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Testing Species Boundaries in the Staphylinid Beetle Genus Mocyta (Insecta, Coleoptera, Staphylinidae)

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Testing Species Boundaries in the Staphylinid Beetle Genus Mocyta (Insecta, Coleoptera, Staphylinidae) Testing Species Boundaries in the Staphylinid Beetle Genus Mocyta (Insecta, Coleoptera, Staphylinidae) Master of Science in Ecology and Evolution Eilen Bjotveit Josefsen Natural History Museum University of Oslo 2014 ©Eilen Bjotveit Josefsen 2014 Eilen Bjotveit Josefsen Testing Species Boundaries in the Staphylinid Beetle Genus Mocyta http://www.duo.uio.no Trykk: Reprosentralen, Universitetet i Oslo II Acknowledgements This work was in part supported by SIU grant HNP-2012/10118 to Vladimir Gusarov. Thank you, Vladimir Gusarov for giving me this opportunity and for introducing me to the interesting world of athetine beetles. Your bottomless knowledge and sense of humour is highly esteemed. Lutz Bachmann, thank you for helping me with the molecular work, for the tips and tricks regarding my thesis and for making me laugh. Thank you, Hallvard Elven for suggestions and interesting conversations. Maria Mavrikidi, thank you for re-extracting and re-sequencing the ‘troublesome’ specimens. To all at the DNA lab at NHM, your assistance and company is invaluable. Special thanks go to Judith Osswald for introducing me to the lab, Eve Zeyl and Virginia Mirre for technical support, and lab manager Audun Schrøder-Nielsen for always having time to spare. Nikolai Yunakov and Peter Hammond, the specimens you provided for this study are highly appreciated. To Renee Labbee and Jeremy DeWaard at the Biodiversity Institute of Ontario, Mikko Pentinsaari at the University of Oulu, Oscar Vorst at Biodiversity Centre in Leiden and Jérôme Morinière at GBOL, thank you all for providing CO1 sequences from Mocyta specimens for my initial analyses, and thank you James Robertson at Bold for the referrals. I would also like to express my gratitude to Lars Jermiin, Henrik Enghoff, Boy Overgaard Nielsen, Jan Pedersen, Frank Jensen and Alexey Solodovnikov for trying to locate a copy of Viggo Mahler’s thesis “Acrotona studies”. Trude Magnussen and Kristine Dobbe, thank you for your support, company and long coffee breaks. To all at NHM, thank you for being so welcoming! June Susanne Berg, my dearest friend, thank you for your patience and discussions regarding this thesis. My beloved mother, you are an endless source of support and encouragement. Erik, no words can express my gratitude, without you, this thesis would never have existed. Oslo, 4th of August 2014 III IV Abstract The taxonomy of the staphylinid genus Mocyta was explored using molecular markers, in addition to a thorough investigation of the published literature regarding this genus and its proposed morphospecies. The aim was to test if the recognized morphospecies differ genetically, and if popular species delimitation methods can be used to delineate these species. Furthermore, the most widespread and also most common species in Norway, Mocyta fungi, was examined in more detail to test if any geographic patterns are reflected in the genetic variation, and if this variation correlates with ecological preferences. This study also aimed to explore if the parthenogenetic populations of M. fungi are restricted geographically. In total, 111 Mocyta specimens, representing 12 morphospecies from 17 countries, were included in the analyses. Both maximum likelihood and Bayesian inference were employed on two PCR amplified molecular markers (the mitochondrial cytochrome oxidase subunit 1 and the internal transcribed spacer 2 of the nuclear ribosomal gene cluster) to investigate the phylogenetic relationship of these morphospecies. In addition, calculations of mean intra- and interspecific genetic distances were used to delimit species. Online service for delimiting species using Bayesian implementation of the PTP model (bPTP) was also applied. Haplotype networks were used to investigate the genetic variation among specimens of M. fungi. The maximum likelihood and Bayesian inference analyses grouped the morphospecies together in well-supported clades, but the species delimitation methods applied failed to confirm this. The bPTP method estimated many species, exceeding the number of morphospecies, but the supports for these were generally low. There were no evident geographic patterns reflected in the genetic variation of M. fungi, nor any correlation between genetic variation and ecological preferences. Biased representation of specimens from different countries made it impossible to confidently compare the genetic variation between specimens from Norwegian populations and specimens from populations elsewhere. Even though all 33 specimens of M. fungi were females, the parthenogenetic populations of M. fungi could not be examined outside Norway because too few non- Norwegian specimens were included. V VI Table of Contents Introduction ........................................................................................................................... 1 Subfamily Aleocharinae Fleming, 1821 ............................................................................. 1 Tribe Athetini Casey, 1910................................................................................................. 1 Genus Mocyta Mulsant & Rey, 1874 ................................................................................. 2 Species Concepts ............................................................................................................... 3 Molecular Markers and Species Boundaries ..................................................................... 4 Purpose of this Study ......................................................................................................... 5 Material and Methods ........................................................................................................... 7 The Species of the Genus Mocyta ..................................................................................... 7 Collecting Methods and Strategy ...................................................................................... 7 Sorting Collected Samples ................................................................................................. 7 Identification ..................................................................................................................... 8 Specimen Selection............................................................................................................ 9 Specimen Preparation ....................................................................................................... 9 DNA Extraction ................................................................................................................ 10 Primer Testing and DNA Amplification ............................................................................ 10 Cleaning and Sequencing ................................................................................................. 11 Trimming and Assembling ............................................................................................... 11 Aligning ............................................................................................................................ 11 Analysing .......................................................................................................................... 12 Datasets ....................................................................................................................... 12 Neighbor-Joining Trees ................................................................................................ 12 Model Tests ................................................................................................................. 12 Maximum Likelihood ................................................................................................... 12 Bayesian Inference ...................................................................................................... 13 VII Intra- and Interspecific Genetic Variation ....................................................................... 14 Haplotype Networks ........................................................................................................ 14 Data Management ........................................................................................................... 15 Results ................................................................................................................................. 17 The Species of the Genus Mocyta ................................................................................... 17 Mocyta fungi (Gravenhorst, 1806) .............................................................................. 17 Mocyta amplicollis (Mulsant & Rey, 1873) .................................................................. 20 Mocyta amblystegii (Brundin, 1952) ........................................................................... 20 Mocyta orbata (Erichson, 1837) .................................................................................. 21 Mocyta orphana (Erichson, 1837) ............................................................................... 21 Mocyta negligens (Mulsant & Rey, 1873) ................................................................... 22 Mocyta gilvicollis (Scheerpeltz, 1949) ......................................................................... 23 Mocyta clientula (Erichson, 1839) ............................................................................... 23 Mocyta fussi (Bernhauer, 1908) .................................................................................. 25 Mocyta breviuscula (Mäklin in Mannerheim, 1852)
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