Correspondence

Island crickets (Mogoplistinae: Orne- geological (and evolutionary) time-scales, pre-human clade age, and one additional bius). PLoS ONE, 11, e0148971. and eddies and counter-currents may event is considered unconfirmed. Wilme, L., Waeber, P.O. & Ganzhorn, J.U. facilitate transport against the main cur- We find that 42 of the 48 WIO long-dis- (2016a) Human translocation as an rent direction. We review the cases of the tance dispersal events (87.5%) followed the alternative hypothesis to explain the extant and extinct WIO giant prevailing marine current, whereas the direc- presence of giant tortoises on remote and suggest that the current distribution tion of only three (6.3%) is contrary to the in the south-western Indian of all lineages can be convincingly current. Three events with unknown region . Journal of Biogeography. explained by overseas dispersal. of origin are omitted. Furthermore, our list Wilme, L., Waeber, P.O. & Ganzhorn, J.U. shows 42 confirmed short-distance dispersal (2016b) Marine used to assist Keywords , biogeography, events within . We thus count a Austronesian sailors reaching new , giant , , minimum number of 90 successful dispersal islands. Comptes rendus biologies, 339, marine currents, Mascarenes, overseas dis- events across long and short distances that is 78–82. persal, , Western reflected in the known extant and extinct diversity of WIO terrestrial .

SUPPORTING INFORMATION MULTIPLE OVERSEAS A CLOSER LOOK AT MARINE Additional Supporting Information may be DISPERSAL EVENTS OF CURRENTS found in the online version of this article: TERRESTRIAL REPTILES With few exceptions, terrestrial organisms Appendix S1 Additional methodological Phylogenetic studies of the last decades totally depend on surface currents and information for Fig. 1 and Video S1. have revealed that overseas dispersal is winds for passive transport while drifting Video S1 Surface current dynamics in the much more common than expected earlier, at sea, and many biogeographical studies Indian Ocean 2007–2008. and that it has occurred in a wide range of discuss overseas dispersal along marine Video S2 , Aldabra- geological periods (de Queiroz, 2005). The currents versus ‘counter-current’ dispersal chelys gigantea, swimming across a pond islands of the Western Indian Ocean as a factor of similar or higher importance on Aldabra , Seychelles. (WIO) region are a global hotspot of over- than distance (de Queiroz, 2005; Nathan seas dispersal, and a rich literature on the et al., 2008; Townsend et al., 2011). We topic has been produced. Crottini et al. created a map of oceanic surface currents Editor: Sonya Clegg (2012) reviewed the complex dispersal his- (Fig. 1) which shows that the westward tory of biota from to Madagascar, supports drifting doi:10.1111/jbi.12893 and Vences et al. (2003) analysed and from north Madagascar to the submerged reviewed the overseas dispersal of amphib- islands between Madagascar and the ians, which is relatively common in the (the Geyser and Leven Banks), region. We restrict our review to studies Aldabra, Glorioso and the Comoros. relevant to terrestrial reptiles of the WIO Further north along the African coast the region except Madagascar, listed in current turns back eastward near the equa- Table 1. Native terrestrial reptiles inhabit tor. Overseas dispersal from Madagascar to How marine currents the Mascarenes (, and the Mascarenes would be ‘counter-current’ influenced the widespread Reunion Island), the Seychelles including and therefore be seen as unlikely, if not natural overseas dispersal of Aldabra, the Comoros and several smaller impossible. reptiles in the Western Indian islands (Fig. 1). Existing literature identi- However, the marine circulation of the Ocean region fies continental Africa, Madagascar and the WIO is more complex than indicated by Eastern Indian Ocean region as major the main currents alone. The region of the source areas for the colonization of the Mascarene Basin between Madagascar and ABSTRACT WIO. This yields a range of minimal the Mascarenes is dominated by eddies and gyres potentially allowing transport in In a recent contribution to this journal, straight-line distances for dispersal in the > > various directions. The South Equatorial Wilme et al. (2016) proposed that the region from 130 to 5000 km, but true Countercurrent varies seasonally, favouring giant tortoises of the islands of the Wes- travelling distances may be much longer. In dispersal from Africa (and Madagascar) to Indian Ocean (WIO: Aldabra, the addition to these long-distance dispersal the Seychelles only in the winter. Moreover, Mascarenes, and the ) events, overseas dispersal also happened Carton & Giese (2008) and Schott et al. might have originated from translocation within archipelagos across shorter distances. (2009) detected an ‘Eastward Flow’ from by early Austronesian sailors. Prompted Under these criteria we find 35 long-distance the region south-east of Madagascar at by this paper we review recent literature dispersal events that are clearly supported by c. 25 ° south with a northward component. and show that natural overseas dispersal evidence from genetic studies, that is, diver- The occurrence of this flow means that dis- was remarkably widespread in the colo- gent allopatric clades that are clearly old persal from Madagascar to Mascarenes may nization history of terrestrial reptiles in enough to exclude the possibility of human not always be ‘counter-current’. the WIO region. Almost 90% of the suc- translocation. Further 12 events are inferred Furthermore, events such as glaciations, cessful colonization events are supported from currently recognized based changes in freshwater influx, tectonic activity by prevailing marine surface currents. on morphological data and dated subfossils, and volcanism may change the oceanic However, these currents may change over suggesting deep genetic divergence and

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Table 1 Cases of natural long-distance overseas dispersals of reptiles in the Western Indian Ocean region. Unless otherwise marked, the origin of natural overseas dispersal was inferred from genetic data. The region of origin of the ancestors, the current distribution and the shortest straight-line distance between these regions today are given. The column ‘Along current?’ informs whether the direction of a dispersal event follows the prevailing present-day oceanic currents or not. ‘SDDE’ denotes the minimum number of Short-Distance Dispersal Events (i.e. between islands of an ) following the long-distance dispersal event that are required to explain the observed biogeographical situation. This number is always zero for the granitic Seychelles because these islands were connected during periods of low sea levels. (†) refers to extinct or populations. * Overseas dispersal inferred from phylogenetic relationships based on morphological data from , subfossils, or museum specimens; no genetic data available. ** No specific age estimates are available for these taxa from the Seychelles (but see Gamble et al., 2012). Some of them may be old enough to pre-date the break-off of the Seychelles micro-continent and may have originated from vicariance rather than dispersal. *** Unconfirmed case. **** Cryptoblepharus spp. have a particularly complex biogeographical history in the WIO. After an initial dispersal from the Eastern Indian Ocean region, almost all islands of the WIO were colonized and are now inhabited by endemic clades. This case therefore comprises several long-distance dispersal events. Some relationships are confirmed by genetic data, while others are inferred through morphological studies.

Distance Along Taxon name Origin Distribution [km] current? SDDE Reference

Aldabrachelys gigantea Madagascar Aldabra 410 Yes - Austin et al. (2003) Oplurus sp.* Madagascar Aldabra (†) 410 Yes - Arnold (1976) Geckolepis sp.* Madagascar Aldabra (†) 410 Yes - Arnold (1976) Amphiglossus sp.* Madagascar Aldabra (†) 410 Yes - Arnold (1976) Trachylepis sp.* Madagascar or Aldabra (†) > 360 Yes - Arnold (1976) Comoros Paroedura sp.* Madagascar or Aldabra (†) > 360 Yes - Arnold (1976) Comoros sp.* Origin unknown Aldabra (†) 410? Yes? - Arnold (1976) (Madagascar?) Aldabrachampsus dilophus* Origin unknown Aldabra (†) 410? Yes? - Brochu (2006) (Madagascar?) Cylindraspis spp. Origin unknown Mascarenes (†) 700? No? 2 Austin & Arnold (2001) (Madagascar?) Gongylomorphus spp. Madagascar? Mascarenes (partly †) 700 No? 1* Austin & Arnold (2006) Bolyeriidae SE-? Mascarenes (partly †) > 4300? Yes? - Reynolds et al. (2014) Phelsuma spp. Madagascar Mascarenes (partly †) 700 No 3 Austin et al. (2004); Rocha et al. (2009) Nactus spp.* Eastern Indian Ocean Mascarenes (partly †) > 5000 Yes 2 Arnold & Bour (2008) Leiolopisma spp. Eastern Indian Ocean Mascarenes (partly †) > 5000 Yes 1 Austin & Arnold (2006) Archaius tigris Africa Seychelles > 1200 Yes - Townsend et al. (2011) Lycognathophis seychellensis** Africa Seychelles > 1200 Yes - Vidal et al. (2008); Pyron et al. (2013) Urocotyledon spp.** Africa Seychelles > 1200 Yes - Rocha et al. (2011), Gamble et al. (2012) Trachylepis spp. Africa Seychelles > 1200 Yes - Lima et al. (2013) Ailuronyx spp.** Madagascar or Africa Seychelles > 1200 Yes - Gamble et al. (2012), Pyron et al. (2013) Lamprophis geometricus** Africa Seychelles > 1200 Yes - Kelly et al. (2011) Pamelaescincus + Origin unknown Seychelles > 1200? Unknown - Austin & Arnold (2006) Janetaescincus** Phelsuma spp. Madagascar Seychelles > 1000 Yes - Rocha et al. (2009) Crocodylus porosus* SE-Asia? Seychelles (†) > 3000? Unknown - Gerlach & Canning (1994) Geckolepis humbloti Madagascar Comoros 290 Yes 3 Hawlitschek et al. (2016) Lycodryas spp. Madagascar Comoros 290 Yes 3 Hawlitschek et al. (2012) Trachylepis comorensis Africa Comoros 290 No 3 Rocha et al. (2010) Paroedura sanctijohannis Madagascar Comoros 290 Yes 2 Hawlitschek & Glaw (2013) Ebenavia cf. inunguis Madagascar Comoros 290 Yes 2 Hawlitschek et al. (in press) Phelsuma spp. (v-nigra Madagascar Comoros 290 Yes 3 Rocha et al. (2007) complex) Amphiglossus johannae Madagascar Comoros 290 Yes 3 Hawlitschek et al. (2013) Madatyphlops spp. Madagascar Comoros 290 Yes 2 Hawlitschek et al. (comorensis complex) (unpublished) Furcifer polleni Madagascar (Comoros) 290 Yes - Rocha et al. (2005); Tolley et al. (2013)

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Table 1 Continued

Distance Along Taxon name Origin Distribution [km] current? SDDE Reference

Liophidium mayottensis Madagascar Mayotte (Comoros) 290 Yes - Hawlitschek et al. (2012) Paroedura stellata Madagascar Mayotte (Comoros) 290 Yes - Hawlitschek & Glaw (2013) Ebenavia cf. inunguis Madagascar Mayotte (Comoros) 290 Yes - Hawlitschek et al. (in press) Phelsuma nigristriata Madagascar Mayotte (Comoros) 290 Yes - Rocha et al. (2009) Furcifer cephalolepis Madagascar Grand Comoro 520 Yes - Rocha et al. (2005); Tolley (Comoros) et al. (2013) Oplurus cuvieri comorensis* Madagascar Grand Comoro 520 Yes - Munchenberg€ et al. (2008) (Comoros) Madatyphlops (?) sp.*** Origin unknown Grand Comoro 520 Yes - Hawlitschek et al. (Madagascar?) (Comoros) (unpublished) Phelsuma comorensis Madagascar Grand Comoro 520 Yes - Rocha et al. (2007, 2009) (Comoros) Ebenavia cf. inunguis Madagascar Pemba Island > 1200 Yes - Hawlitschek et al. (in press) (Zanzibar Arch.) Phelsuma parkeri Madagascar Pemba Island > 1200 Yes - Rocha et al. (2009) (Zanzibar Arch.) Amphiglossus valhallae* Madagascar or Glorioso Island 180 Yes - Brygoo (1983) Comoros Zonosaurus madagascariensis Madagascar Glorioso + > 180 Yes - Brygoo (1985) insulanus* Islands Aldabrachelys sp.* Madagascar Glorioso Island (†) 180 Yes - Bour (1994) Phelsuma andamanense WIO Andaman Islands > 5000 Yes - Austin et al. (2004); Rocha et al. (2009) Lygodactylus insularis* Madagascar or Africa Juan de Nova Island > 130 Yes - Boettger (1913) Cryptoblepharus spp.**** Eastern Indian Ocean WIO > 5000 Yes > 10 Rocha et al. (2006); Horner (2007)

circulation regimes on geological time-scales much older than the extant islands of this of WIO giant tortoises with the idea that (Rahmstorf, 2002; Berne et al.,2004;Stanford group (Bradler et al., 2015). founder effects may enable the rapid evo- et al., 2011). Changes in global sea levels lution of morphological traits, such as occurred as consequences of the size and carapace shape. This may have climate oscillations and also earlier, back to DISTRIBUTION AND ORIGIN OF led scientists to recognize the various tor- c. 3Ma(Rohlinget al., 2014). In the WIO, GIANT TORTOISES IN THE WIO toise populations, including the morpho- these changes led to the emergence of the REGION logically clearly distinct species of Seychelles-Mascarene Ridge (SMR), either as Cylindraspis, as different species. However, a micro-continent of up to 130,000 km² sur- Globally, native populations of giant tor- the genetic divergences, at least among face or a chain of large islands, depending toises have only survived on the Galapagos the various Cylindraspis species from the on the extent of sea level regression (Braith- Islands and Aldabra. In the Pleistocene, Mascarenes, indicate clade ages far older waite, 1984; Warren et al., 2010). The SMR however, giant tortoises were much more than any human seafaring activity (Austin may have influenced strength and direction widespread on islands and continents, and & Arnold, 2001). Furthermore, the of the South Equatorial current as Madagas- extinct populations are known from the assumption of Wilme et al. (2016) is con- car does today (Backeberg & Reason, 2010; Canaries and some islands (Auf- tradicted by well-studied cases of the Badal & Rughooputh, 2010). This can be fenberg, 1974; Hansen et al., 2010). We see human-mediated translocation of other observed in a similar fashion in the west- the common occurrence of giant tortoises chelonians to WIO islands ( ward current of the Indo-Malayan region on oceanic islands as an indication of their zombensis to Madagascar, Kindler et al., (Gordon, 2005), where islands create eddies comparably good dispersal ability. They 2012; Pelomedusa subrufa to Madagascar, and deflect currents from their original are capable of tolerating the absence of Vargas-Ramırez et al., 2010; cas- direction. In addition to possible modifica- freshwater and food for weeks while their tanoides and Pelusios subniger to Madagas- tion of the currents, the enlargement of robust carapace and skin protect them car and the Seychelles, Fritz et al., 2013). the terrestrial surface made the SMR a from the tropical sun and saltwater, and In all these cases the translocated popula- much easier ‘target’ for colonization. Many they have also repeatedly been observed tions are morphologically and genetically organisms surviving on the Seychelles or swimming in the open sea (Gerlach et al., almost identical to their ancestral popula- Mascarenes may have previously colonized 2006). tions. We discuss the giant tortoise popu- the SMR, as has been shown for stick Wilme et al. (2016) justified their lations of the WIO islands, namely of the Mascarenes that may be hypothesis of the human-mediated origin Aldabra, the Mascarenes, Glorioso Island

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Figure 1 A simplified map of oceanic surface currents of the Western Indian Ocean. Light grey zones mark areas presumed to be terrestrial surface during sea level regressions. Arrows indicate surface current directions, circles represent eddies (water vortices). The dotted line between Aldabra and the Comoros marks a region with increased circulation speed. AC = , EACC = East African Coastal Current, MC = Current, NEMC = North-east , SEC = South Equatorial Current, SECC = South Equatorial Countercurrent, SEMC = South-. The SECC runs north of the equator in summer and south (closer to the Seychelles) in winter. ‘EF’ = Eastward Flow according to Carton & Giese (2008) and Schott et al. (2009). The map was created in QGIS 2.8.1 based on data and information from Carton & Giese (2008), Schott et al. (2009), and Ali & Huber (2010), and from http://earth.nullschool.net and http://www.esr.org/oscar_index.html. The true conditions of today’s marine currents are more complicated and can be viewed in the ‘marine currents’ section of http://earth.nullschool.net. and the Comoros, and the Granitic Sey- setting of marine currents may be de Biologia Evolutiva (CSIC-UPF), chelles, individually (see Appendix S1 in explained when taking into account Barcelona, Spain, 3Institut de Ciencies del Supporting Information) and find that regional patterns of marine currents and Mar (CSIC-ICM), Barcelona, Spain natural overseas dispersal is generally the their shifts over seasonal and geological *Correspondence Oliver Hawlitschek, Institut most convincing explanation of their phy- time-scales. de Biologia Evolutiva (CSIC-UPF), Passeig logeographical situation. Marıtim de la Barceloneta 37, 08003 Barcelona, Spain. E-mail: oliver. ACKNOWLEDGEMENTS [email protected] CONCLUSIONS We thank the editor S.C., the chief editor We present ample evidence that natural P.L. and two anonymous referees for their REFERENCES overseas dispersal of reptiles, including immensely helpful comments. Ali, J.R. & Huber, M. (2010) Mammalian giant tortoises, was a relatively common Oliver Hawlitschek1,2,*, on Madagascar controlled phenomenon in the WIO and serves to Sergio Ramırez Garrido 3 and by ocean currents. Nature, 463, 653– explain the origin of most tortoise popu- Frank Glaw1 lations in the region. Dispersal events 656. which may seem improbable or even 1Zoologische Staatssammlung Munchen€ Arnold, E.N. (1976) reptiles from impossible against the global present-day (ZSM-SNSB), Munich, Germany, 2Institut Aldabra Atoll, Indian Ocean. Bulletin of

1438 Journal of Biogeography 44, 1426–1440 ª 2016 John Wiley & Sons Ltd Correspondence

the British Museum of Natural History, Verlagsbuchhandlung, N€agele und Gerlach, J., Muir, C. & Richmond, M.D. 29,83–116. Sproesser, Stuttgart. (2006) The first substantiated case of Arnold, E.N. & Bour, R. (2008) A new Bour, R. (1994) L’etude des animaux dou- trans-oceanic tortoise dispersal. Journal Nactus () and a new blement disparus: les tortues geantes of Natural History, 40, 2403–2408. Leiolopisma skink (Scincidae) from La subfossiles de Madagascar. Memoires et Gordon, A.L. (2005) The of Reunion, Indian Ocean, based on recent Travaux de l’Institut de Montpellier, 19, the Indonesian Seas and their through- fossil remains and ancient DNA 1–253. flow. Oceanography, 18,14–27. sequence. Zootaxa, 1705,40–50. Bradler, S., Cliquennois, N. & Buckley, Hansen, D.M., Donlan, C.J., Griffiths, C.J. Auffenberg, W. (1974) Checklist of fossil T.R. (2015) Single origin of the Mas- & Campbell, K.J. (2010) Ecological his- tortoises (Testudinidae). Bulletin of the carene stick insects: ancient radiation on tory and latent conservation potential: State Museum, 18, 121–251. sunken islands? BMC Evolutionary Biol- large and giant tortoises as a model for Austin, J.J. & Arnold, E.N. (2001) Ancient ogy, 15, 196. taxon substitutions. Ecography, 33, 272– mitochondrial DNA and morphology Braithwaite, C.J.R. (1984) Geology of the 284. elucidate an extinct island radiation of Seychelles. Biogeography and ecology of Hawlitschek, O. & Glaw, F. (2013) The Indian Ocean giant tortoises (Cylin- the Seychelles Islands (ed. by D.R. Stod- complex colonization history of noctur- draspis). Proceedings of the Royal Society dart), pp. 17–38. The Hague, Boston. nal (Paroedura) in the Comoros B: Biological Sciences, 268, 2515–2523. Brochu, C.A. (2006) A new miniature Archipelago. Zoologica Scripta, 42, 135– Austin, J.J. & Arnold, E.N. (2006) Using horned crocodile from the Quaternary 150. ancient and recent DNA to explore rela- of Aldabra Atoll, Western Indian Ocean. Hawlitschek, O., Nagy, Z.T. & Glaw, F. tionships of extinct and endangered Copeia, 2006, 149–158. (2012) Island evolution and systematic Leiolopisma skinks (Reptilia: Scincidae) Brygoo, E. (1983) Systematique des lezards revision of Comoran : why and in the . Molecular Phy- scincides de la region malgache. X. Rap- when still make sense. PLoS logenetics and Evolution, 39, 503–511. ports de Gongylus johannae Gunther€ ONE, 7, e42970. Austin, J.J., Arnold, E.N. & Bour, R. 1880, des Comores, et de Sepsina valhal- Hawlitschek, O., Nagy, Z.T., Berger, J. & (2003) Was there a second adaptive lae Boulenger 1909, des Glorieuses, avec Glaw, F. (2013) Reliable DNA barcoding radiation of giant tortoises in the Indian les especes malgaches. Bulletin du performance proved for species and Ocean? Using mitochondrial DNA to Museum National d’Histoire Naturelle island populations of Comoran squa- investigate speciation and biogeography Paris, 5, 651–660. mate reptiles. PLoS ONE, 8, e73368. of Aldabrachelys (Reptilia, Testudinidae). Brygoo, E. (1985) Les Gerrhosaurinae de Hawlitschek, O., Scherz, M.D., Straube, N. Molecular Ecology, 12, 1415–1424. Madagascar, Sauria (Cordylidae). & Glaw, F. (2016) Resurrection of the Austin, J.J., Arnold, E.N. & Jones, C.G. Memoires du Museum National d’Histoire Comoran fish scale gecko Geckolepis (2004) Reconstructing an island radia- Naturelle ser. A, 134, Paris, 1–65. humbloti Vaillant, 1887 reveals a disjunct tion using ancient and recent DNA: the Carton, J.A. & Giese, B.S. (2008) A reanal- distribution caused by natural overseas extinct and living day geckos (Phelsuma) ysis of ocean climate using simple ocean dispersal. Organisms Diversity & Evolu- of the Mascarene islands. Molecular Phy- data assimilation (SODA). Monthly tion, 16, 289–298. logenetics and Evolution, 31, 109–122. Weather Review, 136, 2999–3017. Hawlitschek, O., Toussaint, E.F.A.T., Gehr- Backeberg, B.C. & Reason, C.J.C. (2010) A Crottini, A., Madsen, O., Poux, C., Strauss, ing, P.S., Ratsoavina, F.M., Cole, N., connection between the South Equato- A., Vieites, D.R. & Vences, M. (2012) Crottini, A., Nopper, J., Lam, A.W., rial Current north of Madagascar and Vertebrate time- elucidates the bio- Vences, M. & Glaw, F. (in press) Gecko Eddies. Geophysi- geographic pattern of a major biotic phylogeography in the Western Indian cal Research Letters, 37, L04604. change around the K-T boundary in Ocean region: the oldest clade of Ebe- Badal, M. & Rughooputh, S. (2010) Madagascar. Proceedings of the National navia inunguis lives on the youngest formation around South West Mas- Academy of Sciences USA, 109, 5358– island. Journal of Biogeography. carene Plateau (Indian Ocean) as evi- 5363. Horner, P. (2007) Systematics of the denced by satellite “global ocean colour” Fritz, U., Branch, W.R., Gehring, P.S., -eyed skinks, Cryptoblepharus data. Western Indian Ocean Journal of Harvey, J., Kindler, C., Meyer, L., Du Wiegmann (Reptilia: : Scinci- Marine Sciences, 8, 139–145. Preez, L., Siroky, P., Vieites, D.R. & dae) – an Australian-based review. The Berne, S., Rabineau, M., Flores, J.A. & Vences, M. (2013) Weak divergence Beagle, 3,21–198. Sierro, F. (2004) The impact of quater- among African, Malagasy and Seychellois Kelly, C.M.R., Branch, W.R., Broadley, nary global changes on strata formation: hinged terrapins (Pelusios castanoides, P. D.G., Barker, N.P. & Villet, M.H. (2011) exploration of the shelf edge in the subniger) and evidence for human- Molecular systematics of the African Northwest . Oceanog- mediated oversea dispersal. Organisms snake family Lamprophiidae Fitzinger, raphy, 17,92–103. Diversity and Evolution, 13, 215–224. 1843 (Serpentes: Elapoidea), with partic- Boettger, O. (1913) Reptilien und Amphi- Gamble, T., Greenbaum, E., Jackman, T.R., ular focus on the genera Lamprophis Fit- bien von Madagascar, den Inseln und Russell, A.P. & Bauer, A.M. (2012) zinger 1843 and Mehelya Csiki 1903. dem Festland Ostafrikas. Reise in Osta- Repeated origin and loss of adhesive toe- and Evolution, frika in den Jahren 1903-1905. Wis- pads in geckos. PLoS ONE, 7, e39429. 58, 415–426. senschaftliche Ergebnisse. Vol. 3. Gerlach, J. & Canning, K.L. (1994) On the Kindler, C., Branch, W.R., Hofmeyr, M.D., Systematische Arbeiten (ed. by A. Voeltz- crocodiles of the Western Indian Ocean. Maran, J., Siroky, P., Vences, M., Har- kow), pp. 269–375. Schweizerbartsche Phelsuma, 2,56–60. vey, J., Hauswaldt, J.S., Schleicher, A.,

Journal of Biogeography 44, 1426–1440 1439 ª 2016 John Wiley & Sons Ltd Correspondence

Stuckas, H. & Fritz, U. (2012) Molecular Rocha, S., Posada, D., Carretero, M.A. & Vargas-Ramırez, M., Vences, M., Branch, phylogeny of African hinge-back tor- Harris, D.J. (2007) Phylogenetic affinities W.R., Daniels, S.R., Glaw, F., Hofmeyr, toises (Kinixys): implications for phylo- of Comoroan and East African day M.D., Kuchling, G., Maran, J., Papen- geography and taxonomy (Testudines: geckos ( Phelsuma): multiple natu- fuss, T.J., Siroky, P., Vieites, D.R. & Testudinidae). Journal of Zoological Sys- ral colonisations, introductions and Fritz, U. (2010) Deep genealogical lin- tematics and Evolutionary Research, 50, island radiations. Molecular Phylogenetics eages in the widely distributed African 192–201. and Evolution, 43, 685–692. helmeted terrapin: evidence from mito- Lima, A., Harris, D.J., Rocha, S., Miralles, Rocha, S., Vences, M., Glaw, F., Posada, D. chondrial and nuclear DNA (Testudines: A., Glaw, F. & Vences, M. (2013) Phylo- & Harris, D.J. (2009) Multigene phy- : Pelomedusa subrufa). genetic relationships of Trachylepis skink logeny of Malagasy day geckos of the Molecular Phylogenetics and Evolution, species from Madagascar and the Sey- genus Phelsuma. Molecular Phylogenetics 56, 428–440. chelles (Squamata: Scincidae). Molecular and Evolution, 52, 530–537. Vences, M., Vieites, D.R., Glaw, F., Brink- Phylogenetics and Evolution, 67, 615–620. Rocha, S., Carretero, M.A. & Harris, D.J. mann, H., Kosuch, J., Veith, M. & Munchenberg,€ T., Wollenberg, K., Glaw, F. (2010) Genetic diversity and phyloge- Meyer, A. (2003) Multiple overseas dis- & Vences, M. (2008) Molecular phy- netic relationships of Mabuya spp. persal in amphibians. Proceedings of the logeny and geographic variation of (Squamata: Scincidae) from western Royal Society B: Biological Sciences, 270, Malagasy iguanas (Oplurus and Chalaro- Indian Ocean islands. Amphibia-Reptilia, 2435–2442. don). Amphibia-Reptilia, 29, 319–327. 31, 375–385. Vidal, N., Branch, W.R., Pauwels, O.S.G., Nathan, R., Schurr, F.M., Spiegel, O., Stei- Rocha, S., Harris, D.J. & Posada, D. (2011) Hedges, S.B., Broadley, D.G., Wink, M., nitz, O., Trakhtenbrot, A. & Tsoar, A. Cryptic diversity within the endemic Cruaud, C., Joger, U. & Nagy, Z.T. (2008) Mechanisms of long-distance prehensile-tailed gecko Urocotyledon (2008) Dissecting the major African dispersal. Trends in Ecology and inexpectata across the Seychelles Islands: snake radiation: a molecular phylogeny Evolution, 23, 638–647. patterns of phylogeographical structure of the Lamprophiidae Fitzinger (Serpen- Pyron, R.A., Burbrink, F.T. & Wiens, J.J. and isolation at the multilocus level. tes, Caenophidia). Zootaxa, 66,51–66. (2013) A phylogeny and revised classifi- Biological Journal of the Linnean Society, Warren, B.H., Strasberg, D., Bruggemann, cation of Squamata, including 4161 spe- 104, 177–191. J.H., Prys-Jones, R.P. & Thebaud, C. cies of lizards and snakes. BMC Rohling, E.J., Foster, G.L., Grant, K.M., (2010) Why does the biota of the Mada- Evolutionary Biology, 13, 93. Marino, G., Roberts, A.P., Tamisiea, gascar region have such a strong Asiatic de Queiroz, A. (2005) The resurrection of M.E. & Williams, F. (2014) Sea-level and flavour? Cladistics, 26, 526–538. in historical biogeogra- deep-sea-temperature variability over Wilme, L., Waeber, P.O. & Ganzhorn, J.U. phy. Trends in Ecology and Evolution, 20, the past 5.3 million years. Nature, 508, (2016) Human translocation as an alter- 68–73. 477–482. native hypothesis to explain the presence Rahmstorf, S. (2002) Ocean circulation Schott, F.A., Xie, S.-P. & McCreary, J.P.J. of giant tortoises on remote islands in and climate during the past 120,000 (2009) Indian Ocean circulation and cli- the south-western Indian Ocean. Journal years. Nature, 419, 207–214. mate variability. Reviews of Geophysics, of Biogeography, 1–7, doi:10.1111/ Reynolds, R.G., Niemiller, M.L. & Revell, 47,1–46. jbi.12751. L.J. (2014) Toward a Tree-of-Life for the Stanford, J.D., Rohling, E.J., Bacon, S., boas and pythons: multilocus species- Roberts, A.P., Grousset, F.E. & Bolshaw, level phylogeny with unprecedented M. (2011) A new concept for the paleo- SUPPORTING INFORMATION taxon sampling. Molecular Phylogenetics ceanographic evolution of Heinrich Additional Supporting Information may be 71 – and Evolution, , 201 213. event 1 in the North Atlantic. Quater- found in the online version of this article: Rocha, S., Carretero, M.A. & Harris, D.J. nary Science Reviews, 30, 1047–1066. (2005) Mitochondrial DNA sequence Tolley, K.A., Townsend, T.M. & Vences, Appendix S1 Discussion on the endemic data suggests two independent coloniza- M. (2013) Large-scale phylogeny of cha- tortoise lineages on Western Indian Ocean tions of the Comoros archipelago by meleons suggests African origins and islands. chameleons of the genus Furcifer. Belgian diversification. Proceedings of the Journal of Zoology, 135,39–42. Royal Society of London B: Biological Rocha, S., Carretero, M.A., Vences, M., Sciences, 280, 20130184. Editor: Sonya Clegg Glaw, F. & Harris, D.J. (2006) Decipher- Townsend, T.M., Tolley, K.A., Glaw, F., € ing patterns of transoceanic dispersal: Bohme, W. & Vences, M. (2011) East- doi:10.1111/jbi.12940 the evolutionary origin and biogeogra- ward from Africa: palaeocurrent- phy of coastal lizards (Cryptoblepharus) mediated chameleon dispersal to the in the Western Indian Ocean region. Seychelles islands. Biology Letters, 7, Journal of Biogeography, 33,13–22. 225–228.

1440 Journal of Biogeography 44, 1426–1440 ª 2016 John Wiley & Sons Ltd