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Proceedings of the Entomological PROC. ENTOMOL. SOC. WASH. 107(4), 2005, pp. 750-761 BAIMAIA, A NEW SUBGENUS FOR ANOPHELES KYONDAWENSIS ABRAHAM, A UNIQUE CRABHOLE-BREEDING ANOPHELINE IN SOUTHEASTERN ASIA Ralph E. Harbach, Rampa Rattanarithikul, and Bruce A. Harrision (REH) Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. (email: [email protected]); (RR) Museum of World Insects, 72 Nimamhemin 13, Huay-Kaeo Road, Chiangmai 50200, Thailand (email: insects. [email protected]); (BAH) Public Health Pest Management, North Carolina Depart- ment of Environment and Natural Resources, 585 Waughtown Street, Winston-Salem, NC 27107 ([email protected]) Abstract. —Baimaia, n. subg., is introduced as a new subgenus of Anopheles for the unusual crabhole species, Att. kyondawensis Abraham, in Southeast Asia. A diagnosis of the subgenus is provided that features unique anatomical characters of the adult, larval, and pupal stages of the type species. The larva of An. kyondawensis is redescribed and the previously unknown adult female, adult male, and pupa are described in detail. The affinities of Baimaia and An. kyondawensis are discussed in terms of their position in the phylogeny of Anophelinae, and their bionomics and distribution are reviewed. Key Words: Culicidae, Anophelinae, taxonomy, mosquito Abraham (1947) described Anopheles pools that follow flooding or rains." In {Anopheles) kyondawensis from larvae 1979, another larva was collected from a found in shallow ground pools along stream in Huai Kop, Sai Yok District of streams near the village of Kyondaw Kanchanaburi Province in western Thailand (Moulmein Township, Mon State) in south- (Harrison et al. 1991). Likewise, it is likely ern Myanmar. The species was not encoun- that this larva may have been dislodged tered again until 1966 when a single larva from a crabhole because the margin of the was found in a crabhole at Ban Pha Man stream where it was captured was lined located near the Laos border in Nan Prov- with these habitats at water level. Go et al. ince of Thailand (Harrison and Scanlon (2004) found one larva of An. kyondawen- 1975). Harrison and Scanlon (1975) sug- sis in a shaded pool along a stream in the gested that the larvae collected by Abraham vicinity of Innwaing near the type locality in Myanmar may have been swept out of during collections made in Myanmar be- freshwater crabholes by high water because tween May 1998 and March 2000. These they were found in association with larvae authors did not indicate whether the pool he identified as An. (Cellia) leucosphyrus was associated with crabholes. Material ex- Donitz [probably An. baimaii Sallum and amined during the present study, however, Peyton (recently described in Sallum et al. confirms that An. kyondawensis does in fact 2005) since it is the dominant species of the breed in burrows created by freshwater Leucosphyrus Group in Myanmar], which crabs. This material consists of larvae, and "normally occur in small temporary ground adults reared from larvae and pupae col- VOLUME 107, NUMBER 4 751 lected from crabholes in Ban Tham Sua, slowly evolving single-copy nuclear white Tak Province, located in northwestern Thai- gene, Krzywinski and Besansky (2003) hy- land. pothesized that Bironella diverged from the Reid and Knight (1961) included An. main lineage of Anopheles following the kyondawensis in the Culiciformis Group of earlier separation of Chagasia. Although subgenus Anopheles based on the reduced this hypothesis is not supported by the mor- setae 5,6,7-C, and other setae of the larval phological and molecular phylogenetic head capsule. Harrison and Scanlon (1975) studies of Sallum et al. (2000 and 2002, re- considered this placement tentative until the spectively), it is not inconsistent with the adult and pupal stages were known. Fol- results of the more recent cladistic analysis lowing the discovery of the previously un- of Harbach and Kitching (2005), one aim known adult and pupal stages, we initiated of which was to investigate the phyloge- studies of this species, and after having neiic position of An. kyondawensis. This considered all of the unique features in the latter analysis placed An. kyondawensis as adult, larval, and pupal stages noted below, sister to Bironella + all other Anopheles, we concluded that An. kyondawensis does with Chagasia as sister to these three taxa. not belong in any currently recognized spe- Although this arrangement of taxa, ex- cies group of Anopheles. pressed parenthetically as Chagasia + {An. The traditional classification of subfami- kyondawensis + {Bironella + other Anoph- ly Anophelinae included three genera: eles)), raises questions concerning the bio- Anopheles Meigen, Bironella Theobald, geography of anophelines, support for this and Chagasia Cruz. The phylogenetic re- set of relationships (assessed using Bremer lationships of these genera, based on a cla- and relative Bremer support) is strong and distic analysis of morphological data (Har- indicates that both An. kyondawensis and bach and Kitching 1998), reflect the intui- Bironella are independent lineages relative tive hypothesis (Ross 1951) that Anophel- to the rest of Anophelinae (see Harbach and inae is a monophyletic clade comprised of Kitching (2005) for a full assessment of re- Chagasia in a sister-group relationship to lationships and character support). These Bironella + Anopheles. Molecular phylog- results agree with the suggestion by Sallum enies inferred from nuclear and mitochon- et al. (2002) that ""Bironella may be plau- drial gene sequences also support this hy- sibly regarded as a subgenus of Anophe- pothesis of relationships (Besansky and les'', and imply in accordance with appli- Fahey 1997; Foley et al. 1998; Krzywinski cation of the principle of equivalent rank et al. 2001a, b), but more recent studies (Hennig 1966) that An. kyondawensis based on both morphological (Sallum et al. should also be afforded subgeneric rank. 2000, Harbach and Kitching 2005) and mo- Hence, a new subgenus is proposed herein lecular data (Sallum et al. 2002) suggest for this species. that Anopheles is a paraphyletic assemblage Materials and Methods relative to Bironella. In the absence of sup- port for the generic status of Bironella, Sal- This study is based on a small number of lum et al. (2000) formally synonymized larvae, and adults reared from larvae and/ this taxon with Anopheles s.s. This synon- or pupae collected from crabholes (as in- ymy, however, is not supported (see below) dicated above), and the holotype larva of by the later studies of Sallum et al. (2002) An. kyondawensis deposited in The Natural and Harbach and Kitching (2005), which History Museum (NHM), London (see Ma- indicate that Bironella should be regarded terial examined following the species de- as a subgenus of Anopheles. scription). Because the medium in which Taking account of independent lines of the holotype was nKuuited on a microscope evidence, especially sequence data for the slide had turned black with age. the speci- — — — 752 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON men was removed following the procedures al and inner setae, and the gonostylus is of Brown and De Boise (2005) and re- flattened and mitten-shaped distally and mounted in Euparal on the same slide. The lacks a gonostylar claw. Males also have head and dissected mouthparts of the larva uniquely developed maxillary palpi, which were mounted under a separate coverslip. are straight and very nearly cylindrical with Following stereoscopic examination, the palpomeres 4 and 5 barely swollen and only head of the only available female was re- slightly flattened. Adults lack thoracic scal- moved, cleared in 5% NaOH for 2 h at ing, and females have a dense covering of 50°C, and mounted, with the mouthparts long sensilla between the antennal whorls separated from the head capsule, in euparal that depart a fuzzy appearance to the anten- on a microscope slide for more detailed nae. The immature stages are found in crab- study. The genitalia of 2 available males holes, or in pools after having been washed were also dissected and likewise cleared out of these habitats. Pupae have a trumpet and mounted on individual microscope that appears undifferentiated and interme- slides. Pinned adults were examined under diate between angusticorn and laticorn, spi- simulated natural light; dissections, larvae, racular scars of abdominal segments II-VII and larval and pupal exuviae were studied and seta 9-II-VIII borne ventrally, seta 1- with differential interference contrast op- III plumose, and a long fringe of spicules tics. Measurements and counts were made on the inner and outer margins of the pad- from all available specimens. Numbers in dle. Larvae have strongly inwardly curved parentheses represent modes, when appar- antennae, setae 5,6,7-C reduced, long, sin- ent, of the reported ranges. The anatomical gle, compressed or flattened, somewhat lan- terminology and abbreviations used in the ceolate setae on the thorax and abdomen, descriptions and illustrations, respectively, and setae 6-IV-VI as long as setae 6-I-III. follow Harbach and Knight (1980, 1982). Baimaia is monobasic: see in the System- The symbols ?, 6, Le, Pe, and L used in atics section following the description of the literature summary and material exam- An. kyondawensis for discussion. ined sections for An. kyondawensis repre- Etymology. Baimaia is a patronymic sent female, male, larval exuviae, pupal ex- honoring Prof. Visut Baimai of Mahidol uviae, and fourth-instar larva, respectively. University, Bangkok, for his many impor- An asterisk (*) after one of these symbols tant contributions to our knowledge of the in the literature summary section indicates cytogenetics and systematics of Anopheles at least part of the life stage was illustrated mosquitoes in southeastern Asia. We have in the publication cited. chosen to Latinize Visut's surname by add- ing the feminine suffix "-a" rather than the Taxonomic Treatment masculine "-M5'" because Baimaia is more euphonious and easier to pronounce {BT-mT- Anopheles subgenus Baitnaia, Harbach, Rattanarithikul, and Harrison, new fJ).
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