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Diptera: Culicidae) Using DNA Barcoding and Polymerase Chain Reaction-Restriction Fragment Length Polymorphism

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Diptera: Culicidae) Using DNA Barcoding and Polymerase Chain Reaction-Restriction Fragment Length Polymorphism 1 Phylogenetics and molecular identification of the Ochlerotatus communis complex (Diptera: Culicidae) using DNA barcoding and polymerase chain reaction-restriction fragment length polymorphism Hooman H. Namin,1 Mahmood Iranpour, Barbara J. Sharanowski Abstract—The Ochlerotatus communis (De Geer, 1776) complex consists of four cryptic mos- quito species in North America, including: O. communis, Ochlerotatus churchillensis (Ellis and Brust, 1973), Ochlerotatus nevadensis (Chapman and Barr, 1964), and Ochlerotatus tahoensis (Dyar, 1916). Most of the morphological characters used for the identification of these species are quantitative and overlap across species. Here we evaluated the efficacy of DNA barcoding for identification of three members of the communis complex (O. nevadensis is not included in this study) and developed diagnostic restriction fragment length polymorphism (RFLP) patterns for O. communis and O. churchillensis. A phylogeny of 23 Ochlerotatus Lynch Arriba´lzaga, 1891 species was inferred using mitochondrial cytochrome c oxidase subunit I gene sequences. All species included in our analysis within the O. communis complex were delineated using cytochrome c oxidase subunit I barcodes. However, this complex was recovered as paraphyletic with respect to Ochlerotatus abserratus (Felt and Young, 1904) and Ochlerotatus implicatus (Vockeroth, 1954), indicating the need for increased genetic and taxonomic sampling to infer the phylogenetic rela- tionships of these taxa. The RFLP profile for multiple field specimens of O. communis was distinct from all RFLP patterns for O. churchillensis, and this method can be used as an efficient molecular method for the identification these species. Re´sume´—Le complexe d’Ochlerotatus communis (De Geer, 1776) comprend quatre espe`ces cryptiques de moustiques, soit O. communis, O. churchillensis (Ellis et Brust, 1973), O. nevadensis (Chapman et Barr, 1964) et O. tahoensis (Dyar, 1916). La plupart des caracte`res morphologiques utilise´s pour l’identification de ces espe`ces sont quantitatifs et il y a du recoupement entre les espe`ces. Nous e´valuons l’efficacite´ de la codification a` barres d’ADN pour l’identification de trois membres du complexe de communis (O. nevadensis n’est pas inclus dans notre e´tude) et mettons au point des patrons RFLP pour O. communis et O. churchillensis. Nous avons de´duit une phyloge´nie de 23 espe`ces d’Ochlerotatus Lynch Arriba´lzaga, 1891 a`partirdesse´quences du ge`ne mitochondrial de la sous-unite´I de la cytochrome c oxydase. Il a e´te´ possible de de´limiter toutes les espe`ces du complexe de communis incluses dans notre analyse a` l’aide des codes de barres de COI. Cependant le complexe s’est ave´re´ paraphyle´tique en ce qui a trait a` O. abserratus (Felt et Young, 1904) et O. implicatus (Vockeroth, 1954), ce qui indique qu’il faudra un plus important e´chantillonnage ge´ne´tique et taxonomique afin de de´duire les relations phyloge´ne´tiques de ces taxons. Le profil RFLP de nombreux e´chantillons de terrain d’O. communis se distingue des patrons RFLP d’O. churchillensis et la me´thode peut servir d’outil mole´culaire efficace pour l’identification de ces espe`ces. Introduction species in North America, including: O. communis (De Geer, 1776), O. churchillensis (Ellis and According to Ellis and Brust (1973), the Brust, 1973), Ochlerotatus nevadensis (Chap- Ochlerotatus communis complex consists of four man and Barr, 1964), and Otahoensis tahoensis Received 22 September 2012. Accepted 1 May 2013. H.H. Namin,1 M. Iranpour, B.J. Sharanowski, Department of Entomology, Faculty of Agricultural and Food Sciences, University of Manitoba, Winnipeg, Manitoba, Canada R3 T 2N2 1Corresponding author (e-mail: [email protected]). Subject editor: Patrice Bouchard doi:10.4039/tce.2013.60 Can. Entomol. 00: 1–10 ᭧ 2013 Entomological Society of Canada 2 Can. Entomol. Vol. 00, 2013 (Dyar, 1916). These species are morphologically their autogenous ovarian development. This cryptic and thus identification based on morphol- character may be useful to separate populations ogy is exceedingly difficult. Most characteristics of O. churchillensis from all other members of are diagnostic for only one species within the the complex (Ellis and Brust 1973). Morpho- complex or for a specific developmental stage or metric characteristics of the metatarsal claws are sex. For example, the shape of the larval comb the primary characters used for the identification scales is only diagnostic for the identification of of adult females, although the measurements are O. nevadensis in the fourth larval instar (Ellis not discrete with considerable overlap across and Brust 1973). In the adult stage, male mem- species (Brust and Munstermann 1992). bers of this complex may be distinguished by The combination of reproductive differences the size of genital gonocoxites (Ellis and Brust in addition to small morphological differences, 1973). Females, however, are much more difficult particularly for males, originally led Ellis and to identify. The shape of the metatarsal claw is Brust (1973) to describe O. churchillensis as a diagnostic for O. communis from other members distinct species from O. communis. Brust and of the complex, but this character is not always Munstermann (1992) suggested isozyme analysis reliable for distinguishing the other species (Brust as a method for distinguishing members of the and Munstermann 1992). O. communis complex. This method, however, Species within the O. communis complex have lacks consistency and requires very fresh speci- interesting distributions suggesting that both mens, thereby limiting the use of this technique as allopatric and sympatric speciation have played a routine tool for species identification (Schutz a role in their evolution. Ochlerotatus communis and Eldridge 1993). Currently, there is no reliable has a Holarctic distribution and has been collected method for quickly and accurately separating throughout much of North America (Wood et al. O. communis from O. churchillensis. 1979). Ochlerotatus nevadensis has been reported Accurate and rapid identification of pathogens from southeastern British Columbia, Canada and and vectors is essential in epidemiological studies of northwestern parts of the United States of America mosquito-borne diseases. Ochlerotatus communis is (Belton 1983) but has only been collected in a vector of Jamestown Canyon virus (JCV) and mountainous regions suggesting that this species snow shoe hare virus (SSHV) in North America might be reproductively isolated from other species (McLean et al. 1981; Heard et al. 1990), but the within the complex (Belton 1982; Schutz and clinical significance of O. churchillensis has not yet Eldridge 1993). Ochlerotatus tahoensis has a been studied. Both viruses are widely distributed restricted range of distribution from the Sierra across temperate areas of North America, and sev- Nevadas to the southern Cascades and Klamath eral boreal Aedes Meigen, 1818 and Ochlerotatus mountains in California, United States of America Lynch Arriba´lzaga, 1891 species are the primary but has only been collected from high elevations vectors (Grimstad 1988; Andreadis et al. 2008; (1500–3000 m; Schutz and Eldridge 1993). Bennett et al. 2012). Although JCV antibodies have Ochlerotatus tahoensis can be distinguished been detected in moose, elk, bison, mule deer, from O. nevadensis based on larval morpholo- domestic bovine and equine, white-tailed deer is the gical characters and from all species in the only known amplifying host in the natural trans- complex by collection locality (Darsie 1995). mission cycle of the virus (Grimstad 1988; Rust Ochlerotatus communis and Ochlerotatus et al. 1999). The common symptoms of JCV infec- churchillensis have a sympatric distribution in tion in humans ranges from nonspecific mild febrile northeastern and southeastern Manitoba and illnesses, acute central nervous system infection, western Alberta, Canada (Ellis and Brust 1973; and respiratory system involvement (Grimstad Wood et al. 1979) and thus are particularly dif- 1988, 2001). SSHV was isolated for the first time ficult to identify. These two species have similar from snowshoe hare (Lepus americanus Erxleben, larval habitats as both species can be found in 1777; Mammalia: Leporidae) in Montana, United snow-melt pools within or at the margins of States of America in 1959 (Burgdorfer et al. 1961). deciduous and coniferous forests (Ellis and Brust However, its antibodies also have recently been 1973). However, O. churchillensis females have detected in several small vertebrates and domestic smaller salivary glands than O. communis due to animals such as equine, cattle, dog, and chicken ᭧ 2013 Entomological Society of Canada Namin et al. 3 (Bennett et al. 2012). The symptoms of SSHV known localities for Ochlerotatus churchillensis) infection in humans include fever, headache, using aspirators between May and July 2011 vomiting, meningitis, and encephalitis (Fauvel (Table 1). The specimens of O. churchillensis, et al. 1980). Developing molecular markers could collected by Brust (Table 1), were obtained from improve the reliability of identification of the the Wallis–Roughley Museum of Entomology cryptic species in the O. communis complex and (University of Manitoba, Winnipeg, Manitoba, greatly facilitate mosquito vector surveillance. Canada). Additional specimens of this species Unlike morphological characters, molecular tools were obtained from the collected materials of can be applied for the identification of species in all a project
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