PALAEOECOLOGICAL AND BIOCHRONOLOGICAL STUDIES OF RIVERSLEIGH, WORLD HERITAGE PROPERTY, OLIGO-MIOCENE FOSSIL LOCALITIES, NORTH-WESTERN QUEENSLAND, AUSTRALIA Kenny J mes TRAVOUILLON Thesis submitted for the degree of Doctor of Philoso.hy t the University of Ne0 South W les, Austr li August, 2008 i N1 d2ordre Ann3e 2008 THESE .r3sent3e devant l2UNIVERSITE CLAUDE BERNARD - LYON 1 .our l2obtention du DIPLOME DE DOCTORAT 8 rr9t3 du 7 o;t 2006 et rr9t3 du 6 j nvier 2005) Souten nce : Novembre 2008 . r M. Kenny J mes TRAVOUILLON TITRE : ETUDES PALEOECOLOGIQUES ET BIOCHRONOLOGIQUES DES GISEMENTS FOSSILIFERES DE L2OLIGO-MIOCENE DE RIVERSLEIGH, WORLD HERITAGE PROPERTY, NORD-OUEST DU QUEENSLAND, AUSTRALIE Directeur de thBse : Dr Serge Legendre Prof Mich el Archer Dr SuC nne H nd JURY : Dr. Serge Legendre Directeur de ThBse Pr. Mich el Archer Directeur de ThBse Pr. Christo.he Lecuyer ED min teur Dr. Gilles Esc rguel ED min teur Dr. So.hie Montuire R ..orteur Pr. Phili. Gingerich R ..orteur Dr. M nuel Hern ndeC Fern ndeC Ra..orteur ii THE UNIVERSITY OF NEW SOUTH WALES ThesisEDissert tion Sheet Surname or Family name: Tr vouillon First name: Kenny Other name/s: J mes Abbreviation for degree as given in the University calendar: PhD School: BEES Faculty: Science itle: PALAEOECOLOGICAL AND BIOCHRONOLOGICAL STUDIES OF RIVERSLEIGH, WORLD HERITAGE PROPERTY, OLIGO-MIOCENE FOSSIL LOCALITIES, NORTH-WESTERN QUEENSLAND, AUSTRALIA Abstr ct 350 0ords m Dimum: Riversleigh, World Heritage Property, located in North-western Queensland, Australia, contains over 200 fossil bearing localities from the Oligo-Miocene. The study presented here aims at finding new methods to improve the accuracy of palaeoecological and biochronological studies and describe the palaeoenvironmental and chronological settings of the Riversleigh fossil deposits. One of the methods developed in this thesis, Minimum Sample Richness (MSR), determines the minimum number of species that must be present in a fauna to allow meaningful comparisons using multivariate analyses. Using MSR, several Riversleigh localities were selected for a palaeoecological study using the cenogram method to determine the palaeoenvironment during the Oligo-Miocene. Finally, the Numerical ages method was used to refine the relative ages of the Riversleigh localities and a re-diagnosis of the Riversleigh Systems is proposed. Decl r tion rel ting to dis.osition of .roject thesisEdissert tion . hereby grant to the University of New South Wales or its agents the right to archive and to ma/e available my thesis or dissertation in whole or in part in the University libraries in all forms of media, now or here after /nown, sub0ect to the provisions of the 1opyright Act 1348. retain all property rights, such as patent rights. also retain the right to use in future wor/s ,such as articles or boo/s- all or part of this thesis or dissertation. also authorise University Microfilms to use the 350 word abstract of my thesis in 7issertation Abstracts .nternational ,this is applicable to doctoral theses only-. 8888888888888888 8888888888888888 888888888888888 Sign ture Witness D te he University recognises that there may be e9ceptional circumstances re:uiring restrictions on copying or conditions on use. Re:uests for restriction for a period of up to 2 years must be made in writing. Re:uests for a longer period of restriction may be considered in e9ceptional circumstances and re:uire the approval of the 7ean of Graduate Research. FOR OFFICE USE ONLY 7ate of completion of re:uirements for Award: THIS SHEET IS TO BE GLUED TO THE INSIDE FRONT COVER OF THE THESIS iii ______________________________________________________________________ RESUME en fran>ais Riversleigh, World Heritage Property, dans le nord-ouest du )ueensland en Australie, contient plus de 200 localit?s fossilif@res de l‘Oligo-Mioc@ne. LA?tude pr?sentée ici vise C trouver de nouvelles m?thodes pour am?liorer lDe9actitude des ?tudes pal?o?cologique et biochronologique et d?crire les param@tres paléoenvironnementau9 et chronologi:ue des localit?s de Riversleigh. LDune des m?thodes d?velopp?es dans cette th@se, Minimum Sample Richness (MSR), d?termine le nombre minimum dDesp@ces qui doivent Etre pr?sents dans une faune pour permettre des comparaisons significatives avec les analyses multivariées. En utilisant MSR, plusieurs localit?s de Riversleigh ont ?t? choisies pour une étude pal?o?cologique utilisant la m?thode des c?nogrammes permettant de d?terminer les paléoenvironnements pendant l‘Oligo-Mioc@ne. Enfin, la m?thode des âges num?ri:ues a ?té utilisée pour raffiner l‘Fge relatif des localit?s de Riversleigh et une r??valuation des Systems de Riversleigh est proposée. ______________________________________________________________________ NAME AN7 A77RESS OF THE AUS RAL.AN LABORA ORY: School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia. ______________________________________________________________________ .N . ULE E A7RESSE DE LDU.F.R. OU DU LABORA O.RE FRANCA.S: UMR 5125 PEPS, CNRS, Fr nceG Universit3 Lyon 1, C m.us de L Dou , Bt. G3ode, 69622 Villeurb nne cedex, Fr nce ______________________________________________________________________ IEH WORDS Palaeoecology, Biochronology, Riversleigh, Australia, Quercy, France, 1enogram, MSR, Numerical Ages ______________________________________________________________________ MO S-1BES Paléo?cologie, Biochronologie, Riversleigh, Australie, Quercy, France, 1?nogramme, MSR, Age num?rique ______________________________________________________________________ iv AcIno0ledgements . would like to than/ the many people that provided assistance and support throughout my Ph7 thesis. First, I would like to than/ my supervisors Professor Michael Archer and Dr Serge Legendre, my co-supervisor Dr Sue Hand and our UNSW lab manager Hen/ Godthelp. I would like to than/ Michael Bedward of the New South Wales National Par/s and Wildlife Service for writing the R routines essential to chapter 2, for his advice and for reading the manuscript. I also would like to than/ Dr Gilles Escarguel for providing his help and advice for chapter 2 and 3. I want to than/ the Palaeo lab in Sydney for their help and support, namely Pip Brewer, Dr Mina Bassarova, 7r Iaren Roberts, Dr Yamila Gurovich, Jac/ie Nguyen, Hayley Bates, Elizabeth Price, Dr Kirsten Crosby, 7r Steven Wroe, Dr Vera Weisbec/er, Dr Iaren Blac/, Dr Anna Gillespie, and especially Dr Julien Louys, Robin Bec/, and Dr Ric/ Arena whose help has significantly improved my thesis. I want to than/ the students at the University of Lyon, namely Jeremy Martin, 7r Ben0amin Greselle, Lincent Fernandez, 7r Marie-Anne Heran, 7r Caroline Sassier, Guillaume Suan, and all other students that I met and gave me their support. I than/ Prof. Bill Sherwin, 7r Peter Ban/s and 7r Alistair Poore for their advice. I than/ the South Australian Department for Environment and Heritage (particularly Andrew Graham-, Queensland National Par/s and Services (particularly Noleen Kunst), VictoriaAs Department of Sustainability and Environment for providing the National Par/ data needed for this study. Than/s to 7r Iaren Roberts, Dr Karen Blac/, Hayley Bates, Dr Anna Gillespie, Dr Anne Musser and Pip Brewer for providing dental measurements of the ta9a they are studying. I would like to than/ Dr Bernard Coo/e, Scott Hoc/null and 7r Gilbert Price for their help during my visit to the Queensland Museum. Than/s to 7r Ian Graham and Dr 7ioni Cendon for helping find references. Finally, I would like to than/ my family for their support and especially my girlfriend Kara Giles, who has supported me throughout this thesis. v T ble of contents CHAPTER 1: INTRODUCTION 1 1.1 R.LERSLE.GH WORL7 HER. AGE AREA: B.O1HRONOLOGH AN7 PALAEOE1OLOGH 2 1.2 A.MS 4 1.3 1HAP ER OU L.NE 5 1.4 REFEREN1ES 8 CHAPTER 2: FINDING THE MINIMUM SAMPLE RICHNESS (MSR) FOR MULTIVARIATE ANALYSES: IMPLICATIONS FOR PALAEOECOLOGY. 11 2.1 ABS RA1 12 2.2 .N RO7U1 .ON 12 2.3 ME HO7S AN7 MA ER.ALS 14 2.3.1 Parent Lists 14 2.3.2 Minimum Sample Richness (MSR) 15 2.3.3 Analyses 16 2.4 RESUL S 14 2.5 7.S1USS.ON 23 2.4 REFEREN1ES 24 CHAPTER 3: THE USE OF MSR (MINIMUM SAMPLE RICHNESS) FOR SAMPLE ASSEMBLAGE COMPARISONS IN CONJUNCTION WITH A TAJONOMIC DISTINCTNESS MEASURE 29 3.1 ABS RA1 30 3.2 .N RO7U1 .ON 30 3.3 MA ER.AL AN7 ME HO7S 34 3.3.1 Data sets 34 3.3.2 luster analysis 36 3.3.3 MSR calculation 38 3.3.4 Taxonomic distinctness 39 3.4 RESUL S N 7.S1USS.ON 42 3.4.1 Preliminary cluster analysis 42 3.4.2 ase study 1: (uercy and Limagne Area 43 3.4.3 ase study 2: Riversleigh 58 3.5 1ON1LUS.ONS 45 3.4 REFEREN1ES 44 3.7 APPEN7.P 70 CHAPTER 4: EJPLAINING THE GAPS IN MAMMALIAN BODY MASS DISTRIBUTIONS (CENOGRAMS) AND THE ECOLOGICAL IMPACT OF INTRODUCED SPECIES IN AUSTRALIA 81 4.1 ABS RA1 82 4.2 .N RO7U1 .ON 83 4.3 MA ER.ALS AN7 ME HO7S 87 4.3.1 Mammal species database 8, 4.3.2 Environmental data 89 4.3.3 enograms 92 4.3.4 Arboreal taxa 94 4.3.5 .ody mass distribution 94 4.3.6 Analysis 95 4.4 RESUL S 35 4.5 7.S1USS.ON 114 4.5.1 Limitations 114 4.5.2 Australia/s shifted gap 118 4.5.3 Explaining the gap 119 4.5.4 Revision of the cenogram method 122 vi 4.5.5 The impact of introduced predators 124 4.5.6 Mid1domain effect 126 4.4 1ON1LUS.ON 124 4.7 REFEREN1ES 127 CHAPTER 5: PALAEOECOLOGICAL ANALYSES OF RIVERSLEIG2S OLIGO-MIOCENE SITES: IMPLICATIONS FOR OLIO-MIOCENE CLIMATE CHANGE IN AUSTRALIA 134 5.1 ABS RA1 135 5.2 .N RO7U1 .ON 135 5.3 MA ER.ALS AN7 ME HO7S 133 5.3.1 Materials 139 5.3.2 .ody mass estimate 141 5.3.3 enogram and .ody Mass Distribution methods 141 5.3.4 Discriminant 2unction Analysis (D2A) of .ody Mass Distribution data 144 5.4 RESUL S 145 5.4.1 2aunal 3one A enograms and .ody Mass Distributions 145 5.4.2 2aunal 3one .