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Zootaxa, Systematic Ambiguity in the Well-Established Model System Bernasconi, M V; Berger, D; Blanckenhorn, W U (2010). Systematic ambiguity in the well-established model system insect Scathophaga stercoraria (Diptera: Scathophagidae): sister species S. soror revealed by molecular evidence. Zootaxa, 2441:27-40. Postprint available at: http://www.zora.uzh.ch University of Zurich Posted at the Zurich Open Repository and Archive, University of Zurich. Zurich Open Repository and Archive http://www.zora.uzh.ch Originally published at: Zootaxa 2010, 2441:27-40. Winterthurerstr. 190 CH-8057 Zurich http://www.zora.uzh.ch Year: 2010 Systematic ambiguity in the well-established model system insect Scathophaga stercoraria (Diptera: Scathophagidae): sister species S. soror revealed by molecular evidence Bernasconi, M V; Berger, D; Blanckenhorn, W U Bernasconi, M V; Berger, D; Blanckenhorn, W U (2010). Systematic ambiguity in the well-established model system insect Scathophaga stercoraria (Diptera: Scathophagidae): sister species S. soror revealed by molecular evidence. Zootaxa, 2441:27-40. Postprint available at: http://www.zora.uzh.ch Posted at the Zurich Open Repository and Archive, University of Zurich. http://www.zora.uzh.ch Originally published at: Zootaxa 2010, 2441:27-40. Zootaxa 2441: 27–40 (2010) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2010 · Magnolia Press ISSN 1175-5334 (online edition) Systematic ambiguity in the well-established model system insect Scathophaga stercoraria (Diptera: Scathophagidae): sister species S. soror revealed by molecular evidence MARCO VALERIO BERNASCONI1*, DAVID BERGER1 & WOLF U. BLANCKENHORN1 1Zoological Museum, Institute of Evolutionary Biology and Environmental Studies, University of Zurich, Winterthurerstrasse 190, CH- 8057 Zurich, Switzerland E-mails: [email protected]; [email protected]; [email protected] *corresponding author Abstract Even for well-established insect model systems, such as the yellow dung fly Scathophaga stercoraria (Linnaeus) (Diptera: Scathophagidae), there may be hidden systematic ambiguities that require clarification. Dung flies from the Afrotropical Region have been considered (i) as con-specific and not different from all the other Holarctic Scathophaga stercoraria; (ii) as a local and peculiar African subspecies of S. stercoraria (Scathophaga stercoraria soror Wiedemann), or (iii) as a separate valid species (Scathophaga soror Wiedemann). Our study represents an attempt, based on mitochondrial (COI, 12S, and 16S), nuclear (ITS2) as well as microsatellite markers, to clarify this problem. Results strongly suggest that S. soror is a separate taxon from S. stercoraria. Due to the importance of S. stercoraria as a model system for studies in ecology, behaviour and evolution, the systematic position of S. soror (relative to S. stercoraria) is not solely of interest for systematists, but for evolutionary ecologists as well. Key words: Systematics, Phylogeny, Speciation, Cytochrome oxidaese I, COI, 12S, 16S, ITS2, internal transcribed spacer 2, microsatellites Introduction Scathophagid flies, with about 400 species described, are mainly confined to the Holarctic Region and are more northern in overall distribution than any other family of Diptera (e.g., Sack 1937; Vockeroth 1987; but see also Šifner 2003, 2008, 2009). Only a few species occur in South Africa and at high altitudes in East Africa, the Andes, and the Oriental Region (see references cited in Bernasconi et al. 2000a). Many adult scathophagids are predators (e.g., on Simuliidae; Werner et al. 2006), and several Arctic species are regularly observed on carrion and mammalian dung. Some species breed in rotting seaweed. The eggs are attached to the leaf surface or inserted into plant tissues, dung, or other substrates. Most of the larvae are phytophagous, while others are carnivorous in dung or coprophagous (Gorodkov 1986; Vockeroth 1987; Šifner 2008). Individuals of most species thus perform the ecologically important function of resource recycling. Within this family, the species Scathophaga stercoraria (Linnaeus) has served since the early 1960s in numerous ecological, behavioural and evolutionary investigations, particularly as model system for studies of sperm competition, cryptic female choice, and life history evolution (summarised in Ward 2007). The species is included in the first tier of the Flytree (Assembling the Diptera Tree of Life) project for this reason (http:// www.inhs.illinois.edu/research/FLYTREE/). From a morphological point of view there is no unanimity within the entomological community about the taxonomic status of the South African dung flies. They have been considered (i) as con-specific and not different from all the other S. stercoraria worldwide (e.g., Šifner 2008), (ii) as a peculiar subspecies (Scathophaga stercoraria soror Wiedemann, e.g., Vockeroth 1958), or even (iii) as a separate species (Scathophaga soror Wiedemann, e.g., Werner et al. 2006). Accepted by J. O’Hara: 23 Mar. 2010; published: 30 Apr. 2010 27 The relationships in this family are now generally well understood (Bernasconi et al. 2000a, 2000b, 2001; Kutty et al. 2007, 2008), but the systematic position (and identity) of S. (stercoraria) soror remains enigmatic, as this taxon has never been included in any molecular systematic or phylogenetic analysis of the family. Our study represents therefore an attempt, based on mitochondrial (COI, 12S, and 16S), nuclear (ITS2) as well as microsatellite markers, to clarify this problem. Material and methods Samples A total of 50 specimens representing 21 species of Scathophagidae were included in the present study (Table 1). Whenever possible, various specimens belonging to the same species, but from different geographic origin, were included to gain information about the intra-specific genetic diversity of the species examined. In particular, six individuals of S. (stercoraria) soror from three different South African localities, as well as 14 specimens of S. stercoraria from Europe, Asia, and North America were included. Four Gimnomera species were used for outgroup comparison (see Bernasconi et al. 2000a; Kutty et al. 2007). TABLE 1. Overview of samples and species of Scathophagidae used in this study. Specimen voucher Species Origin GenBank accession number (sample id in Fig. 1 & COI 12S 16S ITS2 Table 7) Bernasconi 0286 (P) Ceratinostoma ostiorum United Kingdom AF180792 DQ656914 DQ648668 EU884226 (286) (Curtis, 1823) Bernasconi 0308 (P) Gimnomera cerea Old Chelsea, Quebec, AF181009 DQ656917 DQ648671 NA (308) Coquillett, 1908 Canada Bernasconi 0227 (P) Gimnomera cuneiventris Abisko, Sweden AF181011 DQ656918 DQ648672 NA (227) (Zetterstedt, 1846) Bernasconi 0222 (P) Gimnomera dorsata Abisko, Sweden AF181008 DQ656919 DQ648673 EU884220 (222) (Zetterstedt, 1838) Bernasconi 0291 (P) Gimnomera tarsea Whitlaw Moss, United AF181010 DQ656920 DQ648674 EU884229 (291) (Fallén, 1819) Kingdom Bernasconi 0250 (P) Scathophaga analis Prague, Czech AF180783 DQ656942 DQ648696 NA (250) (Meigen, 1826) Republic Bernasconi 0287 (P) Scathophaga calida United Kingdom AF180787 DQ656943 DQ648697 EU884227 (287) Curtis, 1832 Bernasconi 0019 (P) Scathophaga cineraria Switzerland AF180784 DQ656944 DQ648698 EU884209 (19) (Meigen, 1826) Bernasconi 0005 (P) Scathophaga furcata Zurich, Switzerland AF180775 NA NA EU884204 (5) (Say, 1823) Bernasconi 0013 (P) Scathophaga furcata Zurich, Switzerland AF180776 NA NA EU884205 (13) Bernasconi 0042 (P) Scathophaga furcata Zurich, Switzerland AF180777 DQ656945 DQ648699 EU884212 (42) Bernasconi 0267 (P) Scathophaga incola Gronubakken, Norway AF180786 DQ656946 DQ648700 EU884224 (267) (Becker, 1900) Bernasconi 0268 (P) Scathophaga inquinata Hordaland, Norway AF180781 DQ656947 DQ648701 EU884225 (268) (Meigen, 1826) ......continued on the next page 28 · Zootaxa 2441 © 2010 Magnolia Press BERNASCONI ET AL. TABLE 1 (continued) Specimen voucher Species Origin GenBank accession number (sample id in Fig. 1 & COI 12S 16S ITS2 Table 7) Bernasconi 0294 (P) Scathophaga inquinata Finland AF180782 NA NA EU884231 (294) Bernasconi 0218 (P) Scathophaga litorea Leuchtturm Kjolnes, AF180789 DQ656948 DQ648702 EU884219 (218) (Fallén, 1819) Norway Bernasconi 0038 (P) Scathophaga lutaria Zurich, Switzerland AF180778 NA NA EU884210 (38) (Fabricius, 1794) Bernasconi 0046 (P) Altbüron LU, Scathophaga lutaria AF180779 DQ656949 DQ648703 EU884214 (46) Switzerland Bernasconi 0108 (P) Scathophaga lutaria Har Meron, Israel AF180780 NA NA EU884215 (108) Bernasconi 0293 (P) Scathophaga obscura United Kingdom AF180790 DQ656950 DQ648704 EU884230 (293) (Fallén, 1819) Bernasconi 0266 (P) Scathophaga pictipennis Kvam, Norway AF180785 DQ656951 DQ648705 EU884223 (266) (Oldenberg, 1923) Bernasconi 00S1 (P) Scathophaga soror Kwa Zulu-Natal, EU884256 EU884234 EU884244 EU884192 (S1) Wiedemann, 1818 South Africa Bernasconi 00S2 (P) Kwa Zulu-Natal, Scathophaga soror EU884257 EU884235 EU884245 EU884193 (S2) South Africa Bernasconi 00S3 (P) Cape Province, South Scathophaga soror EU884258 EU884236 EU884246 NA (S3) Africa Bernasconi 00S4 (P) Cape Province, South Scathophaga soror EU884259 EU884237 EU884247 EU884194 (S4) Africa Bernasconi 00S7 (P) Western Cape, South Scathophaga soror EU884262 EU884240 EU884250 EU884197 (S7) Africa Bernasconi 00S8 (P) Western Cape, South Scathophaga soror EU884263 EU884241 EU884251 EU884198 (S8) Africa Bernasconi 00S5 (P) Scathophaga stercoraria Hokkaido, Japan EU884260 EU884238 EU884248 EU884195 (S5) (Linnaeus, 1758) Bernasconi 00S6 (P) Scathophaga stercoraria Hokkaido, Japan
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