-il- Taxonomy and systematlcs of the genu:¡ uperolela Gray (Rnura: Leptodactylldae) by tjlargaret Davles, BSc(Hons) (Tas. ), usc (A.N.u' ) Department of ZoologY unlversity of Adelaide A thesls submitted to the Unlverslty of Adelalde for the degree of Doctor of ehllosoPhY April 1987 ù '^r,l,rtl ,= 1,,, ln, To Hike, Graeme' Llnda and Angus TabIe of Contents Volume I Page 1. Summary I l-1. Declaration ili lll . Acknotrledgrnents iv I. Introductlon T 2. Materials and Hethods 12 3. Taxonomlc Studies l4 3.1 Account of characters I4 3.2 Taxonorulc decisions 24 3.3 Checkllst of specles of Uperoleia 25 3.4 Key to UPero1eia ln Australla 26 3.5 Specles accounts 28 3.5 .l cenus Uperolela 28 3.5 .2 Uperolela marmorata GraY, l84l 3t 3.5 .3 Uperoleia laeviqata Kefersteln, 1867 32 3.5 .4 Uperoleia m'ioberqi (Andersson, I9I3) 43 3.5 .5 Uperoleia ruqosa (Andersson, 1916) 47 3.5 .6 Uperoleia russelll (Loverldge, Ì933) 6T 3.5 orientalis (Parker, 1940) 63 3.5 .8 Uoerolela arenicola lYler. Davies and Hartin' I98I 64 3.5 .9 Uperoleia borealis Tyler, Davies and Martin' I98l 67 3.5 . l0 Uperoleia crassa Tyler, Davies and Hartin, 1981 7t 3.5 . II Uperoleia lnundata Tyler, Davies and Martin' I98I ?6 3.5 . 12 Upe rolela lLthomoda Сler, Davies and llartin' t98l 84 3.5 .13 Uoerolela micromeles Tyler, Davies and Inlartin, I98I 99 3.5 .I 4 UoeroLela minima lrler, Davles and Hartin, I98l IOI 3.5 . 15 Uperolela talpa Îy'Ier, Davies and t¡lartin, t98I t03 3.5 .16 Uperolela aspera Tyler, Davles and lilartin' t98I 108 3.5 . 17 Uperoleia trachvderma lYler, Davies and ltlartin' I98I LL2 3.5 .18 Uperolela qlandulosa Davies, Hahony and Roberts, 1985 It8 3.5 . 19 Uperoleia martini Davles and LlttleJohn, 1986 t25 3.5 .20 Uperoleia tvlerl Davles and Llttlejohn, 1986 r31 3.5 .2r Uoerolela capitulata Davies, HcDonald and Corben, 1986 t39 3.5 .22 Uperolela fusca Davles, t{cDonald and Corben. 1986 I46 3.5 .23 Uperolela llttlelohnl Davles, McDonaId and corben, I986 I57 3-5 .24 Uperolel-a mlmula Davies, HcDonald and Corben, 1986 t65 3.6 Dlscusslon t74 Page 4. The ontogeny of Bone 184 4.I Introduction 184 4.1.1 Documentatlon of Ehe ontogeny of bone I84 4.I.2 The effect of ontogeny of osteologlcal features used in taxonomlc accounts 185 4. I.3 Heterochrony and paedomorphos ls in Uoerolela 187 4.1.4 The role o{ ontogeny ln the determination of character polarlty r90 4. I .5 Relationships oE Uperolela t9l 4.2 Materials and Methods 19r 4.3 Results 200 4.3.r uperoleia inundata 200 4.3.2 Uperolela trachvderma 200 4.3.3 Uperoleia lithomoda 202 4.3.4 Uperoleia laevlgata - post metamorphlc develoPment 209 4.3 .5 Uperoleia ruqosa - post metamorphlc development 2L2 4.3.6 Ranl-deI Ia slqnlfera 215 4.3 .7 Pseudophryne blbronl 220 4.3 .8 CornparaElve data and analysis 229 4.4 Discusslon 229 5. PhylogenetLc relatlonshlps wlthln Uperoleia 243 5.1 InEroduction 243 5.2 Haterials and methods 247 5.3 Results 250 5.3.1 Establlsh¡nent of an outgroup and of monophyly of the taxa under conslderatlon 250 5.3.2 Account of characters 25t 5.4 Discusslon 260 6. Dlstrlbution patterns wlthln Uperolela 265 6.I Introduction 265 6.2 Results 267 6.3 Dlscusslon 270 ?. References 277 Volume II l. IllustratLons, Flgs I-I23 2. Appendlces Appendlx I Data matrlces derlved from ontogenetlc studles l.l Uperolela laevÍqata sample I t.2 Uperoleia laevigata sanple 2 1.3 Uperoleia ruqosa r.4 RanldeIla slqnlfera 1.5 Pseudophvrne bibroni Appendix 2 Account oÊ characters wlthin Ranl-delIa Girard and Pseudophrvne Fltzinger Appendlx 3 Binary coded data matrix for phylogenetlc analysis Appendix 4 Publicatlons derived wholly or ln part from material contained withln this thesis 4.I [rler, ]1.J. , tfatson, G.F. and Davies, l{. ( 1983) AdditLons to the Erog fauna of the Northern Terrltory. Trans. R. Soc. S. Aust. 107, 243-245. 4.2 Tyler, H.J., Davies, M. and Martin, A.A. (1983) the frog fauna of the Barkly Tableland, Northern Territory. Trans. R. Soc. S. Aust. 107, 237-242. 4.3 Davies, M. (1984) osteology of the myobatrachine frog Arenophryne rotunda Tlzler (Anura: Leptodactylldae) and comparlsons wlth other myobatrachlne genera. Aust. J. ZooI. 32, 789-802. 4.4 1lzler, ![.J. and Davles, M. (1984) Uperolela Gray (Anura: Leptodactylldae) ln New Gulnea. Trans. R. Soc. S. Aust. 103, L23-L25. 4.5 Davies , l,[. and HcDonald, K.R. (1985) A redeflnltlon of Uperoleia ruqosa . Trans. R. Soc. S. Aust. I09, 37-42. 4.6 Davies, !1., llahony, M. and Roberts, J.D. (1985) A new spectes of Uperolela (Anura: Leptodactylldae) from the Pllbara Region, llestern Australla. Trans. R. Soc. S. Aust. 109, 103-108. 4.7 !'ler, H.J., Davies, H. and tlatson, G.F. (1986) The frog fauna of Groote Eylandt, Northern Terrltory, Australla. ZooL. J. Llnn. Soc. 88, 9l-I0I. 4.8 Davies, M. and LlttleJohn, H.J. (1986) Frogs of the genus Uperoleia cray (Anura: Leptodactylldae) ln southeastern Australla. Trans. R. Soc. s. Aust. Il0, t tt-t43. 4.9 Davles. M., ltlcDonald, K.R. and Corben, C. (f986) The genus UÞerolela Grav (Anura: Leptodactylidae) ln Queensland AustraÌla. Proc. R. Soc. Vlct. 98, 147-188. I SUMMARY Taxonomy and systematlcs of the genus Uperoleia Gray (Anura : Leptodactylldae) The teptodactylld genus Uperoleia Gray comprlses small, morphologtcally conservative, fossorlal specles of frogs that are dlstrlbuted throughout Australia in the wet/dry tropics, the Pllbara and eastern Australia. They penetrate New Gulnea ln the lowland areas across Torres Strait. In thls study representatives of the genus are examtned wlth respect to external morphology, osteology and. where posslble, caII. seven new species are recognlsed and one ls synonlnnlsed, resultLng ln a genus comprlslng 23 taxa. A study of the ontogeny of bone in representatlves of the genus together wlth two related sPecles Ranld slqnlfera and Sseudophrvne Uibronl)nrovldes descrlptive data on the development of bone ln both premetamorphlc and postmetamorphlc materlal. Taxonomlc declslons made with respect to the recognltlon of two cognate specles pairs, g.- lnundata and u. arenicola, ôrrd u- rugosa and u' capitulata are examined ln the llght of these ontogenetlc data. In the case of the ftrst palr, the problem ls unresolvedl but ln the case of U. ruqosa and 9. capltulata, the Judgments made are vlndlcated. Ontogenetlc data are examlned further ln the llght of the proposal of Tyler et aI. (I98la) that UperoLela ls a paedomorphlc llneage. This premlse ls accepted, and 12 characters are tdentlfied as lnfluenced by heterochrony ln Uperolela. 11 Alberch (1985) hypotheslzed that paedomorphosls should lead to a hlgh Level oE homoplasy ln a cladistic analysis of such a llneage. Preliminary data Êrom such an analysis of the relatlonshlps of taxa wlthln Uperolela do not supPort this hypotheslsl but given the weak nature of the data base. thls result requires further lnvestigatlon withln a more malleable grouP. The dlstrlbution patterns of uperoleia are unique amongst Australian vertebrates and no explanatl-on can be provided for the absence of the genus ln southwestern Australla. Local edaphic factors are important wlthl-n current distrlbutlons.,wlth only one specles, U. trachvderma.belng assoclated wlth a partlcular soll tyPe. -) 111 DECLARATION Thls thesis contains no material accepted for the award of any other degree or dlploma ln thls or any other unlverslty. To the best of my knotrledge thls thesLs contaLns no material prevlously publlshed or wrltten by any other person, except when due reference l-s made tn the text. Should thls thesis be accepted for the award of a hlgher degree, I consent to lts being made avallable for photocopylng and loan. Margaret Davies 1v ACKNOTTLEDGMENTS I have been alded greatly by a number of lndlvlduals and instltutlons durlng the course of this study. l,Iy supervisor and colleague, Hichael J. lrler introduced me to frogs ln 1975 and has nurtured my lnterest ln the l-ntervening years. Mtke has been immensely supportlve throughout the study and together wlth craeme tlatson and Angus Martin, both of the University of Helbourne, we have roamed the country of the north of tlestern Australlan and the Northern Territory durlng the wet season ln search of 'Upes' and other dlverse species, €tfld shared what must be amongst the happlest tlmes of my llfe. Kelth HcDonald of the gueensland National Parks and lflldlife Service has been an ever willing contrlbutor to the programrne by way oE material, calls and the sharing of his extensive knowledge of the genus ln eastern Australla. lilurray Llttlejohn of the Unlverslty of Helbourne and Chris Corben of the Queensland Forestry Dept made avallable thelr extenslve collections of eastern Australlan materlal wlthout which thls study would never have been attempted. The following lnstltutions and thelr respectlve curators or associates also made material avallable for the study and their generoslty ls greatly appreclated: south Australlan t{useum (u.,r. tyter), Queensland Huseum (c. tngram), Huseums of Vlctorla (.r. coventry), Llestern Australian llluseum (c. storr), Northern Territory Huset¡n (tt. King), Australian Museum (4. Greer). csrRo canberra (,1. t¿ombey) , Brltish lluseum (NaturaL Hlstory) (s. ctarke) and the Unlverslty of Kansas, Huseum of Natural History (H.8. Duellman). v Addlttonal material was provlded by Dale Roberts, lllchael Mahony' Allce ltelIs and David carter and ls acknowledged gratefully. peter Baverstock Erom the Evolutionary Blology Unlt of the South Australlan liluseum ran the computer prograÍunes for the phylogenetlc analysis and was an unfalling source of support ln this area. Colleagues ln the Department of Zoology have Provlded me wl-th abundant support and advice and I thank them all.