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Utility of the Mitochondrial Cytochrome Oxidase II Gene for Resolving Relationships Among Black Flies (Diptera: Simuliidae) K

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Utility of the Mitochondrial Cytochrome Oxidase II Gene for Resolving Relationships Among Black Flies (Diptera: Simuliidae) K University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Papers in Plant Pathology Plant Pathology Department 8-2000 Utility of the Mitochondrial Cytochrome Oxidase II Gene for Resolving Relationships among Black Flies (Diptera: Simuliidae) K. P. Pruess University of Nebraska-Lincoln, [email protected] B. J. Adams University of Florida, [email protected] T. J. Parsons University of Nebraska-Lincoln X. Zhu University of Nebraska-Lincoln Thomas O. Powers University of Nebraska-Lincoln, [email protected] Follow this and additional works at: http://digitalcommons.unl.edu/plantpathpapers Part of the Other Plant Sciences Commons, Plant Biology Commons, and the Plant Pathology Commons Pruess, K. P.; Adams, B. J.; Parsons, T. J.; Zhu, X.; and Powers, Thomas O., "Utility of the Mitochondrial Cytochrome Oxidase II Gene for Resolving Relationships among Black Flies (Diptera: Simuliidae)" (2000). Papers in Plant Pathology. 443. http://digitalcommons.unl.edu/plantpathpapers/443 This Article is brought to you for free and open access by the Plant Pathology Department at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Papers in Plant Pathology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Molecular Phylogenetics and Evolution Vol. 16, No. 2, August, pp. 286–295, 2000 doi:10.1006/mpev.2000.0807, available online at http://www.idealibrary.com on Utility of the Mitochondrial Cytochrome Oxidase II Gene for Resolving Relationships among Black Flies (Diptera: Simuliidae) K. P. Pruess,* B. J. Adams,†,‡ T. J. Parsons,†,§ X. Zhu,* and T. O. Powers† *Department of Entomology, University of Nebraska, Lincoln, Nebraska 68583-0816; †Department of Plant Pathology, University of Nebraska, Lincoln, Nebraska 68583; ‡Department of Nematology, University of Florida, Gainesville, Florida 32601; and §Armed Forces DNA Identification Laboratory, Rockville, Maryland 20850 Received August 31, 1999, revised May 3, 2000 Chironomoidea with Chironomidae (midges) and Cera- The complete mitochondrial cytochrome oxidase II topogonidae (biting midges) as sister groups. These gene was sequenced from 17 black flies, representing taxa are included, with Culicidae (mosquitoes) and 13 putative species, and used to infer phylogenetic other families, in the infraorder Culicomorpha. Molec- relationships. A midge (Paratanytarsus sp.) and three ular data, based on 28s rDNA gene sequences mosquitoes (Aedes aegypti, Anopheles quadrimacula- (Pawlowski et al., 1996), did not support these relation- tus, and Culex quinquefasciatus) were used as out- ships, suggesting that Thaumaleidae (solitary midges) group taxa. All outgroup taxa were highly divergent and Simuliidae are sister groups and that Chironomi- from black flies. Phylogenetic trees based on weighted dae may be a sister group to all other Nematocera. parsimony (a priori and a posteriori), maximum like- Using a different 28s rDNA sequence, Friedrich and lihood, and neighbor-joining (log-determinant dis- tances) differed topologically, with deeper nodes be- Tautz (1997) found the Culicomorpha to form a ing the least well-supported. All analyses supported strongly supported monophyletic clade but the rela- current classification into species groups but relation- tionships between families were poorly resolved. ships among those groups were poorly resolved. The The fossil record for Simuliidae is meager. The ear- majority of phylogenetic signal came from closely re- liest known fossil, a pupa from middle Jurassic (170 lated sister taxa. The CO-II gene may be useful for mybp) is considered indistinguishable from modern exploring relationships at or below the subgeneric Prosimulium, and larvae from the lower Cretaceous level, but is of questionable value at higher taxonomic (120 mybp) of Australia are indistinguishable from levels. The weighting method employed gave phyloge- modern Simulium (Crosskey, 1990). Quite likely, the netic results similar to those reported by other au- divergence of modern genera was complete in the thors for other insect CO-II data sets. A best estimate Triassic. of phylogenetic relationships based on the CO-II gene Two subfamilies are commonly recognized in Simu- is presented and discussed in relation to current black liidae, with all but four species placed in the subfamily fly classification. © 2000 Academic Press Simuliinae. Simuliinae is divided into two tribes, Pro- Key Words: black fly; Simuliidae; mitochondrial simuliini and Simuliini, but with disagreement as to DNA; cytochrome oxidase subunit II; phylogeny; char- tribal placement of some genera. The genus Simulium acter weighting. contains over 1000 species (Crosskey and Howard, 1997). The current classification of black flies recog- nizes “species groups” within subgenera without infer- INTRODUCTION ring relationships among those groups (Crosskey and Howard, 1997). Currie (1988) provides an alternative Black flies (Diptera: Simuliidae) are a large group of classification at the generic level. medically and economically important Diptera in the Black fly taxonomy is further complicated by sibling suborder Nematocera which have speciated with little species complexes whose component species cannot be morphological change. A paucity of diagnostic charac- morphologically differentiated in all life stages. Many ters for all life stages complicates species identification sibling species are best characterized by larval salivary and therefore control efforts often target black flies in gland chromosome polymorphisms. Sibling species are general, rather than the few pest species. inferred from lack of heterozygotes in areas of sympa- Previous workers agree that Simuliidae is a mono- try (Rothfels, 1979). Larvae of these sibling species are phyletic group. Wood and Borkent (1989) and Ooster- usually isolated temporally, spatially, or ecologically broek (1995) place Simuliidae in the superfamily (Adler, 1987). Within species groups, a number of cy- 286 1055-7903/00 $35.00 Copyright © 2000 by Academic Press All rights of reproduction in any form reserved. BLACK FLY PHYLOGENETICS 287 TABLE 1 Collection Data for Black Flies Sequenced Species Source Heledon dicentum Dyar & Shannon California, Napa Co., 22 March 1990 Prosimulium onychodactylum Dyar & Shannon Colorado, Jackson Co., 5 Aug. 1989 Cnephia sp. A (putative C. dacotensis Dyar & Shannon) New Hampshire, Strafford Co., 4 May 1990 C. sp. B [putative C. pecuarum (Riley)] Texas, Rains Co., 4 Jan. 1991 C. sp. C (putative C. pecuarum) Texas, Rains Co., 4 Jan. 1991 Simulium (Eusimulium) pilosum (Knowlton & Rowe) (ϭ S. aureum “B”) Nebraska, Lancaster Co., 24 April 1989 S. (Eusimulium) donovani Vargas G1 (ϭ S. aureum “G”) California, Mono Co., 25 June 1991 S. (Eusimulium) donovani G2 California, Mono Co., 25 June 1991 S. (Psilozia) vittatum Zetterstedt sibling IIIL-1 Nebraska, Lancaster Co., 25 March 1988 S. (Psilozia) vittatum “Iceland” Iceland, R. Laxa, 31 Aug. 1990 S. (Psilozia) encisoi Vargas & Diaz Najera New Mexico, Dona Ana Co., 2 June 1992 S. (Psilopelmia) bivittatum Malloch A Nebraska, Loup Co., 12 Oct. 1990 S. (Psilopelmia) bivittatum B Wyoming, Crook Co., 21 May 1994 S. (Simulium) piperi Dyar & Shannon (hunteri group) Nebraska, Scottsbluff Co., 11 July 1989 S. (Simulium) jacumbae Dyar & Shannon (hunteri group) Nebraska, Lancaster Co., 30 Oct. 1992 S. (Simulium) luggeri Malloch (jenningsi group) Nebraska, Loup Co., 12 July 1990 S. (Simulium) confusum Moulton & Adler (jenningsi group) Nebraska, Saline Co., 6 July 1990 Outgroups Paratanytarsus sp. (Chironomidae) Pennsylvania, Chester Co., Jan. 1993 Aedes aegypti (L.) (Culicidae) Ho et al. (1995) (GenBank L34412) Anopheles quadrimaculata Say (Culicidae) Mitchell et al. (1993) (GenBank L04272) Culex quinquefasciatus Say (Culicidae) Ho et al. (1995) (GenBank L34351) tophylogenies based on chromosomal inversions exist MATERIALS AND METHODS (e.g., Rothfels, 1979), but these are unrooted. The cytochrome oxidase subunit II (CO-II) gene is Black Fly Collection and DNA Extraction one of the best-known mitochondrial genes. Cooper et Larval black flies were either preserved in ethanol al. (1991) and Saraste (1990) review the structure and upon collection or returned to the laboratory alive and function of the polypeptide subunits. CO-II contains preserved individually in TE buffer at Ϫ70°C. The both highly conserved and variable regions. The se- midge was obtained as an adult. With two exceptions quence of this gene is potentially useful for phyloge- netic analyses over a wide taxonomic range and has (to be discussed), all specimens were obtained from been explored for that purpose. However, Liu and localities where, according to collecting and rearing Beckenbach (1992) were unable to resolve the evolu- data, other closely related species did not occur. Col- tion of insect orders and concluded that divergence lection data are given in Table 1. DNA was obtained may have occurred too rapidly for resolution using using proteinase K digestion, phenol/chloroform ex- mitochondrial genes. This sequence has been generally traction, and ethanol precipitation as described by useful for reconstructing phylogenies among more Wilkerson et al. (1993). closely related groups (Willis et al., 1992; Beckenbach PCR and Sequencing et al., 1993; Brown et al., 1994; Sperling and Hickey, 1994; Emerson and Wallis, 1995; Spicer, 1995; Smith Using the notation of Simon et al. (1994), primers and Bush, 1997; Foley et al., 1998). With the exception used to amplify the complete CO-II gene were of
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