The Biogeography, Systematics and Conservation Of
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THE BIOGEOGRAPHY, SYSTEMATICS AND CONSERVATION OF PHREATOICIDEAN ISOPODS IN SOUTH AFRICA GAVIN GOUWS Promoter: Prof. B. A. COOK (Centre of Excellence in Natural Resource Management, University of Western Australia) Co-promoter: Dr. C. A. MATTHEE (Department of Zoology, University of Stellenbosch) Dissertation presented for the Degree of Doctor of Philosophy (Zoology) at the Department of Zoology, University of Stellenbosch, South Africa December 2004 Declaration I, the undersigned, hereby declare that the work contained in this dissertation is my own original work and has not previously in its entirety, or in part, been submitted at any university for a degree. All references and help received have been fully acknowledged. Signature:……………………………….. Date:…………………………………… i Abstract Historically, isopods of the suborder Phreatoicidea were thought to be represented in southern Africa by four species belonging to the endemic genus Mesamphisopus. This taxonomy was based on poor collections and the extent of variation among and within populations were poorly understood. In the present study, intensive sampling was undertaken to determine the diversity, distribution and biogeography of phreatoicidean isopods within South Africa. Analyses of allozyme data and mitochondrial DNA sequences (from the 12S rRNA and protein-coding COI genes) were used to examine differentiation among populations, extricate species boundaries (in combination with morphometric and morphological data) and to elucidate the evolutionary relationships among taxa. Additionally, conservation units were identified among the sampled populations and conservation threats highlighted. First, genetic and morphometric differentiation was examined among populations identified morphologically as M. capensis. Collection localities spanned two mountainous regions in the Western Cape and these were separated by a coastal plain remnant. Five morphometrically and genetically distinct species were identified. These taxa are also geographically partitioned in two regions, which were regarded as Evolutionarily Significant Units. Differentiation among populations of the two regions, and similar patterns in other taxa, was attributed to Cenozoic sea-level fluctuations. Second, populations, variably assigned to M. abbreviatus or M. depressus, were examined to determine whether they were conspecific. A large geographic area was sampled to account for intraspecific differentiation. Limited morphometric discrepancies were observed, with individual populations being either similar to the M. abbreviatus or the M. depressus syntypes. Genetic support for the recognition of a cryptic species complex among the sampled populations was equivocal. Substantial genetic differentiation and a lack of gene flow were observed among all populations. Clear patterns of isolation by distance were not detected, and genetic structure appeared to be unrelated to geography or drainage systems. The mosaic pattern of relatedness among populations was best explained by stochastic demographic processes, such as extinction-recolonization events or population bottlenecks. ii Thirdly, detailed taxonomic descriptions and illustrations of six new species, identified genetically and morphometrically among the populations included in the above analyses, were provided. These species were largely distinguished from each other, and the four original species, using a combination of setation, mouthpart, pleopod and uropod features. Lastly, phylogenetic relationships among all ten recognized Mesamphisopus species, and an additional unresolved group of populations, were examined. MtDNA data partitions and a recoded allele frequency matrix were analysed independently and in combination. Topologies indicated unrecognized species-diversity within an unresolved group of populations. Evolutionary relationships, the identification of six biogeographic centres, and the dating of divergences using a relaxed Bayesian clock suggested that differentiation and speciation within Mesamphisopus was largely allopatric or vicariant and driven by Mesozoic sea-level and climate change. Chance long distance dispersal events would, in turn, explain spurious phylogenetic relationships and distributions. This study contributes significantly to the understanding of the diversity and the conservation of the little-studied southern African freshwater invertebrates. Moreover, this study is the first to investigate genetic and morphometric differentiation, and phylogenetic relationships, below the generic level within the Phreatoicidea; thus establishing a methodological and theoretical framework for species delineation and the accurate determination of biodiversity within individual phreatoicidean genera. iii Uittreksel Isopoda van die suborder Phreatoicidea was histories in suidelike Afrika verteenwoordig deur vier spesies wat almal aan die endemiese genus Mesamphisopus behoort. Hierdie taksonomie is op ’n beperkte hoeveelheid versamelings gebaseer en die omvang van variasie tussen (en binne) bevolkings was swak verstaan. In die huidige studie is ekstensiewe versameling onderneem om die verspreiding, diversiteit asook biogeografie van dié Isopoda in Suid Afrika te bepaal. Analises van allosiem data en mitokondriale DNS volgorderbepalings (van die 12S rRNS en die proteïenkoderende COI geen) was gebruik om differensiasie tussen bevolkings te ondersoek, om (in kombinasie met morfometriese en morfologiese data) spesiesgrense te bepaal asook om die evolusionêre-verwantskappe tussen taksa te definieer. Benewens word bewaringseenhede binne die studie-bevolkings geïndentifiseer en moontlike bedreigings uitgelig. Eerstens is genetiese en morfometriese differensiasie tussen bevolkings, wat as M. capensis geïdentifiseer is, ondersoek. Versamelingslokaliteite was versprei oor twee bergagtige streke in die Weskaap wat geskei word deur ’n voormalige kusvlakte. Vyf morfometries- en geneties-afsonderlike spesies is geïdentifiseer. Dié taksa was geografies geskei tot die twee streke, wat elk as ’n Evolusionêre Beduidende Eenheid (ESU) gesien kan word. Differensiasie tussen populasies van die twee streke en vergelykbare patrone binne ander taksa word aan Cenosoïese seevlak veranderings toegeskryf. Tweedens is bevolkings wat as M. abbreviatus óf as M. depressus geïdentifiseer kan word ondersoek om te bepaal of hulle konspesifiek is. Bevolkings is oor ’n groot geografiese gebied versamel om intraspesifieke variasie in aanmerking te neem. Beperkte morfometriese verskille is waargeneem – enkel bevolkings was morfometries identies aan of die M. abbreviatus of die M. depressus sintipes. Genetiese getuienis vir die herkenning van ’n kriptiese spesieskompleks was dubbelsinnig. Bevolkings is gekenmerk deur merkbare genetiese differensiasie en die afwesigheid van geenvloei. Duidelike bewys van isolasie-met- afstand was nie waargeneem nie en genetiese struktuur was nie verwant aan geografiese ligging of riviersisteme nie. Die mosaïese patroon van verwantskappe is moontlik teweeg gebring deur stogastiese demografiese prosesse soos uitsterwing en hervestiging of deur afnames in bevolkingsgrootte. iv Derdens is omvattende taksnomiese beskrywings en illustrasies van ses nuwe spesies wat deur bogenoemde analises geneties en morfometries uitgelig was, verskaf. Dié spesies is van mekaar, asook die ander vier spesies onderskeibaar deur ’n kombinasie van setasie-, monddeel-, pleiopoot- en uropooteienskappe. Laastens is die filogenetiese verwantskappe tussen al tien herkende Mesamphisopus-spesies en ’n groep bevolkings waarvan verhoudings onseker was, ondersoek. MtDNS datastelle en ’n hergekodeerde alleelfrekwensie matriks is afsonderlike en in kombinasie geanaliseer. Topologië het onherkende spesies-vlak diversiteit binne die bogenoemde groep bevolkings aangedui. Evolusionêre verwantskappe, die herkenning van ses biogeografiese gebiede, en die bepaling van tye van divergensie (d.m.v. ’n ontspanne Bayesiaanse molekulêre klok) het aangetoon dat spesiasie binne Mesamphisopus grootliks allopatries was en deur Mesosoïese seevlak- en klimaatsveranderings teweeg gebring is. Toevallige lang-aftstand verspreiding kon dan eienaardige filogenetiese verhoudings en verspreidings verklaar. Dié studie lewer ’n wesenlike bydrae tot die kennis van die diversiteit en tot die bewaring van die onbestudeerde Suid Afrikaanse varswater ongewerweldes. Daarenbowe, is hierdie studie die eerste om genetiese en morfometriese differensiasie benede die genusvlak binne die Phreatoicidea te ondersoek; sodoende word die metodologiese en teoretiese raamwerk vir die herkenning van spesies en die akkurate beskrywing van diversiteit binne afsonderlike genera van die Phreatoicidea geskep. v Acknowledgements First, and foremost, I extend my thanks and gratitude towards my supervisors and mentors, Prof. Barbara Cook and Dr. Conrad Matthee. They both provided immeasurable intellectual input, through helpful advice, insights, suggestions and discussion. Perhaps more importantly, they both provided enthusiastic, unwavering support and understanding. Barbara and Conrad deserve additional thanks for, respectively, being willing to continue providing supervision from abroad, and for being willing to stand in and take another student on board. The National Research Foundation (South Africa) and the University of Stellenbosch are thanked for providing funding, without which this study would not have been completed. The National Research Foundation, University of Stellenbosch, Department of Zoology and Harry Crossley Foundation are