See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/233329088 Wolbachia infections in bees (Anthophila) and possible implications for DNA barcoding Article in Systematics and Biodiversity · December 2011 DOI: 10.1080/14772000.2011.627953 CITATIONS READS 21 254 3 authors, including: Michael Gerth Christoph Bleidorn University of Liverpool Georg-August-Universität Göttingen 48 PUBLICATIONS 150 CITATIONS 123 PUBLICATIONS 2,303 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: Platyzoa View project All content following this page was uploaded by Michael Gerth on 16 September 2014. The user has requested enhancement of the downloaded file. This article was downloaded by: [University of Leipzig] On: 16 September 2014, At: 06:02 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Systematics and Biodiversity Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tsab20 Wolbachia infections in bees (Anthophila) and possible implications for DNA barcoding MICHAEL GERTH a , ANNEMARIE GEIßLER a & CHRISTOPH BLEIDORN a a Molecular Evolution and Systematics of Animals, Institute for Biology , University of Leipzig , Talstr. 33, D-04103, Leipzig, Germany Published online: 25 Nov 2011. To cite this article: MICHAEL GERTH , ANNEMARIE GEIßLER & CHRISTOPH BLEIDORN (2011) Wolbachia infections in bees (Anthophila) and possible implications for DNA barcoding, Systematics and Biodiversity, 9:4, 319-327, DOI: 10.1080/14772000.2011.627953 To link to this article: http://dx.doi.org/10.1080/14772000.2011.627953 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. 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Terms & Conditions of access and use can be found at http:// www.tandfonline.com/page/terms-and-conditions Systematics and Biodiversity (2011), 9(4): 319–327 Perspective Wolbachia infections in bees (Anthophila) and possible implications for DNA barcoding MICHAEL GERTH, ANNEMARIE GEIßLER & CHRISTOPH BLEIDORN Molecular Evolution and Systematics of Animals, Institute for Biology, University of Leipzig, Talstr. 33, D-04103 Leipzig, Germany (Received 15 February 2011; revised 24 August 2011; accepted 25 August 2011) The widespread intracellular bacterium Wolbachia is transmitted exclusively maternally and alters the reproduction of its hosts by different mechanisms. Thereby, inheritance patterns of mitochondrial genomes are modified, possibly confining interpretations of mitochondrial sequence data. Although this phenomenon has been reported before, its conclusions seem to be widely ignored. In the light of recent large-scale barcoding projects relying solely on mitochondrial cox1 sequences, we screened the native German bee fauna (Anthophila) for Wolbachia infections. The screening revealed that 66% of the native German bees and 54% of sphecid wasps are infected by Wolbachia. Many species bore identical or similar infections, suggesting a high rate of horizontal transfer. Supergroup A infections were recovered in most cases; only one species bore a super-group F Wolbachia infection. Because Wolbachia is not only present in 66% of bees but also in the majority of arthropod species, we argue that studies interpreting sequence data of arthropod species cannot rely on mitochondrial data alone – nuclear markers must be incorporated. DNA barcoding using only mitochondrial cox1 will not be sufficient to delimit, identify or discover Wolbachia-infected species, i.e. probably the majority of all animal species. Key words: bees (Anthophila), DNA barcoding, molecular taxonomy, sphecid wasps, Wolbachia Introduction mitochondrial cytochrome oxidase subunit I gene (cox1)is Bees (Anthophila) are a group of aculeate Hymenoptera used as a barcode (Hebert et al., 2003). The backbone of which include over 16,000 described species (Michener, the initiative is a database including sequence and voucher- 2000). The monophyly of bees is well established based on ing information for curated specimens (Ratnasingham and molecular and morphological characters, however, ingroup Hebert, 2007). Within this initiative, different campaigns relationships are still a matter of ongoing debate (Danforth aim to facilitate massive DNA barcoding for selected an- et al., 2006). Many interesting evolutionary phenomena imal taxa or to barcode the complete fauna of certain ge- have been reported from bees, including evolution of euso- ographic regions. In the case of bees, a ‘Bee Barcode of ciality (Andersson, 1984), cleptoparasitism (Cardinal et al., Life Initiative’ (http://www.bee-bol.org/) has been brought Downloaded by [University of Leipzig] at 06:02 16 September 2014 2010) and coevolution with flowering plants (Johnson to life. As a first step, the bee fauna of Nova Scotia (Canada) & Steiner, 2000). Besides these, bees are known for their has already been extensively barcoded (Sheffield et al., important role as pollinators and as such they contribute 2009). Bees are also flagship species of the Barcoding Fauna to an immense part of the worldwide human food supply Bavarica campaign (http://www.faunabavarica.de/). (Greenleaf & Kremen, 2006). This makes research on bee A central dogma of DNA barcoding is the postulation of a biodiversity, conservation and autecology economically im- ‘barcoding gap’ which means that intraspecific genetic vari- portant topics (Losey & Vaughan, 2006). ability of mitochondrial cox1 is significantly lower than its Unsurprisingly, bees became a major target of the Bar- interspecific variability (Hebert et al., 2003, 2004b). The code of Life initiative (http://www.barcodeoflife.org/). The existence of such a ‘barcoding gap’ has repeatedly been aim of this initiative is to provide highly standardized ge- questioned in studies on different animal groups (Meyer netic marker systems for species identification (Golding & Paulay, 2005; Meier et al., 2006; Wiemers & Fiedler, et al., 2009). In the case of animals, a 648 bp region of the 2007) and for theoretical reasons (Hickerson et al., 2006). For some insects, this gap does not seem to exist at all Correspondence to: Michael Gerth. E-mail: michael.gerth@uni- (Virgilio et al., 2010). Moreover, the reliance on a single mi- leipzig.de tochondrial marker has been regarded as not adequate due to ISSN 1477-2000 print / 1478-0933 online C 2011 The Natural History Museum http://dx.doi.org/10.1080/14772000.2011.627953 320 M. Gerth et al. reduced effective population size, introgression, maternal Wenseleers et al., 1998; Russell et al., 2009; Lachowska inheritance, inconsistent mutation rate, pseudogenization et al., 2010). and heteroplasmy (Rubinoff et al., 2006; Song et al., 2008; In our study we simply asked: How prevalent are Wol- Galtier et al., 2009; Magnacca & Brown, 2010). Further- bachia infections in the native German bee fauna? We draw more, the presence of cytoplasmic bacteria can dramatically general conclusions for barcoding studies (for which this alter inheritance patterns of mitochondrial genes (Hurst & bee fauna is a major target) and suggest that it is critical to Jiggins, 2005). include nuclear markers in future DNA barcoding studies There are different lineages of symbionts that are capa- of arthropods. ble of inducing such effects, including Cardinium, Rick- ettsia, Spiroplasma (e.g. Duron et al., 2008; Perlman et al., 2008; Weinert et al., 2009) and, probably most widespread, Materials and methods Wolbachia. This genus of intracellular α-proteobacteria is All specimens were collected between August 2009 and found in arthropods and nematodes. Since its first discov- July 2010. The animals were captured with hand nets or ery in ovaries of Culex pipiens in 1924 (Hertig & Wolbach, snap-cap vials in the vicinity of their nests, their host’s 1924), numerous strains were found across many insect or- nest or on their pollen source. To cover a broad range of ders as well as in isopods, amphipods, ostracods, arachnids taxa, sampling took place at different times of the year and and filarial nematodes (Ros et al., 2009). The high inter- at various locations in Germany. Altogether, 126 bees (75 est in and increasing number of studies about Wolbachia species) and 16 sphecid wasps (13 species) were collected is due to its ability to alter the reproduction of its