v PKylogenetic Studies w of *^ , . §outherrf African Branch -R. .x.A. Tolley M. Cunningham AyM. Bauer G. Alexander J.A. Harrison A.A. Turner M.F. Bates iiii< SANBI Biodiversi * Zootomy HyaauroofC'- l^iV»rr^ University i SANBI Biodiversity Series SANBI Biodiversity Series 5 A plan for phylogenetic studies of southern African reptiles Proceedings of a workshop held at Kirstenbosch, February 2006 Editors: W.R. Branch K.A. Tolley M. Cunningham A.M. Bauer G. Alexander J.A. Harrison A.A. Turner and M.R Bates national biodiversity institute SANBI Pretoria 2006 SANBI SANBI Biodiversity Series The South African National Biodiversity Institute (SANBI) was established on 1 Sep- tember 2004 through the signing into force of the National Environmental Manage- ment: Biodiversity Act (NEMBA) No.10 of 2004 by President Thabo Mbeki. The Act expands the mandate of the former National Botanical Institute to include respon- sibilities relating to the full diversity of South Africa's fauna and flora, and builds on the internationally respected programmes in conservation, research, education and visitor services developed by the National Botanical Institute and its predecessors over the past century. The vision of SANBI is to be the leading institution in biodiversity science in Africa, facilitating conservation, sustainable use of living resources, and human wellbeing. SANBI's mission is to promote the sustainable use, conservation, appreciation and enjoyment of the exceptionally rich biodiversity of South Africa, for the benefit of all people. SANBI Biodiversity Series will publish occasional reports on projects, technologies, workshops, symposia and other activitiesl(u=wtiated by or executed in partnership with SANBI. 3K Technical editor: Gillian de Jager Design & layout: Sandra Turck Cover design: Sandra Turck. Photographs provided by Marius Burger. Citation Branch, W.R., Tolley, K.A., Cunningham, M., Bauer, A.M., Alexander, G., Harrison, J.A., Turner, A.A. & Bates, M.F. (eds) 2006. A plan for phylogenet- ic studies of southern African reptiles: proceedings of a workshop held at Kirsten- bosch, February 2006. Biodiversity Series 5. South African National Biodiversity Institute, Pretoria. Branch, W.R. 2006. Priorities for systematic studies on southern African reptiles. Pp. 2-20 in: W.R. Branch, K.A. Tolley, M. Cunningham, A.M. Bauer, G. Alexan- der, J.A. Harrison, A.A. Turner & M.F. Bates (eds). A plan for phylogenetic studies of southern African reptiles: proceedings of a workshop held at Kirstenbosch, February 2006. Biodiversity Series 5. South African National Biodiversity Institute, Pretoria. ISBN 1-919976-33-7 © Published by: South African National Biodiversity Institute. Obtainable from: SANBI Bookshop, Private Bag XI01, Pretoria, 0001 South Africa. Tel.: +27 12 843-5000. E-mail: [email protected]. Website: www.sanbi.org. Printed by United Litho, 22 Eland Street, Koedoespoort, Pretoria, 0186 South Africa. Contents : Introduction J.A. Harrison CHAPTER 1 Priorities for systematic studies on southern African / reptiles W.R. Branch ^r\&Q CHAPTER 2 Taxonomic units relevant to conservation planning A.M. Bauer nr* CHAPTER 3 Mismatches between morphology and genetics 5^ G. Alexander CHAPTER 4 Methods, techniques and protocols for phylogenetic 34 studies on southern African reptiles K.A. Tolley, M. Cunningham and A.A. Turner CHAPTER 5 A sampling and implementation strategy for phylo- 40 genetic studies on southern African reptiles M. Cunningham, J.A. Harrison and K.A. Tolley 48 List °f workshop participants Introduction The Southern African Reptile Conservation Assessment (SARCA) was launched in May 2005. Its primaryaim isto produce a conservation assessmentforthe reptiles ofSouth Africa, Lesotho and Swaziland within a four-year period (2005-2009). It has the distinction of beingthe firstfaunal project ofthe newly constituted South African National Biodiversity Institute (SANBI) which, in itsformer incarnation asthe National Botanical Institute (NBI), was concerned onlywith plants. SARCA was motivated principally by a clear need for an update ofthe South African Red Listfor reptiles which was last updated in 1988. However, projects ofthis kind aimed at beingcomprehensive in geographical and taxonomic scope, are confronted with a discipline's knowledge gaps. These gaps are particularly acute in southern Africa, wheretaxonomic studies in the last 20 years have revealed thatthe subcontinent is a global hotspot of reptile diversity. Inadequacies in the geographical samplingof reptiles are being addressed by a series ofSARCAsurveys in previously undersampled areas. However, this effortalone cannotspecificallyaddressthe incomplete and sometimes problematic nature ofthe region's reptile alpha taxonomy (the description and naming ofspecies). As conservation assessmentand planningdepend fundamentallyon alpha taxonomy, itwas soon apparentthat SARCA would have to help initiate a programme to resolvethe pressingtaxonomic problems. To this end, a workshop was organised to identify, listand prioritise all known taxonom- ic problems. In addition, the workshop participants would discuss, resolve and describe the methodological questions and the practicalities of methods and resources. This workshop was held from 22-24 February 2006, atSANBI's Biodiversity Research Building, Kirstenbosch, Cape Town. The eventwasfunded bythe South African National Research Foundation (NRF), facilitated by Renee le Roux and hosted by SANBI, with the particular assistance of Krystal Tolley. The participants included Prof. Graham Alexander (Universityofthe Witwatersrand), Mike Bates (National Museum, Bloemfontein), Prof. Aaron Bauer (University ofVil- lanova, Pennsylvania), Dr Bill Branch (PE Museum), Marius Burger (ADU), Dr Michael Cunningham (University ofthe Free State), DrSavel Daniels (UniversityofStellenbosch), James Harrison (ADU), Prof. Margaretha Hofmeyr (University ofthe Western Cape), Johan Marais (University ofthe Witwatersrand), Prof. Le Fras Mouton (University of Stellenbosch), Dr Krystal Tolley (SANBI), AndrewTurner(CapeNature) and rapporteurs Daniel Goedbloed (University ofthe Free State) and Lerina Kaars (University ofthe Free State, SANBI intern). Additional inputwas given duringone session by DrJohn Donald- son (SANBI) and Prof. Les Underhill (ADU). So, with few exceptions, the main research- ers involved in thetaxonomy ofthe region's reptiles were present, and contributed substantivelytothe proceedings ofthe workshop and to this report. This report brings together, in a single document, a comprehensive set ofguidelinesfor a whole section ofsouthern Africa's biodiversity research, and should remain relevant forat least a decade. The herpetologists ofthe region congratulatethe institutions, especiallySANBI and NRF, that have had the vision and commitmentto supportthis endeavour. J.A. Harrison Avian Demography Unit, University of Cape Town, Rondebosch 7701 O SANBI Biodiversity Series 5 (2006) ^CHAPTER 1 Priorities for systematic studies on ^\^^^ southern African reptiles i&K$ and optimisethe taxonomic usefulness of W.R. Branch these surveys, and the subsequentanalysis of Bayworld (formerly Port Elizabeth Museum), voucher material, it is necessaryto reviewthe P.O. Box 13147, Humewood 6013, taxonomic knowledge ofthe regional reptile Port Elizabeth fauna and to identify and highlight problematic taxa. Southern Africa hasthe richest reptile diver- sity in Africa (Bauer 1993; Branch 1999a), Approach with a fauna that currently exceeds 520 A provisional list ofgenera in which cryptic species (Branch 1998; subsequent updates). taxa were known orsuspected to be present, Lizardsform the dominant component ofthis was prepared bythe author. Problematic taxa rich fauna due, in part, to an exceptional radia- included the following: tion ofgeckos in the western arid region. Over • 100 species ofgecko are now known from the • species with subspecies (races) that have subcontinent, while the families Scincidae and not been recently reviewed (seethe chap- Cordylidae are also well represented and the ter by Bauer in this volumefor a discussion amphisbaenid diversity isthe richest in Africa. ofspecies definitions and the subspecies Only one introduced reptile {Rhamphotyphlops concept in herpetology); braminus) has become established. • species with disjunct ranges and geo- graphically isolated populations, which may The current rate ofspecies description shows include cryptictaxa; and r < little indication of reachinga plateau (Branch • species with contiguous ranges, butwith 1999a), even after 250 years ofalmost con- confusing morphological (includingcoloura- tinuous study. This is evident in the increase tion) and/or habitatvariation. in species numbers in recent decades (397 in Branch 1988; 480 in Branch 1998; 520+ in These were discussed at a workshop at- Branch unpubl. checklist). In addition to its di- tended by invited researchers (see Introduc- versity, the southern African reptilefauna also tion for list). Following discussion, the listwas displays high endemicity, particularly in lizards amended for oversights and new insights, (mean 65.3%; Cordylidae 85.5%; Chamaeleo- and expanded to include details of proposed nidae 95%). This endemicity exceeds that of and ongoing research projects. Discussion frogs and freshwaterfish (50-60%) and is then prioritised the identified problem taxa much greaterthan that of birds and mammals for attention (1-5, low-high) and the research Mi (<25%). funding required to resolve these problems (1-5). Problem taxa scored high iftheywere A directed programme of reptile surveys is a known to contain numerous newtaxa, or had componentofthe Southern African Reptile the potential to do so, based on the biological Conservation Assessment (SARCA). The survey and distribution characteristics associated localities were selected by usinga gap analy- with high species richness. High scoresfor sis of known reptile distributions, based on research fundingwere based on a lack of museum records in the majorSouth African existingfundingand a reasonable chance museums. The gap analysis identified areas ofobtainingessential study material. For with known reptile diversities, which were the latter, the resolution ofsometaxonomic significantly lowerthan those predicted byan problems requires extralimital material that analysis ofthe distribution maps published is currently not readily available (e.g. from in Branch (1998). In addition to collecting Angola). Duetothe interrelatedness ofthe distribution data, the surveys mayserve a reptile fauna ofsouthern Africa, the analysis of useful ancillaryfunction by collecting material problematictaxa was not limited tothe SARCA fortaxonomic revisions, as well astissue for region (South Africa, Lesotho and Swaziland). use in molecularstudies. Moreover, theycan be expected to uncover additional taxonomic Results novelties, as well as populations which do not fit easily into current hypotheses ofspecies' Genera containing problematic taxa are sum- distributions and diagnoses. To both direct marised in Appendix 1. 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