setae present along anterior margins of anten- nal segments I–III, lengths of these setae exceeding Description 2× the width of the segments on which they occur. Micropterous male (Fig. 32). Length 2.50–2.60 mm Legs — Fore femur with ventral margin straight, ( x ̅ = 2.53, n =3); width 0.80–0.90 mm ( x ̅ = 8.33, not modified, bearing a row of ~10 very long, pale, n = 3). erect, evenly spaced setae; fore tibiae with inner mar- Color. Ground color dark blackish-brown, marked gin straight, anterior and posterior margins bearing with orange–brown on anterior pronotum and scattered long, erect dark setae, distal grasping comb outer portions of connexiva; patches of short, shin- very short, length about one-fourth that of tibia; ing, silvery pubescence present laterally on prono- middle femur unmodified, anterior margin bear- tum, metanotum, and abdominal tergites II and III ing scattered long, erect dark setae, posterior mar- and anterior and posterior portions of connexiva. gin bearing an evenly spaced row of ~12 very long, Head dull medium brown; tylus shining dark black- pale, erect setae; middle tibia with anterior margin ish brown; eyes dark red; antennae medium brown, bearing scattered long, erect dark setae, posterior with distal half of segment I and basal half of seg- margin bearing a row of evenly spaced, long pilose ment II lighter brown; rostrum dark yellow, dark dark setae of progressively decreasing length, longest brown along medial line, piceous distally. Pronotum and most basal of these setae with length exceeding dark blackish-brown, bearing an elongate trans- 2× width of tibia; hind femur and tibia unmodified, verse orange–brown patch behind anterior margin, both bearing numerous long, erect dark setae along width of this patch subequal to width of vertex. both anterior and posterior margins; claws very long, Metanotum and abdominal tergites dark blackish- slender, gently curving. Connexiva evenly converg- brown, connexiva orange–brown on outer halves ing posteriorly, bowed gently outward. Proportions and ventrally. Legs with femora pale yellowish on of legs as follows: femur, tibia, tarsal 1, tarsal 2 of fore basal halves, dark brown on distal halves; tibiae dark leg, 0.70 : 0.60 : 0.30; of middle leg, 0.85 : 0.75 : brown, each with broad dark yellowish annulation 0.15 : 0.25; of hind leg, 0.90 : 1.10 : 0.20 : 0.25. centrally; tarsal segment I and base of tarsal segment Ventral surface — Ventral thorax and abdomen II yellowish-brown, remainder of tarsal segment II with numerous coarse punctations; prosternum dark brown. Ventral surface uniformly dull, dark deeply sulcate along longitudinal midline between blackish-brown, longitudinal midline of abdomen acetabulae, mesosternum weakly sulcate along lon- narrowly shining; genital segments brown, shining gitudinal midline, metasternum broadly domed, and darker basally. lacking longitudinal sulcus; abdominal ventrite I Structural characters. Head — Of moderate length, oriented vertically, with small, posteriorly projecting declivant anteriorly, with weak impressed median tumescence centrally. Abdominal venter set with nu- line, row of 4 coarse punctations present along each merous long, erect, pale setae; abdominal ventrites lateral margin of vertex adjacent to inner margins of I–VII unmodified, lacking carinae, tumescences or eyes; length 0.45, width 0.60; width of eye/interocu- depressions. lar space, 0.20/0.30. Antennal formula I : II : III : Male genitalia — Male genital segment protuding IV; 0.40 : 0.25 : 0.40 : 0.60. (Fig. 32); proctiger elongate, parallel sided, lacking Pronotum — Long, covering mesonotum, coarse- lateral lobes, apex rounded (Fig. 34). Paramere slen- ly punctate, length/width, 0.45/0.75. Metanotum der, gently curving, apex rounded, slightly bulbous coarsely punctate, length 0.18. Abdominal ter- and hooked (Figs 35, 36). gites II–V dull, coarsely punctate, tergites VI–VIII Micropterous female (Fig. 33). Similar to micropter- smooth, shining; lengths of tergites II–VIII, respec- ous male in general structure and color, with fol- tively: 0.18: 0.18 : 0.15 : 0.15 : 0.15 : 0.25 : 0.20. lowing exceptions: length 2.80–2.95 mm ( x = 2.88, Dorsal surface — Entire dorsum and lateroter- ̅ n = 5); width 0.90–1.10 mm ( x ̅ = 1.00, n = 5); mesono- gites covered with fine appressed pale pubescence, tum and abdominal tergites II and III bowed broadly intermixed with numerous, long, erect dark setae; and gently upward, abdominal tergite IV angled patches of moderately long, recumbent, silvery setae downward, abdominal tergites V–VIII horizontal, present laterally within transverse yellowish-brown deeply recessed below connexiva; coarse punctations patch on anterior pronotum, along lateral margins present on abdominal tergites II and III only; con- of mesonotum and abdominal tergites II and III, and nexiva weakly posteriorly convergent adjacent to Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Fig. 37. Microvelia bokor n. sp., micropterous male, dorsal habitus; specimen from Cambodia, Kampot Prov., stream on Bokor Plateau, CL 6035. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Fig. 38. Microvelia bokor n. sp., male foreleg showing distal expansion of fore tibia; specimen from Cambodia, Kampot Prov., stream on Bokor Plateau, CL 6035. eye/interocular space, 0.10/0.25. Antennal formula acetabulae, mesosternum weakly sulcate along lon- I : II : III : IV; 0.20 : 0.20 : 0.30 : 0.40. gitudinal midline, metasternum broadly domed, Pronotum — Long, covering mesonotum, lacking longitudinal sulcus; abdominal ventrite I posterior lobe coarsely punctate, length/width, oriented vertically. Abdominal venter set with nu- 0.25/0.65. Metanotum length 0.15. Abdominal merous very short, pale, recumbent setae; abdominal tergites dull, without shining areas; lengths of ter- ventrites I–VII unmodified, lacking carinae, tumes- gites II–VII, respectively: 0.15: 0.10 : 0.10 : 0.12: cences or depressions. 0.15 : 0.25. Male genitalia — Male genital segment retracted Dorsal surface — Entire dorsum and laterotergites into abdomen; proctiger slightly expanded later- covered with short, appressed golden pubescence, ally, lacking basal lobes, distal section evenly taper- mostly lacking erect setae; patches of moderately ing with apex slightly produced (Fig. 39); paramere long, recumbent, silvery setae present along laterally slender, elongate, gently curving, apex acute, weakly on pronotum behind each eye, along lateral margins multidentate in certain views (Figs 40, 41); pygo- of mesonotum, narrowly along posterior margin of phore unmodified. abdominal tergite III, as prominent paired (1 + 1) Micropterous female. Similar to micropterous male in patches laterally on tergites IV and V, as small paired general structure and color, including distribution of (1 + 1) patches posterolaterally on tergite VI, and silvery hair patches dorsally, with following excep- along all connexival sutures; wing pads present as tions: connexiva more broadly and evenly bowed small, subtriangular transverse flaps arising beneath outward, convergent posteriorly but leaving abdom- posterolateral margins of pronotum; legs and anten- inal tergite VII completely exposed, posterolateral nae thickly clothed with short pale setae, scattered apices of segment VII not produced, lacking setal long erect brown setae present along anterior mar- tufts, connexival margins of even width throughout. gins of all antennal segments, lengths of these setae Abdominal venter unmodified, set with numerous at least 2× the widths of the segments on which they very short pale setae; gonocoxae tightly appressed arise. along posterior margins, lying at 45° angle. Legs un- Legs — Posterior margins for fore femur, middle modified, fore tibia lacking grasping comb. femur and middle tibia bearing a few longer, erect Macropterous male. Unknown. pale setae; fore femur slightly bowed, expanded and Macropterous female. Unknown. ventrally flattened on distal one-half to accomodate Etymology. The name “bokor” is a noun in apposi- grasping comb (Fig. 38); middle and hind femora, tion and refers to the Bokor Plateau of southwestern tibiae and tarsi unmodified; claws long, slender, gen- Cambodia, the type locality for this new species. tly curving. Proportions of legs as follows: Femur, Discussion. Microvelia bokor is similar to M. peng tibia, tarsal 1, tarsal 2 of fore leg, 050 : 0.45 : 0.22; lyi in many respects, but much smaller in size (body of middle leg, 0.55 : 0.50 : 0.10 : 0.15; of hind leg, length 1.70–1.85 mm, vs 2.20–2.75 mm in M. peng 0.55 : 0.70 : 0.10 : 0.15. lyi), and with a different pattern of silvery setifera- Ventral surface — Venter of thorax and abdo- tion on the dorsal abdomen (compare Figs 25, 26 to men with numerous coarse punctations; prosternum Fig. 37). In addition, the male fore tibia is distinctly deeply sulcate along longitudinal midline between expanded on its distal half, with the grasping comb Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Figs 39–41. Microvelia bokor n. sp., male genitalic structures; specimen from Cambodia, Kampot Prov., stream on Bokor Plateau, CL 6035. 39. Proctiger. 40. Paramere, lateral view. 41. Paramere, alternate oblique view.
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Material examined. Cambodia, Koh Kong Prov.: 1 1 wingless male, nr. Rolak, CL 6041; 13 wingless males, 13 winged female, nr. Thmar Bang, CL 6017; 1 winged wingless females, nr. Krong Chbar Mon, CL 6045. female, 1 wingless male, 2 wingless females, Toeuk Noeng, Discussion. This species may be easily recognized CL 6020. Pursat Prov.: 1 winged male, 3 wingless females, among the Veliidae of Cambodia by the presence nr. Vealveang, CL 6024. Sihanoukville Prov.: 6 wingless of finely bifurcating swimming plumes on both the males, 8 wingless females, nr. Pich Nil, CL 6039. middle and hind tarsi, and by the strongly alternat- Discussion. This species superficially resemblesR. ing pale and dark annulations on the legs (Fig. 51). sumatrensis, bearing stout spines basoventrally on Ecological notes. Originally described from Thailand the male middle femur (Fig. 44), but the paramere (Andersen 2000), and until now known only from is shorter, more robust, and not hooked at apex as that country, Tetraripis zetteli was found as scat- in R. sumatrensis. The Cambodian specimens have tered colonies on various streams in the Cardamom been compared to material from the Central High- Mountain premontane foreland in Kampong Speu lands of Vietnam (CL 4297), and match in all par- Province. In all cases the insects were quite localized ticulars, including the spination of the male hind leg along any given stream, aggregating in dark pock- (Fig. 45) and the stout form of the male paramere ets beneath undercut banks. Repeated splashing and when viewed laterally (Figs 48, 49). scooping of such pockets could sometimes result in Although Figs 7 and 8 in Zettel (2000) are suf- capture of a large number of specimens, indicating ficient to illustrate the difference between the stout that adults may be sheltering above the water line. male paramere in R. inexpectata and the more slen- Tetraripis zetteli is almost undoubtedly nocturnal der, hooked paramere in R. sumatrensis when viewed in the adult stage, although immatures were found laterally, they do not convey the true degree of con- skating during daylight hours amid low emergent cavity at the truncate apex of the paramere in R. vegetation swept by the current along the margins of inexpectata, which is evident in specimens from both a channelized lowland stream near Kampong Speu, Vietnam and Cambodia (Figs 48–50). In addition, in a manner reminiscent of Rhagovelia. there is a certain degree of intraspecific variation in the shape of the paramere among local populations in the Cardamom Mountains (Figs 48, 49). Subfamily Veliinae Brullé
Rhagovelia tebakang J. Polhemus & D. Polhemus Angilia (Adriennella) bispinosa Andersen Figs 43, 46, 47 Angilia (Adriennella) bispinosa Andersen, 1982: 345. Rhagovelia tebakang J. Polhemus & D. Polhemus, 1988: 186. Material examined. Cambodia, Kampong Speu Prov.: 1 male, 2 immatures, Kirirom, CL 6047. Material examined. Cambodia, Koh Kong Prov.: 4 Ecological notes. This species was originally described winged males, 3 wingless males, 3 wingless females, Koh from Thailand, and subsequently recorded from Kong–Pursat, CL 6023. Vietnam and Burma (Polhemus & Polhemus 2003), Discussion. This species was previously known only therefore its presence in Cambodia is not surprising. from Borneo. The Cambodian specimens, however, A single male specimen was taken from a stream on possess all the key characters of this species, includ- the flanks of the Kirirom Plateau, flowing through ing the expanded juga, presence of spinules on the native pine forest but shaded by a narrow band of male hind trochanter (Fig. 46), and an elongate, broadleaf riparian gallery forest. The single adult distally hooked male paramere (Fig. 47). They taken was found in a dark recess beneath a projecting therefore appear to represent the first record of this sandstone boulder along the stream margin. Several species from the Southeast Asian mainland. As with immatures were also taken from dark pockets along Rhagadotarsus borneensis and Hydrometra longicapitus vertical banks of red lateritic alluvium. The imma- discussed previously, this seems to represent another tures of this species have black bodies with strongly instance of a disjunct Sundaland distribution within contrasting white spots. the aquatic Heteroptera fauna of Cambodia.
Tetraripis zetteli Andersen Infraorder Nepomorpha Fig. 51 Family Belostomatidae Leach Tetraripis zetteli Andersen, 2000: 187. Diplonychus rusticus (Fabricius) Material examined. Cambodia, Kampong Speu Prov.: 7 Nepa rustica Fabricius, 1781: 333. wingless males, 3 wingless females, nr. Chambok, CL 6033; Diplonychus rusticus: Laporte, 1833: 18.
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Figs 42–43. Rhagovelia species, dorsal habitus. 42. R. inexpectata, apterous female, specimen from Cambodia, Sihanoukville Prov., stream below Kirirom hydroelectric plant, CL 6039. 43. R. tebakang, apterous female, speci- men from Cambodia, Koh Kong Prov., stream on Koh Kong to Pursat road, CL 6023.
Appasus marginicollis Dufour, 1863: 393; syn. by Family Helotrephidae Esaki & China Distant, 1906: 36. Diplonychus indicus Ventaktesan & Rao, 1980: 299; syn. by J. Polhemus, 1995: 651. Idiotrephes cf. shepardi Papácek & Zettel Idiotrephes shepardi Papacek & Zettel, 2000: Material examined. Cambodia, Kampong Speu Prov.: 208. 1 immature, nr. Krong Chbar Mon, CL 6045. Ecological notes. A single immature of this species was Material examined. Cambodia, Koh Kong Prov.: taken in a wetland along Hwy 4 near Kampong Speu. 1 brachypterous female, Toeuk Noeng, CL 6020. This is the only member of the genus present in Indo- Discussion. Only a single female of this species is china, and the species identification is not in doubt. at hand, rendering its determination as I. shepardi Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Figs 44–46. Rhagovelia species, legs. 44. R. inexpectata, male left middle femur, dorsal view, showing stout spines ventrally near base; specimen from Cambodia, Sihanoukville Prov., stream below Kirirom hydroelectric plant, CL 6039. 45. R. inexpectata, male left hind leg, dorsal view; specimen from Cambodia, Sihanoukville Prov., stream below Kirirom hydroelectric plant, CL 6039. 46. R. tebakang, male left hind leg, dorsal view, note spinules on hind trochanter; specimen from Cambodia, Koh Kong Prov., stream on Koh Kong toDownloaded Pursat road, from Brill.com10/07/2021CL 6023. 06:11:13PM via free access
Figs 47–50. Rhagovelia species, male right paramere, lateral view. 47. R. tebakang, specimen from Cambodia, Koh Kong Prov., stream on Koh Kong to Pursat road, CL 6023. 48–50. R. inexpectata, showing geographic variation in shape of paramere apex. 48. Specimen from Cambodia, O Sarb Morth stream near Vealveang, CL 6024. 49. Specimen from Cambodia, Touk Neong River, CL 6020. 50. Specimen from Vietnam, Quang Ngai Prov., NE of Kontum, CL 4297.
provisional. The shape of the subgenital plate con- Family Micronectidae Jackzewski forms to the illustration for I. shepardi given in Papacek & Zettel 2000, but definitive confirmation Genus Micronecta Kirkaldy will require collection of male specimens. Discussion. The three species listed below can be de- finitively identified from the CambodianMicronecta Hydrotrephes maculatus Papácek & Kovac material at hand. In addition, one further series con- Hydrotrephes maculatus Papacek & Kovac, 2001: sisting of one male and two females from the Sva Slab 317. River (CL 6034), in Kampong Speu Province, pos- sesses male genitalic structures that do not match any Material examined. Cambodia, Kampong Speu Prov.: 1 figure in Lundblad (1933), Nieser & Chen (1999), male, 1 female, Sva Slab, CL 6034. or Nieser (1999, 2002). This taxon has a tan ground Discussion. The male genitalia of the Cambodian color with dark lateral patches along costa, but lacks male specimen at hand agree in all respects with the other significant markings. The right paramere is illustrations provided for this species by Papacek & very long and slender, similar to M. dentifera Nie- Kovac (2001). ser, while the left paramere is distinctively shaped. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Fig. 51. Tetraripis zetteli, apterous male, dorsal habitus; specimen from Cambodia, Kampong Speu Prov., stream near Krong Chbar Mon, CL 6045.
Because the author is not comprehensively familiar Material examined. Cambodia, Kampong Speu Prov.: with all Micronecta species occurring in Southeast 1 macropterous female, Sva Slab, CL 6034. Asia, description of this potential new species is de- Ecological notes. This species was taken from water- ferred pending further analysis. filled potholes in diabase bedrock along the Sva Slab River near Chambok. At this locality it co-occurred Micronecta guttatostriata Lundblad in the same pools along with Micronecta guttatostri Micronecta guttatostriata Lundblad, 1933: 101. ata Lundblad.
Material examined. Cambodia, Kampong Speu Prov.: Synaptonecta issa (Distant) 1 macropterous female, Sva Slab, CL 6034. Synaptonecta issa (Distant, 1910): 350. Ecological notes. This species was taken from water- filled potholes in diabase bedrock along the Sva Slab River near Chambok. At this locality it co-occurred Material examined. Cambodia, Kampong Speu Prov.: in the same pools along with Micronecta quadris 1 macropterous male, Sva Slab, CL 6034. Kampot trigata Breddin. Prov.: 1 brachypterous male, Bokor Plateau, CL 6036. Discussion. The macropterous male specimen from Micronecta polhemusi Nieser the Sva Slab River has fully developed hind wings, Micronecta polhemusi Nieser, 2000: 283. and a dark color pattern consisting of heavy brown mottling and small punctate dark brown dots. Material examined. Cambodia, Koh Kong Prov.: 4 mac- By contrast, the hemelytra of the brachypterous ropterous females, Toeuk Noeng, CL 6020. specimen from the Bokor Plateau has a much paler Ecological notes. This species was taken from water- color pattern, similar to that shown in Fig. 18.2 of filled potholes in sandstone bedrock along the Touek Tran et al. (2015). The differences in coloration were Noeng River in the central Cardamom Mountains. pronounced enough that at first it appeared two sepa- rate species might be at hand, but the male genitalic Micronecta quadristrigata Breddin structures in both specimens conform to those of Micronecta quadristrigata Breddin, 1905: 57. S. issa. Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Ecological notes. The Bokor Plateau specimen was Family Nepidae Latreille taken from a deep, cold pond surrounded by upland rain forest on the nearly level summit of the moun- Subfamily Ranatrinae Douglas & Scott tain (Fig. 10). Cercotmetus asiaticus Amyot & Serville Cercotmetus asiaticus Amyot et Serville, 1843: 441. Family Naucoridae Leach Discussion. A previous record of this species from Heleocoris malayensis D. Polhemus & J. Polhemus southwestern Cambodia was provided by Polhemus Heleocoris malayensis D. Polhemus & J. Polhemus, & Polhemus (2013). It was not recollected during 2013: 676. the present surveys.
Material examined. Cambodia, Kampong Speu Prov.: 1 Cercotmetus brevipes brevipes Montandon female, Sva Slab, CL 6034. Cercotmetus brevipes brevipes Montandon, 1909: 65. Discussion. A single female that has been assigned to this species was taken along the margins of Material examined. Cambodia, Koh Kong Prov.: 1 male, Sva Slab River where it flowed across exposures Koh Kong–Pursat, CL 6023. of diabase bedrock northeast of Chambok village. Ecological notes. A single specimen was taken from This specimen has been identified asH. malayensis the shaded margins of a still pond formed upstream based on the shape of the subgenital plate, the of a road crossing, amid submerged grasses and twigs. apex of which is shallowly concave rather than possessing a V-shaped incision (see discussion in Ranatra thai Lansbury Polhemus & Polhemus 2013). Because the range of Ranatra thai Lansbury, 1972: 334. this species and that of H. ovatus Montandon over- lap in southern Indochina, definitive confirmation of this species in Cambodia must await collection Material examined. Cambodia, Kampong Speu Prov.: of male specimens. 2 males, 2 females, Sva Slab, CL 6034; 1 male, Kirirom, CL 6048. Kampot Prov.: 7 males, 8 females, Bokor Pla- teau, CL 6036. Sihanoukville Prov.: 1 male, Thmar Roung, Naucoris scutellaris Stål CL 6038. Naucoris scutellaris Stål, 1860: 266. Ecological notes. This species has been recorded Thurselinus greeni Distant, 1904: 33; syn. by from the Malay Peninsula northward into Vietnam, Lundblad, 1933: 65. Thailand, and now Cambodia (Tran & Polhemus Naucoris rhizomatus J. Polhemus, 1984: 157; syn. by 2012). Several good series were taken from stand- Polhemus & Polhemus, 2013: 669. ing water habitats at elevations ranging from 30 to 1000 m. Material examined. Cambodia, Kampong Chhnang Prov.: 1 female, Bori Bo, CL 6025. Kampong Speu Prov.: Ranatra cf. gracilis Dallas 2 males, 1 female, Sva Slab, CL 6034; 1 male, 2 females, Ranatra gracilis Dallas, 1850: 10. nr. Rolak, CL 6043. Koh Kong Prov.: 1 male, 1 female, O An Dart, CL 6015; 1 male, Koh Kong–Pursat, CL 6023. Material examined. Cambodia, Kampong Chhnang Pursat Prov.: 1 male, nr. Vealveang, CL 6024. Prov.: 2 immatures, Bori Bo, CL 6025. Koh Kong Prov.: Discussion. The genus Naucoris was last comprehen- 3 immatures, O An Dart, CL 6015; 1 immature, Toeuk sively reviewed for continental Southeast Asia by Pol- Noeng, CL 6020. Pursat Prov.: 2 immatures, nr. Vealveang, hemus & Polhemus (2014). This work provided il- CL 6024. lustrations of the male genitalia for all species known Discussion. All of the immature specimens listed from the region. One other Southeast Asian Naucoris above fall into the gracilis species group based on the species, N. sumatrensis Fieber, is known only from presence of a prominent, conical tubercle on the head the female holotype, apparently from Sumatra, with vertex and a short respiratory siphon. In all of these photographic and line drawing illustrations of that specimens the presence of a median longitudinal ca- type specimen provided by Zettel (2011). rina extending the entire length of the prosternum Ecological notes. Naucoris scutellaris is locally com- indicates that these specimens most likely represent mon in lowland streams draining from the Carda- Ranatra gracilis Dallas (see key in Polhemus & mom Mountains. Individuals occur along the mar- Polhemus 2012), but this cannot be verified until gins of pools, often in areas with submerged leaf adult specimens are obtained from these localities. litter. The fact that only immatures were obtained at Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Figs 52–53. Nychia sappho, macropterous male, specimen from Cambodia, Kampong Speu Prov., Thom River near Rolak Kong Cheung village, CL 6042. Fig. 52. Dorsal habitus. Fig. 53. Ventral habitus. morphological variation that might warrent further east of Rolak village (CL 6043, Fig. 8); and a deep, taxonomic subdivision, but declined to undertake cold, open pond on the summit of the Bokor Plateau this given the uncertain state of species concepts (CL 6036, Fig. 10). in the genus. Given the equivocal morphological evidence, it seems likely that a molecular systematic Enithares ciliata (Fabricius) study will be necessary to properly constrain species Enithares ciliata Fabricius, 1798: 524. concepts within this genus. Ecological notes. Nychia sappho was taken at multiple Material examined. Cambodia, Kampong Speu Prov.: 1 sites in southwestern Cambodia, at altitudes ranging male, nr. Chambok, CL 6033. from 95 to 985 m. The unifying characteristic of all Discussion. Originally described from India, this is the collecting sites was the presence of standing or a widespread species in tropical Asia, with a range very slowly flowing waters. The habitats from which extending from southward to Ceylon and eastward N. sappho was collected included unshaded, water- through Thailand, Vietnam and Malaysia to Su- filled rock holes in a diabase bedrock exposure along matra (Lansbury 1968). Its collection in Cambodia the margins of the Sva Slab River (CL 6034); heav- logically fills a gap in the previously documented ily shaded overflow channels in gravel parallel to the distribution. main channel of the Thom River (CL 6042); slowly Ecological notes. The single individual taken during flowing, partially shaded pools along a rocky stream the current surveys was netted from the margins of Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Fig. 54. Nychia sappho, intact male genitalia, left lateral view, with major structures labeled; specimen from Cam- bodia, Kampong Speu Prov., stream near Chambok village, CL 6034.
Figs 55–56. Nychia sappho, brachypterous forms, dorsal habitus. 55. Female specimen from Cambodia, Kampong Speu Prov., Thom River near Rolak Kong Cheung village, CL 6042. 56. Female specimen from Indonesia, New Guinea, Papua Prov, nr. Timika, CL 7053.
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Figs 57–58. Nychia sappho, brachypterous forms, lateral view of head and thorax showing form of antennae and lateral pronotal fovea. 57. Female specimen from Cambodia, Kampong Speu Prov., Thom River near Rolak Kong Cheung village, CL 6042. 58. Female specimen from Indonesia, New Guinea, Papua Prov, nr. Timika, CL 7053.
an unshaded pool about 1 m deep along a stream Discussion flowing in a sandy bed through dry semi-deciduous Based on the current surveys, coupled with limited mesic forest with stands of riparian bamboo. previous literature records, a total of 11 families, 38 genera, and 78 species of aquatic Heteroptera Enithares mandalayensis Distant are known to occur in Cambodia, with 68 species Enithares mandalayensis Distant, 1910: 331. known from the area covered by the present study. Of these, six species are putatively endemic to Cam- Material examined. Cambodia, Kampot Prov.: 1 male, bodia, although such a conclusion is provisional, Bokor Plateau, CL 6036. given that freshwater systems across all of Cambo- Discussion. Originally described from Burma, E. dia, much less Indochina as a whole, have not been mandalayensis has also been previously recorded comprehensively surveyed. All of the potentially en- from Thailand, Vietnam, and peninsular Malaysia demic species so far recorded from the country have (Lansbury 1968). As such, its presence in Cambodia come from the Cardamom Mountains and adjacent is not surprising. uplands. Ecological notes. A single individual was taken from Based on current knowledge, Cambodia overall a deep, cool pond on the top of the Bokor Plateau does not have a particularly high degree of faunal en- (Fig. 10), in company with numerous individuals demism in most groups of organisms for which data of Anisops nigrolineatus. The specimen was taken is available. For Odonata, another freshwater insect by scooping the net very deeply into the pond, in- group that is comparatively well surveyed in Cambo- dicating that this species my occupy deeper waters dia, 191 species are currently known, with none en- nearer the benthos, in comparison to the Anisops demic (Kosterin et al. 2012; Kosterin 2014; Kosterin which were abundant in the middle of the water & Chartier 2014; Kosterin pers. comm.). Among column. reptiles and amphibians, 245 species are currently Downloaded from Brill.com10/07/2021 06:11:13PM via free access
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Appendix 1: Revised checklist of aquatic and semi-aquatic Heteroptera of Cambodia
Based on records from Zettel et al. (2017) and the although the occurrence of this taxon may be likely present work. New Cambodia country taxon records in Cambodia, and it has been erroneously listed from from the present paper are marked with an asterisk the country in the past, there is no currently con- (*). New Cambodian provincial records from the firmed Cambodian record. present paper are marked with a hash (#). The checklist does not include Halovelia ber grothi Esaki, because as noted by Zettel et al. (2017),
Taxon Provincial distribution
INFRAORDER GERROMOPRHA Family Gerridae Amemboa cambodiana n. sp.* Kampot, Koh Kong, Kampong Chhnang# Amemboa cristata J. Polhemus & Andersen, 1984* Kampong Chhnang# Amemboa javanica Lundblad, 1933* Kampong Speu# Amemboa speciosa J. Polhemus & Andersen, 1984* Pursat# Cylindrostethus costalis Schmidt, 1915 Koh Kong, Siem Reap, Kampong Speu Gnomobates kuiterti Hungerford & Matsuda, 1958* Kampot#, Kampong Speu#, Sihanoukville# Lathriobates johorensis (J. Polhemus & D. Polhemus, 1995)* Kampong Speu#, Sihanoukville# Limnogonus fossarum fossarum (Fabricius, 1775) Phnom Penh, Kampong Speu#, Koh Kong# Limnogonus hungerfordi Andersen, 1975* Koh Kong# Limnogonus nitidus (Mayr, 1865) Phnom Penh, Kampong Speu, Pursat#, Sihanoukville# Limnometra matsudai (Miyamoto, 1967) Siem Reap, Sihanoukville#, Pursat#, Kampot#, Koh Kong# Limnometra octopunctata Hungerford, 1955* Sihanoukville# Metrocoris nigrofascioides Chen & Nieser, 1993 Koh Kong, Kampong Speu Metrocoris tenuicornis Esaki, 1926 Koh Kong Naboandelus signatus Distant, 1910* Kampong Chhnang# Neogerris parvulus (Stål, 1860) Koh Kong, Phnom Penh, Kampong Speu, Kampot# Onychotrechus esaki Andersen, 1980* Kampot#, Koh Kong# Ptilomera hylactor Breddin, 1903 Koh Kong, Pursat# Ptilomera tigrina Uhler, 1860 Koh Kong, Kampong Speu, Sihanoukville# Rhagadotarsus borneensis J. Polhemus & Karunaratne, 1993* Kampong Speu#, Pursat# Rhagadotarsus kraepelini Breddin, 1905* Kampong Speu# Rheumatogonus vietnamensis Zettel & Chen, 1996 Siem Reap, Kampong Speu, Sihanoukville# Ventidius (Ventidioides) karen Lansbury, 1988* Kampong Speu#, Sihanoukville# Ventidius (Ventidius) distanti Paiva, 1918 Kampong Speu, Koh Kong#, Kampong Chhnang#, Pursat#, Sihanoukville# Family Hebridae Hebrus nieseri Zettel, 2004* Pursat# Hebrus cf. polysetosus Zettel, 2004* Kampong Speu# Hyrcanus varicolor Andersen, 1981 Siem Reap Timasius miyamotoi Andersen, 1981 Siem Reap Family Hydrometridae Hydrometra annamana Hungerford & Evans, 1934* Kampong Speu# Hydrometra gilloglyi J. Polhemus & D. Polhemus, 1995 Kampong Speu, Pursat#, Koh Kong# Hydrometra greeni Kirkaldy, 1898 Koh Kong, Kampong Speu Hydrometra jackzewskii Lundblad, 1933* Kampong Speu# Hydrometra longicapitis Torre-Bueno, 1927 Siem Reap, Kampong Speu#, Pursat# Downloaded from Brill.com10/07/2021 06:11:13PM via free access
Taxon Provincial distribution
Hydrometra orientalis Lundblad, 1933 Koh Kong, Phnom Penh, Kampong Speu, Kampot# Family Mesoveliidae Mesovelia horvathi Lundblad, 1933* Kampong Speu#, Koh Kong#, Kampong Chhnang#, Pursat# Mesovelia vittigera Horváth, 1895 Phnom Penh, Pursat# Family Veliidae Angilia bispinosa Andersen, 1982* Kampong Speu# Microvelia (Microvelia) bokor n. sp.* Kampot# Microvelia (Microvelia) kohkong n. sp.* Koh Kong# Microvelia (Microvelia) penglyi n. sp.* Kampong Speu#, Koh Kong#, Pursat# Microvelia (Microvelia) setifera n. sp.* Kampong Chhnang#, Pursat#, Koh Kong# Microvelia (Picaultia) douglasi Scott, 1874 Kampong Speu#, Sihanoukville# Microvelia (Picaultia) falcata Zettel et al., 2017 Siem Reap Perittopus asiaticus Zettel, 2001* Kampot# Rhagovelia inexpectata Zettel, 2000 Koh Kong, Kampong Speu, Pursat#, Sihanoukville# Rhagovelia tebakang J. Polhemus & D. Polhemus, 1988* Koh Kong# Strongylovelia albopicta Zettel & Tran, 2004* Kampong Speu#, Koh Kong# Tetraripis zetteli Andersen, 2000* Kampong Speu# INFRAORDER NEPOMORPHA Family Belostomatidae Lethocerus indicus (Lepeletier & Serville, 1825) No precise provincial records Diplonychus rusticus (Fabricius, 1781) Phnom Penh, Kampong Speu# Family Helotrephidae Hydrotrephes maculatus Papácek & Kovac, 2001* Kampong Speu# Idiotrephes asiaticus Papácek & Zettel, 2000 Siem Reap Idiotrephes cf. shepardi Papácek & Zettel, 2000* Koh Kong# Family Micronectidae Micronecta drepani Nieser, 1999 Siem Reap Micronecta guttatostriata Lundblad, 1933* Kampong Speu# Microneca lemnae Nieser, 1999 Siem Reap Micronecta ludibunda Breddin, 1905 Siem Reap Micronecta polhemusi Nieser, 2000* Koh Kong# Micronecta quadristrigata Breddin, 1905 Phnom Penh, Kampong Speu# Micronecta scutellaris (Stål, 1858) Phnom Penh Synaptonecta issa (Distant, 1910)* Kampong Speu#, Koh Kong# Family Naucoridae Heleocoris bengalensis montandoni Lundblad, 1933 Koh Kong, Siem Reap Heleocoris malayensis D. Polhemus & J. Polhemus 2013* Kampong Speu#, Naucoris scutellaris Stål, 1860 Kampong Chhnang#, Koh Kong, Phnom Penh, Kampong Speu# Family Nepidae Cercotmetus asiaticus Amyot & Serville, 1843 Koh Kong Cercotmetus brevipes brevipes Montandon, 1909* Koh Kong# Cercotmetus compositus Montandon, 1903 Kampong Speu Laccotrephes cf. pfeiferiae Ferrari, 1888 Phnom Penh Ranatra cardamomensis Zettel et al. 2017 Koh Kong Ranatra cf. gracilis Dallas, 1850* Kampong Chhnang#, Pursat#, Koh Kong# Ranatra parmata Mayr, 1865 Koh Kong, Siem Reap Ranatra thai Lansbury, 1972 Phnom Penh, Kampong Speu, Kampot#, Sihanoukville# Ranatra varipes Stål, 1861 Phnom Penh, Kampong Speu, Koh Kong Family Notonectidae Anisops breddini Kirkaldy, 1901 Phnom Penh Anisops nigrolineatus Lundblad, 1933* Kampong Speu#, Kampot# Enithares mandalayensis Distant, 1910 Kampong Speu, Kampot# Enithares ciliata (Fabricius, 1758)* Kampong Speu# Downloaded# from Brill.com10/07/2021# 06:11:13PM Nychia sappho Kirkaldy, 1901* Kampong Speu , Kampot via free access
Appendix 2: Collecting site data for Vietnamese localities referenced in the text
CL 4297 VIETNAM, Quang Ngai Prov., small rocky stream crossing road on E. side of Via Lac Pass, 109 km NE of Kontum on Hwy 24, 305 m, 14°46'27''N, 108°32'18''E, water temp. 26.5°C, 17 March 2001, D.A. Polhemus, J.T. Polhemus and P. Nguyen. CL 4374 VIETNAM, Bac Thai Prov., Cuc Bo stream, 37 km S. of Bac Kan off Hwy 3, 70 m, 21°52'48''N, 105°45'18''E, water temp. 21°C, 24 March 2000, J.T. and D.A. Polhemus. CL 4384 VIETNAM, Nghê An Prov., Kem waterfall, Pu Mat Nature Reserve, S. of Con Cuông, 400 m, 18°58'12''N, 104°48'03''E, water temp. 22°C, 1 April 2000, J.T. Polhemus and P. Nguyen. CL 4387 VIETNAM, Nghê An Prov., Choang River at ford near border of Pu Mat Nature Reserve, S. of Con Cuông, 160 m, 18°59'07''N, 104°50'32''E, water temp. 25.5°C, 2 April 2000, J.T. Polhemus and P. Nguyen.
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