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Revision, Molecular Phylogeny and Biology of the Spider Genus Micaria Westring, 1851 (Araneae: Gnaphosidae) in the Afrotropical Region

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Revision, Molecular Phylogeny and Biology of the Spider Genus Micaria Westring, 1851 (Araneae: Gnaphosidae) in the Afrotropical Region REVISION, MOLECULAR PHYLOGENY AND BIOLOGY OF THE SPIDER GENUS MICARIA WESTRING, 1851 (ARANEAE: GNAPHOSIDAE) IN THE AFROTROPICAL REGION by Ruan Booysen Submitted in fulfilment of the requirements for the degree MAGISTER SCIENTIAE in the Department of Zoology & Entomology, Faculty of Natural and Agricultural Sciences, University of the Free State February 2020 Supervisor: Prof. C.R. Haddad Co-Supervisor: Prof. S. Pekár 1 DECLARATION I, Ruan Booysen, declare that the Master’s research dissertation that I herewith submit at the University of the Free State, is my independent work and that I have not previously submitted it for qualification at another institution of higher education. 02.02.2020 _____________________ __________________ Ruan Booysen Date 2 Contents ABSTRACT ..................................................................................................................... 5 OPSOMMING ................................................................................................................. 7 ACKNOWLEDGEMENTS ............................................................................................... 9 CHAPTER 1 - INTRODUCTION ................................................................................... 10 1.1.) Micaria morphology ............................................................................................... 10 1.2.) Taxonomic history of Micaria................................................................................. 11 1.3.) Phylogenetic relationships .................................................................................... 12 1.4.) Mimicry .................................................................................................................. 13 1.5.) Reproductive biology ............................................................................................. 14 1.6.) Mating plugs .......................................................................................................... 15 1.7.) Study aims and objectives .................................................................................... 17 1.8.) Research Significance .......................................................................................... 18 1.9.) Literature Cited...................................................................................................... 19 CHAPTER 2 – REVISION OF THE AFROTROPICAL MICARIA .................................. 28 2.1.) Introduction ........................................................................................................... 28 2.2.) Materials and Methods .......................................................................................... 30 2.3.) Genus Micaria Westring, 1851 .............................................................................. 33 2.4.) Key to the Micaria species of the African part of the Afrotropical Region: ............. 51 2.5.) Taxonomy ............................................................................................................. 55 2.5.1.) Micaria basaliducta sp. nov. ............................................................................... 55 2.5.2.) Micaria beaufortia (Tucker, 1923) ...................................................................... 60 2.5.3.) Micaria bimaculata sp. nov. ................................................................................ 70 2.5.4.) Micaria bispicula sp. nov. ................................................................................... 75 2.5.5.) Micaria chrysis (Simon, 1910) ............................................................................ 81 2.5.6.) Micaria durbana sp. nov. .................................................................................... 92 2.5.7.) Micaria felix sp. nov. ........................................................................................... 97 2.5.8.) Micaria gagnoa sp. nov. ................................................................................... 106 2.5.9.) Micaria koingnaas sp. nov. ............................................................................... 111 2.5.10.) Micaria latia sp. nov. ...................................................................................... 115 3 2.5.11.) Micaria laxa sp. nov. ...................................................................................... 119 2.5.12.) Micaria medispina sp. nov. ............................................................................. 122 2.5.13.) Micaria parvotibialis sp. nov. .......................................................................... 126 2.5.14.) Micaria plana sp. nov. .................................................................................... 130 2.5.15.) Micaria quadrata sp. nov. ............................................................................... 135 2.5.16.) Micaria quinquemaculosa sp. nov. ................................................................. 139 2.5.17.) Micaria rivonosy sp. nov. ................................................................................ 145 2.5.18.) Micaria salta sp. nov. ..................................................................................... 149 2.5.19.) Micaria scutellata sp. nov. .............................................................................. 153 2.5.20.) Micaria tersissima Simon, 1910 ..................................................................... 157 2.6.) Discussion ........................................................................................................... 161 2.7.) Literature Cited.................................................................................................... 162 CHAPTER 3: SYSTEMATICS OF MICARIA ............................................................... 165 3.1.) Introduction ......................................................................................................... 165 3.2.) Materials and Methods ........................................................................................ 166 3.3.) Results ................................................................................................................ 167 3.4.) Discussion ........................................................................................................... 182 3.5.) Literature Cited.................................................................................................... 184 CHAPTER 4: BIOLOGY, BEHAVIOUR AND MYRMECOMORPHY OF MICARIA, WITH EMPHASIS ON THE AFROTROPICAL SPECIES ...................................................... 186 4.1.) Introduction ......................................................................................................... 186 4.2.) Materials and methods ........................................................................................ 187 4.3.) Results ................................................................................................................ 188 4.4) Discussion ............................................................................................................ 194 4.5.) Literature Cited.................................................................................................... 196 CHAPTER 5: CONCLUDING REMARKS ................................................................... 198 4 ABSTRACT The genus Micaria Westring, 1851 (Araneae, Gnaphosidae) is a group of small (1.85 - 5 mm) ant-like spiders that can be distinguished from other gnaphosids by their piriform gland spigots that are similar in size to the major ampullate gland spigots. According to the World Spider Catalog, there are 105 species of Micaria in the world, of which only four species are known from the Afrotropical Region, namely M. chrysis (Simon, 1910), M. tersissima Simon, 1910,M. beaufortia (Tucker, 1923) and M. ignea (O. Pickard- Cambridge, 1872). The objectives of this study were to revise Micaria in the Afrotropical Region, providing new and updated records for each of the species, evaluating the relationships between them using COI barcoding data, and providing information on their biology, mimetic relationships and feeding ecology. These objectives were met by collecting fresh material from the KwaZulu-Natal, Western Cape, Northern Cape, and Free State provinces in South Africa. Fresh material of M. tersissima and M. chrysis were collected from their type localities, Komaggas and Port Nolloth (Northern Cape Province), respectively, for identification and DNA analyses. Material from eight collections yielded a variety of species of Micaria from countries throughout the Afrotropical Region. Male and female genitalia were dissected and cleaned using a pancreatin solution. The left leg II of male and female representatives of each species was preserved in absolute ethanol and sent to the Canadian Centre for DNA Barcoding (CCDB) for DNA barcoding (COI gene). The data were aligned using Mega X software and molecular analyses were performed using MrBayes for Bayesian Inference (BI) and RaxML for maximum likelihood (ML) analyses. Morphological analysis of the collected and voucher material yielded 17 new species for the Afrotropical Region, namely M. basaliducta sp. nov., M. bimaculata sp. nov., M. bispicula sp. nov., M. durbana sp. nov., M. felix sp. nov., M. gagnoa sp. nov., M. koingaas sp. nov., M. latia sp. nov., M. laxa sp. nov., M. medispina sp. nov., M. parvotibialis sp. nov., M. plana sp. nov., M. quadrata sp. nov., M. quinquemaculosa sp. nov., M. rivo sp. nov., M. salta sp. nov. and M. scutellata sp. nov. The maximum likelihood
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