Mosquito fauna of Lao People's Democratic Republic, with special emphasis on the adult and larval surveillance at Nakai District, Khammuane Province.
The mosquito fauna of the Lao PDR are not well known, except for several scattered reports. (7-14) In this study, we updated the records and checklist of mosquito species from the Lao PDR based on the literature, specimens deposited at the US National Mosquito Collections (USNMC), National Museum of Natural History (NMNH), Smithsonian Institution, Washington, DC, and our latest specimen collections from Khammuane Province, particularly at the Phou Hin Poun National Biodiversity Conservation Area (PHP NBCA). This area, which has a human population of approximately 30,000, is located in a limestone tower karst region of the Annamite Range in Khammuane Province. It is composed mainly of rugged caves, porous karst terrain, and dry evergreen forest and scrubland. It is also the home to a number of rare or newly discovered species of animals. (15-17) We are in the process of confirming the identification of several species of mosquitoes and sand flies, and possibly describing new species from our recent collections in the area.
MATERIALS AND METHODS
Mosquito Field Collection, Museum Specimens and Identification
Specimen collections were conducted from May 1 to May 31, 2012, and from February 21 to March 10, 2014, from various areas in the PHP NBCA (17.99524[degrees] N, 104.82108[degrees] E), Ban Natan, Nakai District, Khammuane Province (Figure 1). Adults were collected using modified Centers for Disease Control and Prevention traps (Figure 2A, B) with light attractants, and were suspended about 1.3 m above ground level on selected sites and inside the caves. Larvae were collected using a standard larval dipper (350 ml, 13 cm diameter: BioQuip, Rancho Dominguez, CA) (Figure 2 C, D) from various habitats including water pockets along edges of rivers, rock holes, temporary pools in between rocks, caves, etc (Figures 2, 3, and 4). They were individually link-reared to adult stage, as morphological voucher specimens for this work. Emergent adults were pinned on paper points, each given a unique collection number, properly labeled, and identified using diagnostic morphological characters. (18-23) Voucher specimens were deposited at the USNMC NMNH, Smithsonian Institution, Washington, DC, USA, and at the Entomology Laboratory, Institut Pasteur du Laos, Vientiane, Lao PDR. In addition, old mosquito specimens at the NMNH repository were examined, and their collection data were recorded.
The summary of mosquito collections from the PHP NBCA, Khammuane Province is presented in Table 1. Figure 1 shows the map of the Lao PDR, with 10 of 16 provinces, Vientiane (capital city) and PHP NBCA (all with asterisks as shown in the map) where adult and larval mosquitoes were collected or reported in the literature. In the PHP NBCA, mosquito habitats included water pockets along edges of rivers, rock holes, temporary pools along the edges of rivers, in between rocks, and in caves (Figures 2-4). A total of 43 mosquito taxa were collected from PHP NBCA in 9 genera (Aedes, Anopheles, Culex, Heizmannia, Mansonia, Orthopodomyia, Topomyia, Toxorhynchites, Tripteroides). Among the 3 genera examined, Aedes (19 species) had the greatest number of species, followed by Culex (8 species) and Anopheles (7 species). Only 18 species out of 43 (42%) were morphologically identified, while the rest (25 species; 58%) need further analyses (including molecular techniques) to clarify their taxonomic identities. Known or potential vectors of human infectious diseases were also collected from PHP NBCA, including Aedes vexans (Meigan), Ae albopictus (Skuse), and several unconfirmed species of Anopheles (Anopheles), An (Cellia), Culex (Culex), and Mansonia.
An updated checklist of mosquitoes in the Lao PDR (Table 2) includes a total of 101 species. They are in 16 genera, namely Aedes (22 species), Anopheles (33), Armigeres (14), Coquillettidia (2), Culex (12), Ficalbia (1), Heizmannia (1), Hodgesia (1), Mansonia (4), Mimomyia (2), Orthopodomyia (1), Topomyia (1), Toxorhynchites (2), Tripteroides (2), Uronotanea (2), Verrallina (1). About 80 of 101 species were reported in the Walter Reed Biosystematics Unit (WRBU) catalog, (23) 2 species found from the Smithsonian/NMNH collections, 17 species from current PHP NBCA collections, and the remaining species from the literature. About 10 species of mosquitoes are new records for the Lao PDR. They include 9 species under 7 subgenera of the genus Aedes and one species in the genus Orthopodomyia (Table 2).
The Lao PDR, like other countries comprising the GMS, has a high biodiversity of vector species, a great number of mosquito species complexes, enormous spatial heterogeneity in distribution patterns, and extensive behavioral plasticity both between and within species (2).
In 1934 (7) and 1938, (8) Anopheles mosquitoes were reported in the Laos PDR (Table 2). In December 1999, malaria vector surveys were carried out by Vythilingam et al (11) in 7 provinces, namely Borikhamxay, Champasak, Luangprabang, Saravane, Savannakhet, Xayaboury, and Sekong, and in the capital city of Vientiane in the Lao PDR. Using bare leg collections from indoors and outdoors from 6 PM to 5 PM, a total of 438 Anopheles mosquitoes belonging to 19 species were obtained. Of these, only 3 species were found infected with oocysts, namely An maculatus Theobald, An dirus Peyton and Harrison, and An minimus Theobald. Anopheles aconitus Doenitz was the predominant species in the 1999 collection, but its vectorial status was unknown. The prevalence of Anopheles and epidemiology of malaria were also reported in the provinces of Xekong (12) and Attapeu. (13,14) In 2014, Hii and Rueda (2) listed 3 species in Anopheles (Anopheles) and 20 species in Anopheles (Cellia) in the Lao PDR, including known and potential malaria vectors in countries of the Mekong Subregion. While there are numerous examples of An dirus mostly feeding outdoors and much earlier in the evening, (24,25) Vythilingam et al (13) reported an unusual stereotypical nocturnal indoor and late feeding behavior in Attapeu province, Laos PDR. In 2002, Tsuda et al (10) conducted an ecological survey of Aedes dengue vectors in the central part of the Lao PDR. A new hydroelectric project, Nam Theun 2, created ideal conditions for Aedes aegypti (Linnaeus) breeding in water storage jars and tires, and Ae albopictus was abundant. (26)
The present study indicates the species diversity of mosquitoes in the Lao PDR. The difficulty in doing morphological comparisons among species warrants further molecular analysis to ascertain taxonomic identities and to clarify hierarchic classifications. With the diversity of the habitats, particularly the caves and surrounding areas, we expect that more unknown species will be collected and described in the near future. Deforestation, water resources and management, (27,28) conventional agricultural practices, and unregulated destruction of many habitats are major human activities that may adversely affect the floral and animal fauna of the Lao PDR, including the creation or elimination of suitable breeding sites of mosquitoes and other arthropods. While habitats in some government protected areas are not hugely damaged yet, continuous inventories of arthropod fauna, particularly those groups (mosquitoes, sand flies, ticks, mites, etc) with known disease vectors, should be conducted to accumulate much needed data for developing strategies to manage and control infectious human diseases. Proper vector surveillance, including ecological surveys, should be performed in areas where human diseases (malaria, dengue, tick-borne viruses, filariasis, etc) are common and severely affect the local human populations. The updated checklist of mosquitoes in this article (including several vector species) may help health personnel in mapping out some risk areas for infectious diseases in the Lao PDR.
We express sincere gratitude and appreciation to Lea Thutkhin for collecting, processing, mounting and pinning specimens, and staff of the Institut Pasteur du Laos for assistance; to Sonexay Ounekham of the Entomology Unit, Center for Malaria, Parasitology, and Entomology, Vientiane, for help in collecting specimens in Khammuane Province. We are grateful to Yiau-Min Huang, Jeffrey Clark, and Belen P. Rueda for reviewing this manuscript and for their valuable comments. Special thanks go to the staff of the Public Health Office, Nakai District, Khammuane Province for cooperation and field assistance.
Partial funding was provided by the US Naval Medical Research Center-Asia through the Global Emerging Infections Surveillance and Response System, a Division of the US Armed Forces Health Surveillance Center, Silver Spring, MD. Institut Pasteur du Laos and the Lao Ministry of Health also contributed to support this project.
This research was performed under a Memorandum of Understanding between the Walter Reed Army Institute of Research and the Smithsonian Institution, with institutional support provided by both organizations.
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Leopoldo M. Rueda, PhD
Khamsing Vongphayloth, MD
James E. Pecor, BS
LCDR Ian W. Sutherland, USN
Jeffrey Hii, PhD
Mustapha Debboun, PhD
Paul T Brey, PhD
Dr Rueda is a Research Entomologist, Principal Investigator, and former Chief of the Walter Reed Biosystematics Unit, Entomology Branch, Walter Reed Army Institute of Research located at the Smithsonian Institution, Museum Support Center, Suitland, Maryland.
Dr Vongphayloth is a Medical Doctor and Entomologist, Institut Pasteur du Laos, Vientiane, Laos PDR
Mr Pecor is a Museum Specialist at the Walter Reed Biosystematics Unit, Entomology Branch, Walter Reed Army Institute of Research located at the Smithsonian Institution, Museum Support Center, Suitland, Maryland.
LCDR Sutherland is the Chief of Entomological Sciences, US Naval Medical Research Center - Asia located at the U.S. Navy Region Center, Sembawang, Singapore.
Dr Hii, formerly a WHO Malaria Scientist, is an Adjunct Principal Research Fellow in the School of Public Health, Tropical Medicine and Rehabilitation Sciences, James Cook University based in Bangkok, Thailand.
Dr Debboun is the Director, Mosquito Control Division, Harris County Public Health & Environmental Services, Houston, TX.
Dr Brey is a Research Entomologist and Director of the Institut Pasteur du Laos, Vientiane, Laos PDR.
Table 1. Summary of collected mosquito adults and larvae in Phou Hin Poun NBCA, Ban Natan, Nakai District, Khammuane Province, Lao PDR (17.99524[degrees]N, 104.82108[degrees]E), from May 1 thru May 31, 2012, and February 21 thru March 10, 2014. Species Sex * Collection no. Aedes (Aedimorphus) 3F LN-048, alboscutellatus 050, 060 (Theobald) Aedes (Aedimorphus) 1F LN-012 sp Aedes (Aedimorphus) 1F LN-047 vexans (Meigen) Aedes (Bothealla) 3F ([dagger]) LN-002, eldridgei Reinert 041, 068 Aedes (Bothealla) sp 3F, 1M LN-018, 022, 023, 069 Aedes (Collessius) 1F LN-013 sp Aedes (Downsiomyia) 1F LN-024 ganapathi Colless Aedes (Downsiomyia) 1F LN-001 harinasutai Knight Aedes (Downsiomyia) 1M LN-008 sp Aedes (Fredwardsius) 2F, 1M LN-015, vittatus (Bigot) 065, 066 Aedes (Hulecoeteomyia) 1F LN-035 chrysolineatus (Theobald) Aedes (Hulecoeteomyia) 1F, 1M LN-036, 037 formosensis Yamada Aedes (Hulecoeteomyia) 1F, 1M LN-031, 046 sp (near reinerti or formosensis) Aedes (Kenknightia) 1F LN-063 dissimilis (Leicester) Aedes (Kenknightia) 1F LN-044 sp Aedes (Stegomyia) 1F LN-043 albopictus (Skuse) Aedes (Stegomyia) 1F LN-039 pseudoscutellaris (Theobald) Aedes (Tewarius) 2F LN-003 pseudonummatus Reinert Aedes sp 4F, 2M LN-011, 013, 019, 020, 026, 042 Anopheles (Anopheles) sp 1F LN-011 (Barbirostris Group) Anopheles (Anopheles) sp 1F LN-049 (Asiaticus Group) Anopheles (Anopheles) sp 1F LN-046 Anopheles (Anopheles) sp 1F LN-062 (Culiciformis Group) Anopheles (Cellia) 1F LN-045 pseudowillmori Theobald Anopheles (Cellia) sp 1F LN-005 (Leucosphyrus Group) Anopheles (Cellia) sp 2F LN-009, 010 Coquillettidia (Coquillettidia) 1F LN-004 ochracea (Theobald) Culex (Culex) sp (Vishnui 1F LN-052 Complex) Culex (Culex) sp (Sitiens Group) 1F LN-054 Culex (Culex) sp 3F LN-028, 055, 071 Culex (Culex) tritaeniorhynchus 1F LN-053 Giles Culex (Culiciomyia) 2F, 1M LN-064, 074,075 nigropunctatus Edwards Culex (Culiciomyia) sp 1F, 1M LN-067, 070 Culex (Eumelanomyia) sp 1F, 2M LN-017, 072, 073 (Temipalpus Complex) Culex (Lophoceraomyia) sp 1F, 1M LN-007, 016 Heizmannia sp 3F LN-025, 038, 059 Mansonia (Mansonioides) uniformes 1F LN-057 (Theobald) Mansonia sp 1F LN-051 Orthopodomyia albipes Leicester 1M LN-033 Orthopodomyia sp 1F LN-032 Topomyia sp 1F LN-030 Toxorhynchites sp 3M ([dagger]) LN-076, 077, 078 Tripteroides sp 2F, 1M LN-021, 040, 058 * F indicates female adult, M indicates male adult. ([dagger]) Larvae collected using plastic larval dipper. Table 2A. Updated checklist of mosquito species from Lao PDR. Species Reference (a) Aedes (Aedimorphus) alboscutellatus 14, 23, X (Theobald) Aedes (Aedimorphus) pipersalatus (Giles) 14, 23 Aedes (Aedimorphus) vexans (Meigen) 14, 23, X Aedes (Bothaella) eldridgei Reinert (b) X Aedes (Collessius) macfarlanei (Edwards) 23 Aedes (Diceromyia) iyengari Edwards 14, 23 Aedes (Downsiomyia) ganapathi Colless (b) X Aedes (Downsiomyia) harinasutai Knight (b) X Aedes (Downsiomyia) niveus (Ludlow) 23 Aedes (Fredwardsius) vittatus (Bigot) 10, X Aedes (Hulecoeteomyia) chrysolineatus 14, 23, X (Theobald) Aedes (Hulecoeteomyia) formosensis X Yamada (b) Aedes (Hulecoeteomyia) reinerti X Rattanarithikul and Harrison (b) Aedes (Kenknightia) dissimilis X (Leicester) (b) Aedes (Neomelaniconion) lineatopennis 19 (Ludlow) Aedes (Paraedes) ostentatio (Leicester) 19 Aedes (Phagomyia) prominens (Barraud) (b) X Aedes (Stegomyia) albopictus (Skuse) 14, X Aedes (Stegomyia) aegypti (Linnaeus) 10, 23, M Aedes (Stegomyia) pseudalbopictus Borel 14, 23 Aedes (Stegomyia) pseudoscutellaris X (Theobald) (b) Aedes (Tewarius) pseudonummatus X Reinert (b) Anopheles (Anopheles) albotaeniatus 11 (Theobald) Anopheles (Anopheles) argyropus 2 (Swellengrebel) Anopheles (Anopheles) baileyi Edwards 23 Anopheles (Anopheles) barbirostris Van 2, 8, 11, 14, 23 der Wulp Anopheles (Anopheles) donaldi Reid 2, 14, 23 Anopheles (Anopheles) sinensis Wiedemann 3, 8 Anopheles (Anopheles) umbrosus (Theobald) 12 Anopheles (Cellia) aconitus Doenitz 2, 7, 11, 14, 23 Anopheles (Cellia) annularis Van der Wulp 2 Anopheles (Cellia) culicifacies Giles 2, 7, 23 Anopheles (Cellia) dirus Peyton and 2, 10, 11, 12, 23 Harrison Anopheles (Cellia) dravidicus Christophers 2, 14, 23 Anopheles (Cellia) harrisoni Harbach 23 and Manguin Anopheles (Cellia) indefinitus (Ludlow) 2, 23 Anopheles (Cellia) jamesii Theobald 2, 23 Anopheles (Cellia) jeyporiensis James 2, 7, 8, 12, 23 Anopheles (Cellia) karwari (James) 2, 11, 12, 23 Anopheles (Cellia) kochi Donitz 2, 8, 11, 14, 23 Anopheles (Cellia) maculatus Theobald 2, 7, 8, 11, 12, 14, 23 Anopheles (Cellia) minimus Theobald 2, 8, 11, 12, 14, 23 Anopheles (Cellia) nivipes (Theobald) 11, 12, 14, 23 Anopheles (Cellia) notanandai 2, 14, 23 Rattanarithikul and Green Anopheles (Cellia) pallidus Theobald 11, 12 (a) X indicates observed, field collection; M indicates observed, Smithsonian/National Museum of Natural History museum collection. (b) New record for Lao PDR. Table 2B. Updated checklist of mosquito species from Lao PDR Species Reference (a) Anopheles (Cellia) pampanai Buttiker and 11, 14, 23 Beales Anopheles (Cellia) philippinensis Ludlow 2, 8, 11, 12, 14, 23 Anopheles (Cellia) pseudowillmori Theobald 2, 14, 23, X Anopheles (Cellia) rampae Harbach and Somboon 29 Anopheles (Cellia) sawadwongporni 14, 23 Rattanarithikul and Green Anopheles (Cellia) splendidus Koidzumi 11, 12, 14, 23 Anopheles (Cellia) subpictus Grassi 2, 11 Anopheles (Cellia) sundaicus (Rodenwaldt) 2 Anopheles (Cellia) tessellatus Theobald 2, 11, 14, 23 Anopheles (Cellia) vagus Donitz 2, 7, 8, 11, 12, 14 Anopheles (Cellia) varuna Iyengar 2, 11, 12, 14, 23 Armigeres (Armigeres) aureolineatus 23 (Leicester) Armigeres (Armigeres) durhami (Edwards) 23 Armigeres (Armigeres) kuchingensis Edwards 23 Armigeres (Armigeres) laoensis Toma and 23, M Miyagi (c) Armigeres (Armigeres) moultoni Edwards 23 Armigeres (Armigeres) setifer Delfinado 14, 23 Armigeres (Armigeres) subalbatus (Coquillett) 14, 23 Armigeres (Armigeres) theobaldi Barraud 14, 23 Armigeres (Leicesteria) annulitarsis 23 (Leicester) Armigeres (Leicesteria) dolichocephalus 23 (Leicester) Armigeres (Leicesteria) flavus (Leicester) 23, M Armigeres (Leicesteria) longipalpis 23 (Leicester) Armigeres (Leicesteria) magnus (Theobald) 23 Armigeres (Leicesteria) pectinatus (Edwards) 23 Coquillettidia (Coquillettidia) crassipes 14, 23 (Van der Wulp) Coquillettidia (Coquillettidia) ochracea 23, X (Theobald) Culex (Culex) fuscocephala Theobald 14, 23 Culex (Culex) gelidus Theobald 23 Culex (Culex) hutchinsoni Barraud 14, 23 Culex (Culex) pseudovishnui Colless 14, 23 Culex (Culex) quinquefasciatus Say 14, 23, M Culex (Culex) tritaeniorhynchus Giles 14, 23, X Culex (Culex) vishnui Theobald 14, 23 Culex (Culex) whitmorei (Giles) 14, 23 Culex (Culiciomyia) nigropunctatus Edwards 14, 23, X Culex (Oculeomyia) bitaeniorhynchus Giles 14, 23 Culex (Oculeomyia) pseudosinensis Colless 14, 23 Culex (Oculeomyia) sinensis Theobald 14, 23 Ficalbia minima (Theobald) 23 Heizmannia (Heizmannia) complex (Theobald) 23 Hodgesia malayi Leicester 23 (a) X indicates observed, field collection; M indicates observed, Smithsonian/National Museum of Natural History museum collection. (c) Holotype male, 1 paratype female, 4 females, 3 whole larvae, and 3 larval exuviae, deposited in the Smithsonian/National Museum of Natural History museum collection. Table 2C. Updated checklist of mosquito species from Lao PDR Species Reference (a) Mansonia (Mansonioides) annulifera (Theobald) 14, 23 Mansonia (Mansonioides) dives (Schiner) 14 Mansonia (Mansonioides) indiana Edwards 14, 23 Mansonia (Mansonioides) uniformes (Theobald) 14, 23, X Mimomyia (Mimomyia) Chamberlaini Ludlow 23 Mimomyia (Mimomyia) hybrida (Leicester) 23 Orthopodomyia albipes Leicester (b) X Topomyia (Topomyia) gracilis Leicester 23 Toxorhynchites (Toxorhynchites) albipes (Edwards) 23 Toxorhynchites (Toxorhynchites) kempi (Edwards) 23 Tripteroides (Rachionotomyia) aranoides (Theobald) 23 Tripteroides (Rachionotomyia) ponmeki Miyagi and Toma 9, 23 Uranotaenia (Pseudoficalbia) nivipleura Leicester 23, M Uranotaenia (Pseudoficalbia) novobscura Barraud 23, M Verrallina (Verrallina) dux (Dyar and Shannon) 23 (a) X indicates observed, field collection; M indicates observed, Smithsonian/National Museum of Natural History museum collection. (b) New record for Lao PDR.
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|Author:||Rueda, Leopoldo M.; Vongphayloth, Khamsing; Pecor, James E.; Sutherland, Ian W.; Hii, Jeffrey; Debbo|
|Publication:||U.S. Army Medical Department Journal|
|Date:||Jul 1, 2015|
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