Belgian Journal of Entomology 75: 1–13 (2018) ISSN: 2295-0214 www.srbe-kbve.be
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Belgian Journal of Entomology
First records of Eulophidae from Cambodia (Hymenoptera: Chalcidoidea)
1 2 3 Zoya YEFREMOVA , Ekaterina YEGORENKOVA & Wouter DEKONINCK
1 School of Zoology, The Steinhardt Museum of Natural History, Tel Aviv University, Tel Aviv 69978, Israel. E-mail: [email protected] 2 Department of Geography and Ecology, Ulyanovsk State Pedagogical University, Ulyanovsk, 432700, Russia. E-mail: [email protected] 3 Royal Belgian Institute of Natural Sciences, Brussels, B-1000 Belgium E-mail: [email protected]
Published: Brussels, August 29, 2018 Citation: YEFREMOVA Z., YEGORENKOVA E. & DEKONINCK W. - First records of Eulophidae from Cambodia (Hymenoptera: Chalcidoidea). Belgian Journal of Entomology, 75: 1–13.
ISSN: 1374-5514 (Print Edition) ISSN: 2295-0214 (Online Edition)
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Front cover: Neotrichoporoides beonus, female, lateral view.
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Belgian Journal of Entomology, 75: 1–13 (2018)
First records of Eulophidae from Cambodia (Hymenoptera: Chalcidoidea)
1 2 3 Zoya YEFREMOVA , Ekaterina YEGORENKOVA & Wouter DEKONINCK
1 School of Zoology, The Steinhardt Museum of Natural History, Tel Aviv University, Tel Aviv 69978, Israel. E-mail: [email protected] 2 Department of Geography and Ecology, Ulyanovsk State Pedagogical University, Ulyanovsk, 432700, Russia. E-mail:[email protected] 3 Royal Belgian Institute of Natural Sciences, Brussels, B-1000 Belgium. E-mail: [email protected]
Abstract
We present a first checklist of Eulophidae from Cambodia, featuring 26 species belonging to 15 genera, and an additional nine species identified to generic level. Eighteen species are recorded for the first time for Southeast Asia. In Cambodia the Eulophidae are represented by four subfamilies: Eulophinae (13 species from 7 genera), Entiinae (1 species from 1 genus), Entedoninae (4 species from 4 genera) and Tetrastichinae (17 species from 8 genera).
Keywords: Entiinae, Entedoninae, Eulophinae, Tetrastichinae, new records, Southeast Asia
Introduction
The inventory of chalcids in Cambodia began in 1991 and until recent only three species of Chalcidoidea were known from Cambodia, belonging to different families: Kriechbaumerella kraussi Narendran, 1989 (Chalcididae) (deposited in USNM United States Museum of Natural History, Washington D.C., USA); Plutarchia bicarinativentris Girault, 1925 (Eurytomidae) (deposited in QMB, Queensland Museum, Brisbane, Australia) and Oodera rufimana Westwood, 1874 (Pteromalidae) (deposited in Oxford Museum, UK). Three additional species (Euplectrus colliosilvus Wijesekera & Schauff, 1994, Euplectrus koebelei Crawford, 1911, and Euplectrus leucostomus Rohwer, 1921) have recently been discovered in Cambodia (YEFREMOVA, 2017). In Southeast Asia, fifty-eight species of Eulophidae are known from Vietnam (YEFREMOVA, 1994; ZHU & HUANG, 2002, 2003; YEFREMOVA & STRAKHOVA, 2009; STRAKHOVA & YEFREMOVA, 2013; YEFREMOVA, 2017), 39 species from Thailand (KAMIJO, 1986; GRAHAM, 1987; BOUČEK, 1988; ZHU & HUANG, 2003; NARENDRAN, 2007; STRAKHOVA & YEFREMOVA, 2013; YEFREMOVA, 2017), nine species from Myanmar (HUSAIN & KHAN, 1986; STRAKHOVA & YEFREMOVA, 2013), and five species from Brunei (BOUČEK 1988; UBAIDILLAH & LA SALLE, 1996). No Eulophidae species have been recorded to date from Christmas Island, East Timor, or Laos. This paper provides new distributional records of Eulophidae (Hymenoptera, Chalcidoidea) for Cambodia and also for Southeast Asia discovered in the entomological collections during a Synthesys visit of the first author at Royal Belgian Institute of Natural Sciences (RBINS) in 2015.
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YEFREMOVA Z., YEGORENKOVA E. & DEKONINCK W. Eulophidae from Cambodia (Hymenoptera: Chalcidoidea)
Map 1. Map of Cambodia showing both localities where of Eulophidae were collected during 2004 and 2006.
Material and methods
This report is based on 46 specimens collected from the Siem Reap Province in Cambodia at Sam Veasna Center for Wildlife Conservation and at the Preah Khan Angkor Temple, located Northeast of Angkor Thom in a degraded semi-evergreen forest, and just west of the Jayatataka baray (about 100 km from Thailand), as well as from the Prey Nob mangrove, 10°38’7”N, 103°47’1”E (Map 1). The material was collected by Frédéric Goes, Inariddh Var, Yothin Oul (volunteers at RBINS during 2005) and Kai-Olaf Krüger between 2004 and 2006, during the GTI project “Biodiversity assessment at three protected areas in Northwest Cambodia” and deposited in RBINS (Brussels, Belgium). During this project, Malaise traps were installed at Sam Veasna Center (3 years) and Preah Khan Angkor Temple (2 years) and more than 50.000 insect specimens were sorted out at family or higher taxonomically level by specialists. Specimens of the superfamily of Chalcidoidea were not sorted out from the microhymenoptera before 2015. The specimens were examined and some of them were illustrated under a Nikon (SMZ800) dissecting microscope (equipped with a camera lucida). Habitus photographs were taken with a Nikon (D 300S) camera.
Abbreviations: M.T. = Malaise trap, RBINS = Royal Belgian Institute of Natural Sciences.
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Taxonomy
Subfamily EULOPHINAE Westwood, 1829 Genus Cirrospilus Westwood, 1832 1. Cirrospilus nigriscutum (Girault, 1915) MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Elephant Site, 20-27.II.2004, M.T., Leg. I. Var. HOSTS: Unknown. DISTRIBUTION: Australia (BOUČEK, 1988); first record for Southeast Asia. Genus Diglyphomorphomyia Girault, 1913 2. Diglyphomorphomyia sp.1. MATERIAL EXAMINED: 1♂, Cambodia, Siem Reap, Preah Khan Angkor Temple, 24.I- 21.II.2006, M.T., Leg. Oul Yothin; 1♂, Cambodia, Siem Reap, Preah Khan Angkor Temple, 20-27.I.2005, M.T., Leg. I. Var. Genus Elachertus Spinola, 1811 3. Elachertus australis (Girault, 1913) MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes; 2♂, Cambodia, Siem Reap, Preah Khan Angkor Temple, 8.II.2005, M.T., Leg. I. Var. HOSTS: Unknown. DISTRIBUTION: Australia (BOUČEK, 1988); first record for Southeast Asia. 4. Elachertus malabaricus Narendran, 2011 MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes. HOSTS: Unknown. DISTRIBUTION: India (NARENDRAN, 2011); first record for Southeast Asia. 5. Elachertus sobrinis (Girault & Dodd, 1915) MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes. HOSTS: Unknown. DISTRIBUTION: Indonesia, Sulawesi (UBAIDILLAH, 2003); Australia (GIRAULT, 1915); first record for Southeast Asia. REMARKS: The species is recognized by having additional carinae on the lateral sides of the propodeal median carina. Genus Elasmus Westwood, 1833 6. Elasmus brevicornis Gahan, 1922 (Fig. 1) MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Kbal Spean, 13-27.III.2005, M.T., Leg. K.O. Kruger. HOSTS: Ectoparasitoid of Gelehiidae, Hesperiidae (Lepidoptera) (YEFREMOVA & STRAKHOVA, 2009; STRAKHOVA & YEFREMOVA, 2013). DISTRIBUTION: Afrotropical and Oriental regions (YEFREMOVA, 2007; YEFREMOVA & STRAKHOVA, 2009).
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YEFREMOVA Z., YEGORENKOVA E. & DEKONINCK W. Eulophidae from Cambodia (Hymenoptera: Chalcidoidea)
Figs 1-4. 1. Elasmus brevicornis, lateral view; 2. Euplectrus colliosilvus, lateral view; 3. Euplectrus koebelei, lateral view; 4. Acrias varicornis, dorsal view.
7. Elasmus grimmi Girault, 1920 MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Preah Khan Angkor Temple, forest edge, 01-08.II.2006, M.T., Leg. I. Var. HOSTS: Unknown. DISTRIBUTION: Australia (GIRAULT, 1920), India (VERMA et al., 2002), Thailand (STRAKHOVA & YEFREMOVA, 2013). 8. Elasmus viridiscutellatus Verma & Hayat, 2002 MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Preah Khan Angkor Temple, 01.VII.2006, M.T., Leg. Oul Yothin. HOSTS: Ectoparasitoid of Cnaphalocrocis medinalis (Guenée) (Lepidoptera, Pyralidae) (VERMA et al., 2002; KAZMIM & GIRISH KUMAR, 2015). DISTRIBUTION: India (VERMA et al., 2002); first record for Southeast Asia. Genus Euplectrus Westwood, 1832 9. Euplectrus colliosilvus Wijesekera & Schauff, 1994 (Fig. 2) MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes. HOSTS: Unknown. DISTRIBUTION: Sri Lanka (WIJESEKERA & SCHAUFF, 1994), Cambodia (YEFREMOVA, 2017).
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10. Euplectrus koebelei Crawford, 1911 (Fig. 3) MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Sam Veasna Center garden, 15-31.I.2004, M.T., Leg. F. Goes. HOSTS: Unknown. DISTRIBUTION: Japan, Malaysia, China, Taiwan, Papua New Guinea (CRAWFORD, 1911; ZHU & HUANG, 2003), Cambodia (YEFREMOVA, 2017). 11. Euplectrus leucostomus Rohwer, 1921 MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Sam Veasna Center, 28.XII.2005, M.T., Leg. I. Var. HOSTS: Parasitoid of larva of Achia sp. (Noctuidae) and larva of Pyralidae (FERRIÈRE, 1941); ectoparasitoid of Lasiocampidae and Noctuidae (Lepidoptera) (ZHU & HUANG, 2003). DISTRIBUTION: Sri Lanka, China, Mexico (ZHU & HUANG, 2003), India (FERRIÈRE, 1941; HERTING, 1977), Malaysia (FERRIÈRE, 1941); Cambodia (YEFREMOVA, 2017). Genus Notanisomorphella Girault, 1913 12. Notanisomorphella sp. (Figs 9-10) MATERIAL EXAMINED: 1♂, Cambodia, Siem Reap, Sam Veasna Center, 28.XII.2004, M.T., Leg. I. Var. HOSTS: Unknown. DISTRIBUTION: Australia, Africa (BOUČEK, 1988), India (NARENDRAN, 2011); first record for Southeast Asia. Genus Noyesius Bouček, 1988 13. Noyesisus nr. testaceus Bouček, 1988 MATERIAL EXAMINED: 1♂, Cambodia, Siem Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes. HOSTS: Unknown. DISTRIBUTION: New Zealand (BOUČEK, 1988); first record for Southeast Asia. REMARKS: Paratype (examined): Female with label: New Zealand, 19 km E of Tapu, 31 January 2018, J. Noyes (Natural History Museum, London, UK) was examined. Antenna with 6 flagellomeres and 3-segmented clava. Propodeum sculptured. Face is covered by numerous setae that were not visible in paratype. Petiole long with lateral projection with one seta. Subfamily ENTIINAE Hedqvist, 1974 Genus Acrias Walker, 1847 14. Acrias varicornis (Girault, 1922) (Fig. 4) MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes. HOSTS: Unknown. DISTRIBUTION: Australia (BOUČEK, 1988); first record for Southeast Asia. Subfamily ENTEDONINAE Förster, 1856 Chrysonotomyia Ashmead, 1904 15. Chrysonotomyia sp. 1. MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Sam Veasna Center, 13-20.XII.2004, M.T., Leg. I. Var.
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Genus Euderomphale Girault, 1916 16. Euderomphale sp.1. MATERIAL EXAMINED: 1♂, Cambodia, Siem Reap, Preah Khan Angkor Temple, 12- 19.XI.2004, M.T., Leg. WMF. Genus Neochrysocharis Kurdjumov, 1912 17. Neochrysocharis beasleyi Fisher & La Salle, 2005 MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes. HOSTS: Unknown. DISTRIBUTION: Indonesia, Vietnam (FISHER & LA SALLE, 2005). Genus Omphale Haliday, 1833 18. Omphale sp. 1. MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Preah Khan Angkor Temple, 1- 18.VII.2006, M.T., Leg. Oul Yothin. Subfamily TETRASTICHINAE Förster, 1856 Genus Aprostocetus Westwood, 1833 19. Aprostocetus kaya Narendran, 2007 MATERIAL EXAMINED: 1♀, 1♂, Cambodia, Siem, Reap, Sam Veasna Center, garden, 15- 31.I.2004, M.T., Leg. F. Goes. HOSTS: Unknown. DISTRIBUTION: India (NARENDRAN, 2007); first record for Southeast Asia. 20. Aprostocetus pausiris (Walker, 1839) MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes. HOSTS: Endoparasitoid of Dasineura leguminicola (Lintner) (Diptera, Cecidomyiidae) (BURKS, 1979; GRAHAM, 1987), Lipara sp. (Diptera, Chloropidae) (VIDAL, 1993). DISTRIBUTION: Nearctic (Canada), Palearctic (BURKS, 1979; GRAHAM, 1987; BOUČEK, 1988); first record for Southeast Asia. 21. Aprostocetus vimania Narendran, 2007 MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes. HOSTS: Unknown. DISTRIBUTION: India (NARENDRAN, 2007); first record for Southeast Asia. 22. Aprostocetus near venustus (Gahan, 1914) MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Preah Khan Angkor Temple, 26.VI.2006, M.T., Leg. Oul Yothin. HOSTS: Contarinia sorghicola (Coquillet), (Cecidomyiidae, Diptera) (YEFREMOVA et al., 2007). DISTRIBUTION: Palearctic and Nearctic regions (GRAHAM, 1987); first record for Southeast Asia.
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Figs 5-8. 5. Nesolynx thymus, lateral view; 6. Nesolynx thymus, face, frontal view; 7. Neotrichoporoides beonus, female, lateral view; 8. Neotrichoporoides beonus, female, dorsal view.
23. Aprostocetus (Ootetrastichus) sp. 1. MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Sam Veasna Center, in garden, 15- 31.I.2004, M.T., Leg. F. Goes. Genus Baryscapus Förster, 1856 24. Baryscapus evonymellae (Bouché, 1834) MATERIAL EXAMINED: 2♀, Cambodia, Siem Reap, Preah Khan Angkor Temple, 18- 25.XII.2005, M.T., Leg. Oul Yothin; 1♀, Cambodia, Siem Reap, Sam Veasna Center, 13- 20.XII.2004, M.T., Leg. I. Var.
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YEFREMOVA Z., YEGORENKOVA E. & DEKONINCK W. Eulophidae from Cambodia (Hymenoptera: Chalcidoidea)
HOSTS: Endoparasitoid of Cynipidae (Hymenoptera), Lasiocampidae, Lyonetiidae, Pyralidae, Yponomeutidae (Lepidoptera) (HERTING, 1977; YEFREMOVA et al., 2007; BOUČEK, 1974; GRAHAM, 1991). DISTRIBUTION: Palearctic (Europe) (BOUČEK, 1974; GRAHAM, 1991); first record for Southeast Asia. 25. Baryscapus sp. 1. MATERIAL EXAMINED: 2♀, Cambodia, Siem Reap, Sam Veasna Center, 13-20.XII.2004, M.T., Leg. I. Var. Genus Citrostichus Bouček, 19880 26. Citrostichus phyllocnistoides (Narayanan, 1960) MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Preah Khan Angkor Temple, 01.VII.2006, M.T., Leg. Oul Yothin. HOSTS: Endoparasiotid of Triozidae (Hemiptera), Gracillariidae, Nepticulidae (Lepidoptera) (DASH & DAS, 1997; BOUČEK, 1988). DISTRIBUTION: Thailand (SAKAKIBARA & MORAKOTE, 1998); Philippines (KAMIJO & IKEDA, 1997). Genus Neotrichoporoides Girault, 1913 27. Neotrichoporoides viridimaculatus (Fullaway, 1955) MATERIAL EXAMINED: 1♂, Cambodia, Siem Reap, Sam Veasna Center, 02-09.III.2005, M.T., Leg. I. Var; 1♀, Cambodia, Siem Reap, Sam Veasna, garden, 2-8.I.2005, M.T., Leg. I. Var. HOSTS: Endoparasiotid of Pseudaletia unipuncta (Haworth) (Noctuidae, Lepidoptera) (PECK, 1963). DISTRIBUTION: Nearctic, Palearctic regions (BURKS, 1979; DOMENICHINI, 1966; GRAHAM, 1987); first record for Southeast Asia. 28. Neotrichoporoides beonus Narendran, 2006 (Figs 7–8) MATERIAL EXAMINED: 2♀, Cambodia, Siem Reap, Sam Veasna Center, 15-31.I.2004, M.T., Leg. F. Goes. HOST: Unknown. DISTRIBUTION: India (NARENDRAN, 2007); first record for Southeast Asia. Genus Nesolynx Ashmead, 1905 29. Nesolynx thymus (Girault, 1916) (Figs 5–6) MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Preah Khan Angkor Temple, 1-8.II.2005, M.T., Leg. I. Var.; 1♀, Cambodia, Siem Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes. HOSTS: Endoparasitoid of Pyralidae (Lepidoptera) (NARENDRAN, 2007); Tachinidae (Diptera) (BANSUDE et al., 2010). DISTRIBUTION: Australia, India, Indonesia, Sri Lanka (BOUČEK, 1988; NARENDRAN, 2007), Myanmar (HUSAIN & KHAN, 1986). Genus Oomyzus Rondani, 1870 30. Oomyzus scaposus (Thomson, 1878) MATERIAL EXAMINED: 1♀, Cambodia, Siem, Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes.; 1♀, Cambodia, Prey Nob mangrove, 10°38’7’’N, 103°47’1’’E, M.T., 8- 12.V.2015, Leg. J. Constant & V. Sougnes. DISTRIBUTION: Cosmopolitan; first record for Southeast Asia.
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Figs 9–10. Notanisomorphella sp. 9. Body, dorso-lateral view; 10. Head with antenna, lateral view.
Genus Quadrastichus Girault, 1913 31. Quadrastichus vacuna (Walker, 1839) MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap, Angkor, 18-25.XII.2005, M.T., Leg. Oul Yothin. HOSTS: Endoparasitoid of Cecidomyiidae (Diptera), Tenthredinidae (Hymenoptera), Lyonetiidae (Lepidoptera) (DOMENICHINI, 1966; GRAHAM, 1991; ASKEW & SHAW, 1974). DISTRIBUTION: Palearctic (Europe) (DOMENICHINI, 1966; GRAHAM, 1991; ASKEW & SHAW, 1974); first record for Southeast Asia. 32. Quadrastichus zenta Narendran, 2007 MATERIAL EXAMINED: 1♀, Cambodia, Siem, Reap, Sam Veasna Center, garden, 23- 30.II.2005, M.T., Leg. I. Var. HOSTS: Unknown. DISTRIBUTION: India (NARENDRAN, 2007); first record for Southeast Asia. 33. Quadrastichus sp. 1. MATERIAL EXAMINED: 1♀, Cambodia, Siem Reap Province, Preah Khan Angkor Temple, 01.VII.2006, M.T., Leg. Oul Yothin. 34. Quadrastichus sp. 2. MATERIAL EXAMINED: 1♀, Cambodia, Siem, Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes. Tetrastichus Haliday, 1844 35. Tetrastichus howardi (Olliff, 1893) MATERIAL EXAMINED: 1♂, Cambodia, Siem, Reap, Sam Veasna Center, garden, 15-31.I.2004, M.T., Leg. F. Goes. HOSTS: Tenebrio molitor L. (Coleoptera, Tenebrionidae), Busseola fusca (Fuller) (Noctuidae), Holocerina smilax (Westwood) (Saturnidae) (Lepidoptera), Apis mellifera L. (Hymenoptera) (POLASZEK & LA SALLE, 1995); Sesamia calamistis Hampson (Lepidoptera, Noctuidae) (POLASZEK & LA SALLE, 2007). DISTRIBUTION: Cosmopolitan.
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YEFREMOVA Z., YEGORENKOVA E. & DEKONINCK W. Eulophidae from Cambodia (Hymenoptera: Chalcidoidea)
Conclusions
In this paper a first overview of all so far collected and identified Eulophidae from Cambodia is reassembled. This checklist represents 66.6% of the fauna so far known from Thailand (39 species according to NOYES (2018)). In Thailand the subfamily Eulophinae (19 species) is more studied than in Cambodia, however for the subfamily Tetrastichinae (13 species in Thailand) this is the reverse. In Vietnam 58 species of Eulophidae have been found so far (40 species in NOYES (2018) and 18 species in YEFREMOVA (2017)), in China 388 species are known and according to NOYES (2018) for the Oriental fauna 1049 species are recorded so far. We hope this first checklist for Cambodia will promote more collecting and research on Eulophidae in the country and Southeast Asia in general as there are still many things to discover, and taxonomy needs revision for many genera.
Acknowledgements
This research received support from the SYNTHESYS Project http://www.synthesys.info/ financed by the European Community Research Infrastructure Action under the FP7 Integrating Activities Programme to travel to and study the collections at the Royal Belgian Institute of Natural Sciences (Belgium) for the first author. Most of the specimens were collected during a GTI project “Biodiversity assessment at three protected areas in Northwest Cambodia” and therefore we want thank Dr Patrick Grootaert who was promotor of the project under the reference CBD/GTI/ys/IC/2005.04. This project was supported through a grant issued by the Belgian Global Taxonomic Initiative National Focal Point that runs under the CEBioS programme, RBINS, with financial support from the Belgian Directorate-General for Development Cooperation (DGD). We thank Yvonnick Gérard for his help in RBINS collections. We thank Roger Burks (University of California, Riverside, USA) for discussion on the identification within the genus Notanisomorphella. We want to thank Dr Patrick Grootaert and Isabelle Coppée for their useful comments on a previous version of the manuscript and Isabelle Sauvage for the lay-out.
References
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