Taphonomic and Palaeoecological

Home , Galadi, Marlu, Marsupial mole, Northern common cuscus, Wakaleo

TAPHONOMIC AND PALAEOECOLOGICAL

INVESTIGATIONS OF RIVERSLEIGH

OLIGO-MIOCENE FOSSIL SITES

MAMMALIAN PALAEOCOMMUNITIES AND THEIR HABITATS

Mina Bassarova

Thesis submitted for the degree of Doctor of Philosophy at the University of New South Wales, Australia

August, 2005 PLEASE TYPE THE UNIVERSITY OF NEW SOUTH WALES Thesis/Dissertation Sheet

Surname or Family name: Bassarova First name: Mina Other name/s: Plamenova

Abbreviation for degree as given in the University calendar: PhD School: BEES Faculty: Science

Title: Taphonomic and palaeoecological investigations of Riversleigh Oligo-Miocene fossil sites – mammalian palaeocommunities and their habitats

Abstract 350 words maximum: (PLEASE TYPE)

The palaeoecology of selected fossil sites from the Riversleigh World Heritage Area, northwestern Queensland, Australia, was studied with the aim of describing the palaeohabitats of the sites through the use of characteristics of mammalian community structure. Taphonomic analyses were carried out to determine whether the study sites

represent allochthonous or autochthonous assemblages. Subsequently, ecological attributes of the mammalian fossil assemblages were inferred from functional morphology. Trophic and locomotor behaviours were used to describe the adaptive structure of communities and a method was established for inferring the locomotor behaviour of fossil taxa from morphometrics of their calcanea. Such ecological attributes of the assemblages can be used in reconstructing habitats. This is possible because modern mammalian community structure (as

represented by ecological diversity/attribute characteristics) has been found to correlate with habitat structure and thus has predictive value, directly applicable in palaeoecology. Modern mammalian faunas from a variety of habitats around the world were used as possible analogues for the Riversleigh fossil faunas. Multivariate statistical techniques were explored for identifying potential similarities between the community structure of the fossil faunas and that of the modern faunas. Annual rainfall was then estimated for the fossil sites through regression analysis allowing climatic inference from the faunal palaeocommunities. On the basis of similarities

in community structure, general habitat or vegetation structure was proposed for the fossil assemblages. The results of the analyses undertaken indicate that Riversleigh early-middle Miocene habitats were densely forested. The late Oligocene Quantum Leap Site local fauna and the late Miocene Encore Site local fauna suggest mixed vegetation, or more open environments. The trend of decreasing annual rainfall through the Miocene and the palaeohabitats of the Miocene sites proposed here fit the general pattern of vegetation and climate change during this period for the Australian continent as a whole.

Declaration relating to disposition of project thesis/dissertation

I hereby grant to the University of New South Wales or its agents the right to archive and to make available my thesis or dissertation in whole or in part in the University libraries in all forms of media, now or here after known, subject to the provisions of the Copyright Act 1968. I retain all property rights, such as patent rights. I also retain the right to use in future works (such as articles or books) all or part of this thesis or dissertation. I also authorise University Microfilms to use the 350 word abstract of my thesis in Dissertation Abstracts International (this is applicable to doctoral theses only).

……………………………………… …………………………………………………… ……….…………………….

Signature Witness Date

The University recognises that there may be exceptional circumstances requiring restrictions on copying or conditions on use. Requests for restriction for a period of up to 2 years must be made in writing to the Registrar. Requests for a longer period of restriction may be considered in exceptional circumstances if accompanied by a letter of support from the Supervisor or Head of School. Such requests must be submitted with the thesis/dissertation.

FOR OFFICE USE ONLY Date of completion of requirements for Award:

Registrar and Deputy Principal

THIS SHEET IS TO BE GLUED TO THE INSIDE FRONT COVER OF THE THESIS N:\FLORENCE\ABSTRACT

Table of contents

1. Introduction 1 1.1. Background 1 1.2. Aims 3 1.3. The sites 4 1.4. Chapter outline 7 1.5. References 10

2. Taphonomy of Oligo-Miocene fossil sites of the Riversleigh World Heritage Area 13 2.1. Introduction 13 2.1.1. Definitions 13 2.1.2. Time-averaging 14 2.1.3. Methods of quantification 17 2.2. Aims 21 2.3. Methods 21 2.4. Results and discussion 30 2.5. Summary 48 2.6. References 50 Appendix A1. Raw data for percentage of various break shapes on limb bones for all size categories 57 Appendix A2. Raw data for skeletal representation for small, medium and large size categories at the fossil sites studied 58 Appendix A3. Specimens used from Camel Sputum Site for age distribution analysis 59

3. The calcaneum – on the heels of marsupial locomotion 61 3.1. Introduction 61 3.1.1. Locomotor behaviour 61 3.1.2. Aims 63 3.1.3. Why calcanea? 63 3.1.4. Ancestry as a confounding factor 65 3.2. Methods 66 3.2.1. Specimens 66 3.2.2. Measurements 66 3.2.3. Locomotor categories 68 3.2.4. Notes on the modern marsupial sample 69 3.2.5. Statistical analysis 70 3.3. Results 71 3.4. Discussion 81 3.5. Conclusions 85 3.6. References 86 Appendix B1. Calcaneum measurements for modern marsupial species represented by more than one specimen 89 Appendix B2. Raw calcaneum measurements for modern marsupial species represented by single specimens 90

ii Appendix B3. Raw calcaneum measurements for modern marsupial species used as ‘unknowns’ in analysis 91 Appendix B4. Raw calcaneum measurements for fossil marsupials 92 Appendix B5. Normal probability plots for all variables 93

4. The diets of Riversleigh fossil mammals 94 4.1. Introduction 94 4.2. Mammalian cranio-dental functional morphology 95 4.3. Trophic categories 99 4.4. References 105

5. Palaeoecology of Riversleigh fossil sites 109 5.1. Introduction 109 5.1.1. Definitions 109 5.1.2. Aims 111 5.1.3. Background 112 5.2. Methods 113 5.2.1. Consideration of potential bias in fossil community representation 113 5.2.2. Modern communities: localities and habitats 118 5.2.3. Ecological diversity analyses 124 5.2.3.1. Variables used 124 5.2.3.2. Mammal taxa 125 5.2.3.3. Statistical analyses – determining habitat types and annual rainfall for fossil communities 129 5.3. Results 132 5.3.1. Consideration of potential bias in fossil community representation 132 5.3.2. Ecological diversity analyses 138 5.3.2.1.Determining broad habitat types from modern communities 138 5.3.2.2.Grouping fossil communities into broad habitat types 142 5.3.2.3.Determining annual rainfall for fossil communities 150 5.4. Discussion 154 5.4.1. Consideration of potential bias in fossil community representation 154 5.4.2. Habitat descriptions based on modern localities 156 5.4.3. Determining habitat types and annual rainfall for fossil communities 165 5.4.4. The influence of abiotic factors on vegetation structure and animal communities 169 5.5. Conclusions 171 5.6. References 173 Appendix C1. Species lists for Australian sites 181 Appendix C2. Species list for New Guinea locality 182 Appendix C3. Species lists for Central American sites 183 Appendix C4. Species lists for South American sites 189 Appendix C5. Species lists for North and Central American sites 191

iii

Appendix C6.1. Trophic and locomotor adaptations for modern marsupial and monotreme species 193 Appendix C6.2. Trophic and locomotor adaptations for modern primate and edentate species from Central and South America 196 Appendix C6.3. Trophic and locomotor adaptations for modern species of Carnivora, Perissodactyla, Artiodactyla and Lagomorphia from North, Central and South America 197 Appendix C6.4. Trophic and locomotor adaptations for modern rodent species, listed by area 199 Appendix C7.1. Split-sample cluster analysis for diet categories for modern sites 203 Appendix C7.2. Split-sample cluster analysis for diet and locomotion categories for modern sites excluding Africa 204 Appendix C8.1. Zskewness values for variables from the ‘Diet Only’ data set 205 Appendix C8.2. Zskewness values for variables from the ‘Diet and Locomotion’ data set 205

6. Australia’s changing climate through the Tertiary: review of different lines of evidence 206 6.1. Climate change 206 6.2. Evidence of climate change 207 6.2.1. Moving continents and changing currents 207 6.2.2. Palynology and plant macrofossils 209 6.2.3. Oxygen isotopes 214 6.2.4. Sea creatures 215 6.2.5. Grossplots 216 6.2.6. Terrestrial faunas 217 6.2.7. Other factors relevant to climate and vegetation 219 6.3. Concluding remarks 221 6.4. References 224

7. Conclusions and further work 228

iv List of figures

Figure 1.1.1. Map of Australia showing location of Riversleigh 1 Figure 1.3.1. Quantum Leap Site and BitesantennarySite 5 Figure 1.3.2. Camel Sputum and Mike’s Menagerie Sites 6 Figure 1.3.3. Ringtail Site 6 Figure 1.3.4. Encore Site 6 Figure 2.3.1. Relative abundance of taxa at each site 26 Figure 2.3.2. Size variation at Camel Sputum Site 26 Figure 2.4.1. Percentage of various levels of bone weathering 31 Figure 2.4.2. Percentage of various break shapes on limb bones 33 Figure 2.4.3. Fragmentation of particular elements 34 Figure 2.4.4. Extent of fragmentation for all elements and all size categories combined 36 Figure 2.4.5. Nambaroo sp. 3 from Quantum Leap Site 37 Figure 2.4.6. Turtle carapace from Quantum Leap Site 37 Figure 2.4.7. Skeletal representation 39 Figure 2.4.8. Hydraulic transport potential groupings 39 Figure 2.4.9. Mortality profiles for dasyurids and Perameles gunni 43 Figure 2.4.10. Mortality profiles for kangaroos 45 Figure 2.4.11. Jurien Caves rodent age distributions 46 Figure 3.2.1. Illustration of raw measurements for calcanea 67 Figure 3.3.1. Canonical variates plot for modern marsupial species 76 Figure 3.3.2. Log10 Body mass against log10 Calcaneum length for modern marsupial sample 80 Figure 5.2.1. Map of locations of modern Australian sites used in palaeoecological analyses 120 Figure 5.2.2. Lawn Hill Gorge and surrounding landscape 121 Figure 5.3.1. Logarithms of species abundance against body mass for primary consumers from Riverseligh fossil sites 134 Figure 5.3.2. Boxplots showing range of relative abundance of species covered by 29 modern Australian faunas for the diet/body mass groups 136 Figure 5.3.3. Cluster analysis results for diet categories for all modern sites 139 Figure 5.3.4. Plot of principal co-ordinates 1 and 2 for diet categories for all modern sites 140 Figure5.3.5. Cluster analysis results for diet and locomotion categories for all modern sites excluding Africa 141 Figure 5.3.6. Plot of principal co-ordinates 1 and 2 for diet and locomotion categories for all modern sites excluding Africa 142 Figure 5.3.7. Canonical variates plot for Diet Only data set 146 Figure 5.3.8. Canonical variates plot for Diet and Locomotion data set 147 Figure 5.3.9. Simple linear regression for Diet Only data set and annual rain 151 Figure 5.3.10. Simple linear regression for Diet and Locomotion data set and annual rain 151 Figure 5.3.11. Percentage of modern habitats with even seasonality 153

List of tables

Table 1.3.1. Study sites and their interpreted ages 4 Table 2.3.1. Taxonomic representation at the sites 24 Table 2.3.2. Hydraulic transport potential groups 28 Table 2.3.3. Tooth wear categories 29 Table 2.4.1. Percentage of specimens exhibiting evidence of possible digestion or tooth marks 42 Table 3.2.1. Descriptions of raw measurements for calcanea 67 Table 3.2.2. Variables used in canonical variates analysis 68 Table 3.3.1. Data for modern species used in canonical variates analysis 72 Table 3.3.2. Data for fossil morphotypes used in canonical variates analysis 73 Table 3.3.3. Summary statistics for modern species 75 Table 3.3.4. Canonical discriminant functions 78 Table 3.3.5. Classification results from canonical variates analysis 78 Table 3.3.6. Percent probabilities for locomotor group allocations for fossil morphotypes 79 Table 4.3.1. Description of trophic categories 100 Table 4.3.2. Diets of species and higher-level taxa as indicated by references for Camel Sputum Site 101 Table 4.3.3. Diets of species and higher-level taxa as indicated by references for Mike’s Menagerie Site 102 Table 4.3.4. Diets of species and higher-level taxa as indicated by references for Quantum Leap Site 102 Table 4.3.5. Diets of species and higher-level taxa as indicated by references for Encore Site 103 Table 4.3.6. Diets of species and higher-level taxa as indicated by references for Bitesantennary Site 103 Table 4.3.7. Diets of species and higher-level taxa as indicated by references for Ringtail Site 104 Table 5.2.1. Trophic level and body mass categories for taxa from the study sites 115 Table 5.2.2. Modern fauna site locations, habitat descriptions and average annual rainfall 119 Table 5.2.3. Vegetation types of the East Gippsland and Grampians regions of Victoria 123 Table 5.2.4. Percentages of taxa in ecological diversity categories for locomotion for modern and fossil localities 126 Table 5.2.5. Percentages of taxa in ecological diversity categories for diet for modern and fossil taxa 127 Table 5.3.1. Altered ecological diversity variables for fossil sites 138 Table 5.3.2. Summary statistics for broad habitat types from Diet Only data set 143 Table 5.3.3. Summary statistics for broad habitat types from Diet and Locomotion data set 144 Table 5.3.4. Canonical discriminant functions for Diet Only data set 148 Table 5.3.5. Canonical discriminant functions for Diet and Locomotion data set 148

vi Table 5.3.6. Classification results from canonical variates analysis of Diet Only data set 149 Table 5.3.7. Classification results from canonical variates analysis of Diet and Locomotion data set 149 Table 5.3.8. Mahalanobis distance-square from group means and percent probabilities for broad habitat type allocations for fossil communities and miss-classified moder communities for the two data sets 150 Table 5.3.9. Estimated annual rainfall five Riversleigh fossil communities based on simple linear regression 153 Table 5.4.1. Combined habitat types for the analyses of Diet and Locomotion data set (D+L) and Diet Only data set (D) 157 Table 6.1. Geological time scale 207 Table 6.2. Summary of evidence for climate and vegetation change in Australia during the late Tertiary 222

vii

ACKNOWLEDGEMENTS

Many people have provided me with assistance and support over many years in the production of this thesis. I would like to thank my supervisor, Professor Michael Archer, my co-supervisor, Dr Sue Hand and Henk Godthelp for invaluable advice and providing me with numerous opportunities for involvement in this field. I am grateful to Anna Gillespie, Karen Black, Stephan Williams and many volunteers at the University of New South Wales for the preparation of fossil material. Students and friends from the Vertebrate Palaeontology Laboratory at the University of New South Wales have been an inspiration to me, in particular, Dr Troy Myers, Dr Kirsten Crosby, Dr Steven Wroe, Dr Ben Kear, Rick Arena, Karen Black, Anna Gillespie, Adam Morell, Karen Roberts, Kenny Travouillon, Julien Louys, Zac Kirkham, Elizabeth Price, Robin Beck, Vera Weisbecker, Jenni Brammall, Pip Brewer, and Susan Henderson. In-depth discussions with Pip Brewer, Rick Arena and Kenny Travouillon have been especially valuable to me. Thank you to Dr Kay Behrensmeyer for comments and advice on the topic of taphonomy. I am also grateful to Tish Ennis and Dr Sandy Ingleby from the Australian Museum for access to specimens; Dr Dan Faith from the Australian Museum for statistical advice; Dr Bernie Cooke and Scott Hocknull from the Queensland Museum for assistance and access to specimens; Dr Ben Kear for taking measurements of specimens from the South Australian Museum; Gilbert Price and Scott Hocknull for an entertaining time in Brisbane; Dr Peter Banks, Dr Alistaire Poore and Professor Ross McMurtrie and Associate Professor Iain Suthers from the University of New South Wales for help with statistics; Professor Paul Adam from the University of New South Wales for information and advice on vegetation; Henk Godthelp for his great knowledge and many books; and Featherdale Wildlife Park and Taronga Zoo for providing scats for preliminary analyses. Special thanks go to my family for their consistent support and encouragement and patience.

viii

CHAPTER 1

MINA INTRODUCTION

CHAPTER 1

INTRODUCTION

1.1. Background

The Riversleigh World Heritage Area is located near Riversleigh Station homestead

(19o 02’ S, 138o 45’ E) on the Gregory River, approximately 200 km north-west of

Mount Isa, in north-western Queensland, Australia (Archer et al. 1996; Fig. 1.1.1). Over

300 fossil sites, or assemblages, have been identified at Riversleigh, spanning at least the last 25 million years (Archer et al. 1996). The sites occur in Tertiary limestones and were formed in fluvio-lacustrine, cave, sinkhole, or fissure fill situations (Archer et al.

1989). Most of the sites occur on two plateaus – D-site Plateau and Gag Plateau.

15o

35o

1000km

o o 125 140

Figure 1.1.1. Map of Australia showing location of Riversleigh (19o2’, 138o45’)

1 CHAPTER 1

Determination of absolute ages of Riversleigh sites has been a major challenge. This is because only sediments with certain types of lithology can be dated directly, using methods such as magnetostratigraphy and radiometric dating. Magnetostratigraphy requires the presence of rocks containing polarised ferromagnesium particles.

Radiometric dating involves the use of isotopes. Unfortunately, not all sediments contain the appropriate minerals or elements. For Riversleigh it may be possible to obtain U/Pb dates directly from primary calcites, such as those encrusting fossil bones, so a radiometric dating program is currently underway (Archer et al. 1997).

When direct dating methods cannot be applied, relative age can be determined by stratigraphic superposition (gives a minimum age), use of foraminiferal data (in cases where terrestrial faunas are interbedded with marine deposits), stage of evolution interpretations, and biocorrelation (comparison of fossil taxa in undated deposits with others in securely dated deposits). So far, temporal relationships of Riversleigh sites have been determined through stage of evolution studies, biocorrelation (Archer et al.

1989; Woodburne et al. 1985; Woodburne et al. 1994; Archer et al. 1997; Black 1997;

Cooke 1997) and stratigraphic analyses (Creaser 1997).

Riversleigh sites preserve a diverse fauna, including insects (Arena 1997; Duncan et al.

1998), molluscs, amphibians, fish, reptiles, birds and mammals (Archer et al. 1989,

1996). Plant material is not preserved sufficiently to use in the study of palaeoecology, therefore, faunal remains must be used in formulating hypotheses and making interpretations of the past environments.

2 CHAPTER 1

1.2. Aims

The Riversleigh fossil assemblages are significant in a temporal sense because they cover a period of changing climatic conditions, from relatively humid to more arid. The causes of aridity in Australia were a combination of world climatic changes and continental drift (Archer 1984). There is abundant palaeobotanical, faunal, biogeographical and palaeoclimatic evidence to support such a change (e.g. Martin

1998; Frakes 1997; McGowran et al. 2000; Archer et al. 1989; Archer et al. 1998;

Archer et al. 1995; Murray and Megirian 1992; McGowran and Li 1997; Hill et al.

1999; Truswell 1990).

The general aim of this thesis was to infer aspects of ecology from the preserved mammalian fauna at Riversleigh fossil sites. This was addressed through, firstly, determining whether faunal assemblages at the sites chosen for study are likely to represent single communities in space and time; secondly, quantification of ecological components of the preserved fauna in order to characterise mammalian communities through time at Riversleigh; thirdly, ascribing a general habitat description to the fossil mammal communities based on comparisons to modern communities from a variety of habitats; and lastly, recognising the effects of changing climate on mammalian communities and their habitats in the Oligo-Miocene at Riversleigh.

Determination of habitat for fossil assemblages through comparisons to modern analogues is possible because modern mammalian community structure has been found to correlate with habitat type and thus has predictive value, directly applicable in palaeoecology (see Andrews et al. 1979; Damuth 1992; Reed 1998; Andrews and

O’Brien 2000). The underlying principles of the study are uniformitarianism and

3 CHAPTER 1

actualism. Lyman (1994) gives a review of uniformitarian and actualistic methods as they apply to palaeontology and to making inferences about the past based on the present.

1.3. The Sites

The sites in this study were chosen to cover the period of greatest climatic/environmental change represented at Riversleigh (Table 1.3.1). All site locality data have been determined using a differential global positioning system and can be accessed via permit from the Queensland Museum and the Queensland National Parks and Wildlife Service, Brisbane.

Age (millions of Excavation size Epoch Sites years) (m3) 2 Pleistocene 3.4 Late Pliocene 5.2 Early Pliocene 6 8 Late Miocene Encore 3.84 10.4 12 14 Middle Miocene Ringtail ? 16.3

18 3.5 Bitesantennary

20 Camel Sputum 50 Early Miocene 22 Mike’s 2.7 23.3 Managerie 24 26 Late Oligocene Quantum Leap 1 28 29.3

Table 1.3.1. Study sites, their excavation dimensions and their interpreted ages (site ages from Archer et al. 1997 and Creaser 1997). Dashed line indicates upper age boundary is uncertain.

Quantum Leap (Fig. 1.3.1. A) and Bitesantennary Sites (Fig. 1.3.1. B) are located on northern D-site Plateau, separated by a distance of approximately 500 m; Camel Sputum and Mike’s Menagerie Sites (Fig. 1.3.2) are on central D-site Plateau, situated 5.4 m

4 CHAPTER 1

apart from each other; Ringtail Site (Fig. 1.3.3) is on northern Gag Plateau; and Encore

Site (Fig. 1.3.4) is on southern Gag Plateau.

Figure 1.3.1. A, Quantum Leap Site facing north-west, and B, Bitesantennary Site, facing south-east. The close-up view of Bitesantennary Site shows the cave floor (orange coloured rocks) and remainder of cave walls (grey rocks above).

Dimensions of the excavations at the sites are as follows: Camel Sputum is approximately 25 m2 with an average depth of 2 m; Mike’s Menagerie is approximately 9 m2 with an average depth of 0.3 m; Encore is approximately 6 m2 and

0.64 m deep; Bitesantennary is approximately 5 m2 and 0.7 m deep; all fossil material from Quantum Leap Site was obtained from a boulder approximately 1 m in diameter

5 CHAPTER 1

(A. Gillespie pers. comm., 2000); and the dimensions of Ringtail Site could not be measured because the excavation boundaries have become obscured over time, although its depth does not exceed one metre.

Figure 1.3.2. Camel Sputum (yellow circle) and Mike’s Menagerie (blue circle) Sites, facing north.

Figure 1.3.3. Ringtail Site, facing north-west.

Figure 1.3.4. Encore Site, facing west.

6 CHAPTER 1

Bitesantennary Site is interpreted to be a cave deposit because of the diversity and the huge number of microchiropteran specimens preserved, and also, there is a flowstone floor with a stalagmite at the base of the deposit (Hand 1997). Mike’s Menagerie Site may have belonged to the same subterranean system as Camel Sputum because they are linked by lithologically similar fossiliferous deposits. The Ringtail Site local fauna contains the monotreme Obdurodon dicksoni. It also contains the crocodilian

Trilophosuchus rackhami (Willis 1993) and other aquatic taxa, for example, lungfish, molluscs and frogs. The fossils at this locality are considered to have been deposited under low energy fluvial conditions, probably in a tufa barrage pond (Arena 2004).

Encore Site may represent a cave deposit (Arena 2004) and the environment of deposition of Quantum Leap Site is uncertain.

Geology of various Riversleigh sites has been investigated by Tedford (1967), Archer et al. (1989), Megirian (1992), Arena (1997), Morrell (2002), Price (2002) and Roberts

(2004). Also, Creaser (1997) considers sedimentology of northern D-site Plateau and southern Gag Plateau. Arena (2004) examines geology and lithology further and postulates models/theories on the processes and environments of deposition of

Riversleigh sites.

1.4. Chapter Outline

Chapter 1 is a general introduction with brief description of Riversleigh and the main aims of the research.

Chapter 2 examines taphonomy of the assemblages from the six Riversleigh sites.

Taphonomic analyses of fossil skeletal remains identify faunal assemblages as single or

7 CHAPTER 1

mixed communities and thus enable further palaeoecological studies involving habitat reconstructions to be carried out. Addressing the question of whether or not assemblages preserve spatially mixed communities involves discriminating between what was buried near the place of death and what was transported from elsewhere. It is also necessary to consider over what time frame assemblages accumulated and if temporal mixing of different communities is likely. To this end, specimens from the chosen sites were examined for evidence of exposure and transport, for example, abrasion, breakage, weathering, digestion, predator biases and skeletal part preservation.

Also, age-class distribution analysis of several peramelemorphians and a subfamily of macropodids was carried out for one of the sites to determine if the mortality profile might be attritional or catastrophic. The results contained in Chapter 2 have been published (Bassarova 2004).

Chapter 3 describes a method for determining the mode of locomotion of fossil taxa.

This method involes morphometric analysis of marsupial calcanea. The aim was to find a relationship between measurements of the calcanea and the locomotor behaviour of extant marsupial species that could subsequently be used in determining the locomotor behaviour of fossil marsupial taxa from Oligo-Miocene deposits at Riversleigh. Extant species were grouped into locomotor categories on the basis of observation and literature, and measurements of their calcanea were run through a canonical variates analysis. The locomotor categories distinguished in this study (arboreal, scansorial, quadruped terrestrial, and hopping) highlight differences between species in their use of available substrates and are thus informative with regards to the structural components of their habitat. The analysis reveals that locomotor behaviour of extant marsupials can be distinguished on the basis of calcaneum measurements and therefore, the locomotor

8 CHAPTER 1

behaviour of fossil marsupial taxa can be predicted from measurements of their calcanea. The results of this analysis form one component of the data used in the palaeoecological study (Chapter 5).

Chapter 4 reviews methods of inferring diets of mammals based on dental and cranial morphology. Described fossil taxa from the six Riversleigh study sites are subsequently placed into trophic categories based on details provided by authors in the species descriptions. The distribution of taxa amongst these trophic categories forms another component of the data used in the palaeoecological study (Chapter 5).

Chapter 5 presents palaeoecological analyses. Fossil and modern communities are characterised by ecological diversity measures (proportion of taxa in categories of locomotor behaviour and trophic habits). Habitat reconstructions for fossil faunas are based on comparisons with modern communities from distinct habitats using a variety of multivariate methods such as cluster analysis, ordination techniques and discriminant functions (canonical variates) analysis.

Chapter 6 examines Tertiary habitats at Riversleigh, with respect to general climatic/environmental changes in Australia during this period as described in the literature.

Conclusions and implications of this research and suggested further work are presented in Chapter 7.

9 CHAPTER 1

1.5. References

Andrews, P., Lord, J. M. and Nesbit Evans, E. M., 1979. Patterns of ecological diversity in fossil and modern mammalian faunas. Biological Journal of the Linnean Society 11, 177-205.

Andrews, P. and O’Brien, E., 2000. Climate, vegetation, and predictable gradients in mammal species richness in southern Africa. Journal of Zoology, London 251, 205-231.

Archer, M. 1984. Evolution of arid Australia and its consequences for vertebrates. Pp. 97-108 in Vertebrate Zoogeography and Evolution in Australia. Eds M. Archer and G. Clayton. Hesperian Press, Perth.

Archer, M., Burnley, I., Dodson, J., Harding, R., Head, L. and Murphy, A. 1998. From Plesiosaurs to People: 100 million years of Australian Environmental History. State of the Environment Australia Technical Paper Series (Portrait of Australia), Department of the Environment, Canberra.

Archer, M., Godthelp, H., Hand, S. J. and Megirian, D., 1989. Fossil mammals of Riversleigh, northwestern Queensland: preliminary overview of biostratigraphy, correlation and environmental change. Australian Zoologist 25: 29-65.

Archer, M., Hand, S. J. and Godthelp, H. 1995. Tertiary environmental and biotic change in Australia. Pp. 77-90 in Paleoclimate and Evolution, with Emphasis on Human Origins. Eds E. S. Vrba, G. H. Denton, T. C. Partridge, L. H. Burckle. Yale University Press, New Haven.

Archer, M., Hand, S. J. and Godthelp, H., 1996. Riversleigh. Second edition. Reed Books. Chatswood, NSW.

Archer, M., Hand, S. J., Godthelp, H. and Creaser, P., 1997. Correlation of the Cainozoic sediments of the Riversleigh World Heritage Fossil Property, Queensland, Australia. Pp. 131-152 in Actes du Congress BiochroM’97. Eds J. P. Aguilar, S. Legendre and J. Michaux. Mem. Trav. E.P.H.E., Inst. Monpellier.

Arena, D. A., 1997. The palaeontology and geology of Dunsinane site, Riversleigh. Memoirs of the Queensland Museum 41(2): 171-179.

Arena, D. A., 2004. The geological history and development of the terrain at the Riversleigh World Heritage Area during the middle Tertiary. PhD thesis. University of New South Wales, Sydney.

Black K., 1997. Diversity and biostratigraphy of the Diprotodontoidea of Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41(2): 187-192.

Cooke, B. N., 1997. Biostratigraphic implications of fossil kangaroos at Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41(2): 295-302.

Creaser, P., 1997. Oligocene-Miocene sediments of Riversleigh: the potential significance of topography. Memoirs of the Queensland Museum 41(2): 303-314.

10 CHAPTER 1

Damuth, J. D., 1992. Taxon-free characterization of animal communities. Pp. 183-203 in Terrestrial Ecosystems Through Time. Eds A. K. Behrensmeyer, J. D. Damuth, W. A. Di Michele, R. Polts, H. D. Sues and S. L. Wing. The University of Chicago Press, Chicago.

Duncan, I. J., Briggs, D. E. G., and Archer, M., 1998. Three-dimensionally mineralized insects and millipedes from the Tertiary of Riversleigh, Queensland, Australia. Palaeontology 41: 835-851.

Frakes, L. A., 1997. Grossplots: a method for estimating the temperature state of the Earth and of Australia, Cretaceous to middle Miocene. Australian Journal of Botany 45: 358-372.

Hand, S., 1997. New Miocene leaf-nosed bats (Michrochiroptera: Hipposideridae) from Riversleigh, Northwestern Queensland. Memoirs of the Queensland Museum 41(2): 335-349.

Hill, R. S., Truswell, E. M., McLoughlin, S. and Dettmann, M. E., 1999. Evolution of the Australian flora: fossil evidence. Pp. 251-320 in Flora of Australia. Volume 1, Introduction. Ed. A. E. Orchard. ABRS/CSIRO, Melbourne.

Lyman, R. L., 1994. Vertebrate Taphonomy. Cambridge University Press. Great Britain.

Martin, H. A. 1998. Tertiary climatic evolution and the development of aridity in Australia. Proceedings of the Linnean Society of New South Wales 119: 115-136.

Megirian, D. 1992. Interpretation of the Miocene Carl Creek Limestone, northwestern Queensland. The Beagle 9: 219-248.

McGowran, B., Archer, M., Bock, P., Darragh, T. A., Godthelp, H., Hageman, S., Hand, S. J., Hill, R., Li, Q., Maxwell, P. A., McNamara, K. J., MacPhail, M., Mildenhall, D., Partridge, A. D., Richardson, J., Shafik, S., Truswell, E. M., and Warne, M. 2000. Australasian palaeobiogeography: the Palaeogene and Neogene record. Memoirs of the Association of Australasian Palaeontologists 23: 405-470.

McGowran, B. and Li, Q. 1997. Australian Cainozoic biogeography: environmental framework for a southern-temperate province on a cooling planet. Geological Society of Australia Abstracts 48: 52-54.

Morrell, A. 2002. The geology and palaeontology of Keith’s Chocky Block Site, Riversleigh World Heritage Fossil Property, northwestern Queensland. Honours thesis. University of New South Wales, Sydney.

Murray, P. and Megirian, D. 1992. Continuity and contrast in the middle and late Miocene vertebrate communities from the Northern Territory. The Beagle 9: 195-218.

Price, E. 2002. A study of the geology of the Riversleigh area, northwestern Queensland. Honours thesis. University of New South Wales, Sydney.

11 CHAPTER 1

Roberts, K. 2004. The taphonomy and geological context of Price Is Right Site, Riversleigh World Heritage Area, northwestern Queensland. Honours thesis. University of New South Wales, Sydney.

Truswell, E. M., 1990. Australian rainforests: the 100 million year record. Pp. 7-23 in Australian tropical rainforests: science, value and meaning. Eds L. J. Webb and J. Kikkawa. CSIRO Publications, Melbourne.

Woodburne, M. O., MacFadden, B. J., Case, J. A., Springer, M. S., Pledge, N. S., Power, J. D., Woodburne, J. M. and Springer, K. B., 1994. Land mammal biostratigraphy and magnetostratigraphy of the Etadunna Formation (late Oligocene) of South Australia. Journal of Vertebrate Paleontology 13: 483-515.

Woodburne, M. O., Tedford, R. H., Archer, M., Turnbull, W. D., Plane, M. D. and Lundelius, E. L., 1985. Biochronology of the continental mammal record of Australia and New Guinea. Special Publication of South Australian Department of Mines and Energy 5: 347-363.

CHAPTER 2

TAPHONOMY OF RIVERSLEIGH OLIGO-MIOCENE FOSSIL SITES

CHAPTER 2

TAPHONOMY OF OLIGO-MIOCENE FOSSIL SITES OF THE RIVERSLEIGH WORLD HERITAGE AREA

2.1. Introduction

2.1.1. Definitions

Taphonomy is the study of the total effect of predeath circumstances and postmortem processes on organisms and on their potential for preservation in the geological record.

It includes manner of death; decay and decomposition; transportation (for example by predators, scavengers or water movements); chemical and physical modification; and burial and post-burial processes (Cadee 1991; Martill 1991; Behrensmeyer 1991;

Behrensmeyer and Kidwell 1985; Lyman 1994). Fossil accumulations, or death assemblages, are a result of ‘natural sampling’ through taphonomic processes

(Behrensmeyer 1984) and as such, may be affected with respect to taxic representation and/or quantitative measures of faunal remains. Such measures include, for example, taxonomic abundances and skeletal element representation (Lyman 1994). Taxonomic representation may be affected due to taphonomic processes obscuring the spatial association of elements such that taxa from different communities occur together in one fossil site (Hill 1979).

Identification of death assemblages as single or mixed communities is necessary for palaeoecological studies that involve habitat reconstructions based on mammalian ecological characteristics at the community level. Such a distinction can be made through taphonomic analyses that discriminate between what was buried near the place of death and what was transported from elsewhere (Behrensmeyer 1984), thereby addressing the question of whether a fossil site preserves spatially mixed communities

CHAPTER 2

(allochthonous or autochthonous assemblage of organisms). Allochthonous refers to the part of the sample that “…did not inhabit the environment of deposition and arrived there after active or passive transport” (Saunders 1977: 68, cited in Lyman 1994); autochthonous refers to locally derived fossils - those that have not been transported to the site of deposition from other habitats or environments.

The collection of specimens extracted from a fossil site for study in the laboratory is a sample of the taphonomically determined ‘natural sample’, which is in turn a sample from one or more once living communities or populations (Behrensmeyer 1984; Lyman

1994). It is important to establish whether the final sample used in a palaeoecological study is representative of the original biota and, if not, whether it is adequate to address certain palaeoecological questions by allowing biases to be identified and taken into account. Behrensmeyer (1991), Lyman (1994) and Martin (1999), among others, review analytical techniques used in taphonomy to investigate distributions of fossils and identify and determine the extent of any differences between a death assemblage and the biotic community from which it is drawn.

2.1.2. Time-averaging

In addition to determining if animal remains at one site are locally derived, it is necessary to consider over what time frame they accumulated and if temporal mixing of different communities is likely, prior to referring to them as a ‘community’ and basing inferences about habitat on them.

Time-averaging is the mixing of non-contemporaneous material in a single sedimentary stratum, so ecologically unrelated populations or communities may possibly occur

CHAPTER 2

together (Kidwell and Flessa 1996; Olszewski 1999). This generally occurs when sediment accumulation rates are slower than population turnover and can result in death assemblages that differ from the actual communities that existed (Kidwell and Flessa

1996). Usually, the result is an enhanced diversity of taxa compared to that which existed at any single moment (Behrensmeyer et al. 2000). The difference between the time of death of the oldest and youngest organisms in an assemblage gives the scale of time-averaging (Kidwell and Flessa 1996) and this scale may or may not significantly impact on our palaeobiological understanding (Behrensmeyer et al. 2000). For example, the extent of time-averaging may obscure habitat distinctions, thus leading to the false conclusion of ecological uniformity or stability (Cutler et al. 1999). On the other hand, fossil assemblages may be distinct where shifts in habitat have been slow relative to sedimentation rates and extent of time-averaging (Cutler et al. 1999).

Direct dating of fossils can be used to specify whether or not the animals in a death assemblage lived contemporaneously (Hadly 1999). However, in situations where fossils have not been dated directly, such as the Riversleigh fossil samples, the extent of time-averaging has to be estimated. Taphonomic uniformity, or lack thereof, within a site and the environment and mode of deposition of the site can be used to assess time- averaging (Behrensmeyer and Kidwell 1985). Modes of deposition for Riversleigh sites

(Arena 2004) are considered in the following taphonomic analysis for an indication of the extent of time-averaging. Also, taphonomic uniformity is investigated through observations of preservation quality within sites.

Preservation quality is a sensitive indicator of time-averaging as it has been found to differ even between periods of a few years (Fürsich and Aberhan 1990). A single

CHAPTER 2

assemblage exhibiting high variability in bone surface modifications (or preservation quality) can indicate a complex taphonomic history, which tends to correlate with greater time-averaging (Behrensmeyer et al. 2000). An additional point to consider is that fragile fossils are less likely to be spatially or temporally mixed with each other

(Kowalewski 1996, 1997), whereas more durable remains will potentially last long enough to become temporally or spatially mixed with remains from other communities

(Behrensmeyer 1984).

Kowalewski (1996) distinguishes two types of time-averaging: absolute and relative.

Absolute time-averaging denotes a scale of temporal mixing and is expressed in absolute terms. Relative time-averaging is expressed as ‘significant’ or ‘insignificant’ and depends on the time-scale of the process of interest – when the time-scale of the process is less than the scale of temporal mixing, time-averaging is significant, and conversely, it is insignificant when the time-scale of the process is greater than the scale of temporal mixing (Kowalewski 1996).

Associated with time-averaging are both advantages and disadvantages. Short-term seasonal and spatial variations in species abundance can be concealed by time- averaging but long-term palaeoenvironmental patterns may be better reflected

(Behrensmeyer and Kidwell 1985; Fürsich and Aberhan 1990). Also, rare taxa have more chance of being preserved in time-averaged deposits because of the extended temporal range (Cutler et al. 1999) which benefits the interpretation of faunal diversity

(Hadly 1999). Time-averaging (over a period of hundreds to thousands of years), during periods of relative ecological stability, can result in more reliable sampling than instantaneous deposits, with up to 93% of the mammal fauna in a habitat being

CHAPTER 2

represented by the time-averaged sample (Hadly 1999). If Riversleigh accumulations represent hundreds or thousands of years, there would be no problems associated with interpretation assuming that habitats and communities remained stable over this time period.

Possible disadvantages of time-averaging are: the time factor, in combination with various biological and environmental factors, produces changes in palaeocommunity parameters such as taxonomic composition, relative abundance and diversity of species

(Fürsich and Aberhan 1990). Species diversity will be higher in a temporally mixed sample compared to the average diversity at any one moment during accumulation of the sample (Peterson 1977; Behrensmeyer et al. 2000). Also, time-averaging can generate a false pattern if, for example, time-slices from different climates and their associated habitat types are preserved (Kowalewski 1996). When fossils from changing communities (due to climate and habitat changes) are lumped into one time-averaged sample, the non-contemporaneity of the organisms may make speculations about niche partitioning unreliable (Fürsich and Aberhan 1990).

2.1.3. Methods of quantification

Quantification of taxa and elements from a fossil site allows a vast array of analyses, both taphonomic and palaeoecological, to be undertaken. In the following taphonomic study, elements were counted in order to establish skeletal part frequencies represented at the sites and thus determine if transport or some form of preferential preservation has taken place. The relative abundance of ordinal-level taxa was calculated to identify possible taxonomic biases in preservation. With respect to palaeoecological and

CHAPTER 2

community analyses, calculation of relative abundance of taxa provides important information in terms of community structure and ecological patterns/organisation.

Many different fossil quantification methods exist. Badgley (1986) argues that these methods should be selected on a case-by-case basis, based on the sedimentary environment and processes of accumulation of a bone assemblage, because different counting methods can result in very different numerical results. In the case of

Riversleigh fossils, postcranial elements cannot be used for relative abundance of taxa counts because it is, at this stage, impossible to assign them to a taxon with certainty.

The reason for this is that many Riversleigh species- and generic-level taxa, described chiefly from craniodental specimens, are so far unknown from anywhere else and skeletal remains at Riversleigh sites are, in most cases, disassociated. Dental elements, on the other hand, are appropriate for the purpose of calculating relative abundance of taxa.

NISP (number of identifiable specimens per taxon) and MNI (minimum number of individuals) are the most commonly used quantitative indices (Lyman 1994). Modified versions of these, and other counting methods, include MNE (minimum number of skeletal elements per taxon), WAE (weighted abundance of elements), MAU (minimum number of animal units), matched-pairs method, and relative frequency estimates

(Holtzman 1979; Grayson 1984; Badgley 1986; Lyman 1994; and references therein).

The following study uses the NISP method, applied to isolated molar teeth, dentaries and maxillae, to obtain the relative abundance of taxa. Skeletal element representation

CHAPTER 2

was calculated in a similar way to the NISP method, however, the totals of different element types were calculated irrespective of taxon.

There are assumptions inherent in each of the quantification methods listed above. The

NISP count is the total of all identifiable specimens of a taxon in a sample; it assumes all specimens are independent and are equally affected by chance breakage (Grayson

1984). Consequently, a criticism of this method is sample inflation as a result of counting fragments of elements which may not be independent, but may be pieces of the same element (Grayson 1973, 1984). There are other problems associated with the NISP method (listed in Grayson 1984). Some (those that involve ‘butchering’) are relevant only in archaeological and not palaeontological contexts. The use of only dental elements to obtain the counts for relative abundance of taxa in the following study, overcomes some of the remaining problems (e.g. differential identifiability of elements from different taxa; differential preservation of different elements). NISP gives not the number of individuals in a taxon but the relative abundance for that taxon, because it is improbable that each specimen belongs to a different individual (Badgley 1986).

The remaining quantification methods are briefly described below. MNI is defined as

“that number of individuals which are necessary to account for all of the skeletal elements (specimens) found in the site” (Shotwell 1955: 272). Problems associated with this method include exaggeration of the importance of rare animals (Payne 1972, cited in Grayson 1978; Ducos 1968, cited in Grayson 1984); effects of sample size on MNI values (Grayson 1978); and the assumption that there is a high probability of association between elements for the assemblage under study (Badgley 1986).

CHAPTER 2

MNE takes fragmentation into account, so all fragments that could belong to one element are grouped together (Badgley 1986), leading to a potential error of unknown magnitude from fragments that are difficult to identify to an element.

WAE, the “frequency of elements divided by the number of elements per individual”

(Holtzman 1979: 80), was devised in an effort to minimise bias caused by differential preservation of elements (Badgley 1986). However, this method is seen as inappropriate because the correction factor (specification of how many elements can be preserved per taxon) is variable and subjective; it can differ between taxa and even between individuals of one taxon in different depositional environments (Badgley 1986).

MAU’s, introduced by Binford (1984), are counts of elements divided by the number of times each element occurs in the skeleton of the taxon involved (Grayson 1984).

Grayson (1984) elaborates on flaws in this method.

The matched-pairs method, developed by Krantz (1968), requires elements in the fossil sample to be matched up based on their pairing in a skeleton. It is an attempt to estimate the original number of animals that may have contributed to the sample (Fieller and

Turner 1982). This method requires the retrieval of all fossil bones from a site (not always possible and not the case for Riversleigh) and there is potential for inaccuracy resulting from falsley matched and unmatched pairs (Grayson 1984).

The RF estimate requires fragments of elements and whole elements to be identified to a taxon. The number of fragments are counted and “…corrected for the frequency of each bone type in the skeleton and for the number of bone types recovered for each

CHAPTER 2

taxon” (Badgley 1986: 329 and Gilbert et al. 1982, cited within). There are many difficulties associated with this method with regards to applying it to fossil material from Riversleigh, the main ones being the inability to assign postcranials to specific taxa, as mentioned above, and the difficulty of recognising which elements fragments may be a part of.

2.2. Aims

The major aims of the taphonomic analyses were to determine whether the six study sites represent allochthonous or autochthonous assemblages and, where possible, whether accumulation was attritional or catastrophic. These aims were addressed through investigation of taxonomic presence and relative abundance at the sites; the extent of exposure of specimens prior to burial; skeletal element representation at sites; extent of predator/scavenger activity; analysis of skeletal completeness according to potential for water transport and sorting of bone elements; and age class distributions of taxa.

2.3. Methods

Fossils are collected from Riversleigh deposits in blocks of limestone matrix, which are transported to the University of New South Wales for processing. All fossil material is extracted from the matrix through dissolution in weak acetic acid. Bone is then separated from other undisolved material and sorted under a microscope.

Mammalian skeletal and cranial remains were used for most aspects of the taphonomic study. Bird, reptile, amphibian, and fish fossils were not considered (except where specifically indicated). The reason for this is the major macrostructural and

CHAPTER 2

microstructural variations between fish, bird, reptile, amphibian and mammal bones and the lack of work focusing on comparisons in the responses of these structurally varied bones to particular taphonomic processes (Lyman 1994).

Lyman (1994: 22 and references therin) provides a summary list of field and laboratory data ideal to have recorded for taphonomic analyses (also see Behrensmeyer 1991). The data obtained for the following taphonomic study include degree of disarticulation; association of elements; relative counts of different elements; taxonomic identification and abundance based on dentition; surface modification attributes of specimens such as weathering extent, breakage pattern, extent of abrasion, predator/scavenger tooth marks, and evidence of digestion. The in situ distribution and orientation of individual bones has not been included because the specimens used were collected and processed over many years and such information was not recorded for the bulk of the material. The collection methods involved selection of fossil-bearing limestone in the field, followed by dissolution of the limestone with dilute acetic acid in the laboratory and screening of insoluble residues which were subsequently sorted for skeletal material. Processing was carried out by Stephan Williams, Anna Gillespie, Karen Black and students and volunteers at the University of New South Wales.

Taxa found at each site were noted for an indication of taxonomic diversity (Table

2.3.1). Myers et al. (2001) provide more details on the fauna of Encore Site. To estimate the relative abundance of taxa at each site (Fig. 2.3.1), dental specimens were counted using the NISP method (Grayson 1984, Badgley 1986). Postcranial elements were not used in obtaining relative abundance of taxa for reasons outlined above.

CHAPTER 2

All specimens were put into size groups (small, medium or large) because taphonomic processes may affect small and large bones in different ways. All fossil specimens (see

Fig. 2.3.2 for an indication of size variation at sites) were assigned to a size group by comparison to corresponding elements of a potoroo (Potorous tridactylus; average

1.1 kg, Johnston 1998) and a swamp wallaby (Wallabia bicolor; average 15 kg,

Merchant 1998). Dimensions measured were length and circumference for limb bones and other elongated elements; occlusal length and width for teeth; and maximum width, length and height for vertebrae, pelves, scapulae, crania and dentaries. Specimens the same size or smaller than corresponding elements of the potoroo were classified as small, those intermediate in size between the potoroo and wallaby were classified as medium, and those larger than the wallaby were classified as large.

CHAPTER 2

Sub- Sub family or Order Superfamily Family Genus Species Q C M R B E order tribe Yalkapari- Yalkapari- Yalkapari- Yalkaparidon coheni (1) dontia dontoidea dontidae Notorycte- Notoryct-

morphia idae Dasyur- Barinya wangala (2) idae Ganbulanyi djadjinguli (3) Wabulacinus ridei (4) Ngamalacinus timmulvaneyi (4) Dasyuro- Thyla- Dasyuroidea Maximucinus muirheadae (5) morphia cinidae Thylacinus macknessi (6) sp. cf. T. macknessi Incertae Mayigriphus orbus (7) sedis

Yaralidae Yarala burchfieldi (8) Peramele- Genus 1 sp. Yaraloidea Incertae morphia Genus 2 sp. sedis Genus 3 sp. V-Didae Wynyard-

iidae Litokoala kanunkaensis (9) Phasco- Nimiokoala greystanesi (10) larctidae Phascolarctos sp. Vombat- cf. Warendja sp. idae Thylaco- Wakaleo vanderleuri (11) Vombat- iformes leonidae Priscaleo

Diproto- Nimbadon lavarackorum (12) dontidae Zygomatu-rinae sp. Neohelos Diproto- tirarensis (12) dontoidea Propalorchestes ponticulus (13) Palorchest-

idae Palorchestes anulus (14) sp.

Phalan- Strigocuscus reidi (15)

Phalanger- geridae Wyulda asherjoli (16) Diproto- oidea Trichosurus sp. (15) dontia Pilkipild-

ridae Tarsiped- Acrobat-

oidea idae Burramy- Burram- Burramys brutyi (17)

oidea yidae Cercatetus Petauridae Pseudochirops Phalan- cf. kutjamarpensis gerida Marlu Pseudo- (18)

Petauroidea cheiridae maxbourkei (19) Paljara nancyhawardae (19) Pildra sp. 2 Incertae Djaludjangi yadjana (20) sedis Macropod-inae Wabularoo naughtoni (21) Macropod- Macropod- Nowidgee matrix (22) Bulunga- oidea idae Bulungamaya delicata (23) mayinae bilamina (22) Gangaroo sp. nov.

Table 2.3.1. Taxonomic representation at the sites. Table continued next page.

CHAPTER 2

Sub- Sub family or Genus Order Superfamily Family Species Q C M R B E order tribe hilarus (24) Wanburoo Macro- Bulunga- sp. podidae mayinae Gumardee Sthenuri-nae Hadronomas cf. Hadronomas sp. Potoro- Potoroinae idae Hypsi- Hypsiprymno- Diproto- Phalan- Macropod- Hypsiprymnodon dontia gerida oidea prym- dontinae nodont- ima (25) Propleopinae Ekaltadeta idae jamiemulvaneyi (26) Ganawamaya acris (27) Balbaroo gregoriensis (23) Balbar- Nambarinae Wururoo sp. idae sp. 5 Nambaroo sp. 3 Ornitho- Mono- rhynch- Obdurodon dicksoni (28) tremata idae

Xenorhinos halli (29) Riversleigha williamsi (30) Brachipposideros watsoni (31) Rhinonicteris Rhinoloph- Hippos- tedfordi (31) Micro- oidea ideridae Chiroptera chiro- Bitesantennary sp. 1 Bitesantennary sp. 3 ptera Rhinonycterini Bitesantennary sp. 4 Bitesantennary sp. 5 Bitesantennary sp. 6 Bitesantennary sp. 7 Bitesantennary sp. 8 Bitesantennary sp. 9 Mega- Macroderma dermat- idae

Table 2.3.1. continued…Taxonomic representation at the sites (grey shading refers to presence of taxa). Q = Quantum Leap; C = Camel Sputum; M = Mike’s Menagerie; R = ringtail; B = Bitesantennary; E = Encore. Number in brackets indicates the reference where the species is described; references are as follows: 1. Archer et al. (1988) 13. Murray (1990) 25. Archer and Flannery (1985) 2. Wroe (1999) 14. Black (1997b) 26. Wroe (1996) 3. Wroe (1998) 15. Flannery and Archer (1987) 27. Cooke (1992) 4. Muirhead (1997) 16. Crosby et al. (2001) 28. Archer et al. (1992) 5. Wroe (2001) 17. Brammall and Archer (1997) 29. Hand (1998a) 6. Muirhead (1992) 18. Woodburne et al. (1987) 30. Hand (1998b) 7. Wroe (1997) 19. Bassarova et al. (2001) 31. Hand (1997) 8. Muirhead and Filan (1995) 20. Brammall (1999) 32. Kear (2002) 9. Springer (1987) 21. Archer (1979) 10. Black and Archer (1997) 22. Cooke (1997b) 11. Clemens and Plane (1974) 23. Flannery et al. (1982) 12. Hand et al. (1993) 24. Cooke (1999)

CHAPTER 2

100 90 80 70 60 50 Percent 40 30 Encore (n=98) 20 Bitesantennary (n=142) Ringtail (n=104) 10 Site (n=NISP) Mike's Menagerie (n=203) 0 Camel Sputum (n=693) Quantum Leap (n=234)

Yalkaparidontia Chiroptera Diprotodontia Monotremata Dasyuromorphia Notoryctemorphia Peramelemorphia Order

Figure 2.3.1. Relative abundane of taxa at each site

5 cm

Figure 2.3.2. Size variation at Camel Sputum Site

CHAPTER 2

Weathering and breakage patterns reveal the extent of aerial exposure of bones prior to fossilisation. Specimens (excluding teeth due to structural differences) from each site were scored for six categories of bone surface deterioration (adapted from

Behrensmeyer 1978 and Coombs and Coombs 1997). To determine when bones from each site were broken (fresh, after aerial exposure, or after fossilisation), breakage of limb bones was categorised into groups (adapted from Shipman 1981 and Coombs and

Coombs 1997). Although it is arguable that fracture shapes indicate when breakage occurred (Shipman 1981), spiral fractures are more common in fresh bones, transverse and longitudinal fractures occur on mineralised bones, and stepped fractures develop after long aerial exposure (Andrews 1990, Behrensmeyer 1991, Coombs and Coombs

1997). The extent of fragmentation was also investigated as an indication of the degree of disturbance or bioturbation of pre-fossilised material at the sites.

To determine if fossils from each site were locally derived, they were examined for: abrasion; evidence of predator/scavenger activity; skeletal element representation; and association of elements. Also, specimens were grouped on the basis of their potential for hydraulic transport. These hydraulic transport potential groups (referred to as 1, 2 and 3; see Table 2.3.2) are based on the findings of Voorhies (1969), Behrensmeyer (1975),

Dodson (1973) and Korth (1979) for medium sized and small mammals. The groups combine the information from these authors but, where appropriate, certain elements are excluded (for example, ribs were removed as there is likely to be a sorting bias against

CHAPTER 2

GROUP ELEMENTS vertebrae 1 sacra sternum scapulae limbs 2 foot/hand bones teeth 3 dentaries

Table 2.3.2. Hydraulic transport potential groups (modified from Voorhies 1969, Behrensmeyer 1975, Dodson 1973 and Korth 1979)

them in the laboratory because they are not taxonomically informative and are hard to recognise when broken). Group 1 includes vertebrae, sacra and sternums - those elements most easily transported by water; group 2 includes scapulae, limbs, foot/hand bones and teeth – elements that are intermediate in their water transport potential; and group 3 includes dentaries – the elements most resistant to water transport. The distributions of fossil specimens (within their size categories) among these groups were compared to the numbers expected for no hydraulic transport. These expected numbers were obtained by calculating the average number of elements in each group from marsupial skeletons (these numbers are generally close to numbers for the average michrochiropteran; there is further discussion of this, with respect to bat remains from

Bitesantennary Site, below).

Age-class distribution analyses were performed in an attempt to determine if assemblages exhibit attritional or catastrophic mortality profiles. An attritional mortality profile indicates an assemblage accumulated over time and reflects the number of individuals in each age class that are dying. Catastrophic mortality profiles may result from instantaneous mortality events. Some assemblages, however, may exhibit a catastrophic mortality profile but may have been accumulated over a long period of time, for example, pitfall trap and some owl pellet deposits which sample randomly

CHAPTER 2

from the living population and reflect the proportion of individuals in each age class in that living population.

The age classes used for the fossils in this study are arbitrary categories determined from the extent of tooth wear (Table 2.3.3) and are based on the assumption that teeth wear down at a constant rate over time. Peramelemorphians (bandicoots) and a subfamily of macropodids (kangaroos) were examined only from Camel Sputum (see

Appendix 1 for specimen list), because of sample size limitations at the other sites. Only

Taxon Wear Life stage Features category 1 Juvenile Totally unworn to slight exposure of dentine on tips of cuspids 2 Sub-adult Dentine exposed along length of crests 3 Adult Dentine exposed within talonid basin; cuspids worn to 90% of their original height 4 Adult Cuspids worn to 75% of their original height 5 Aged Cuspids worn to 50% of their original height; cuspids are same height as MORPHIAN MORPHIAN PERAMELE- interconnecting crests 6 Very old Trigonid and talonid basins worn smooth; crown almost entirely worn away 0 Juvenile No wear 1 Sub- Facets and striae on lophs/lophids adult 2 Young Dentine exposed on protocone and/or metaconule; dentine exposed on adult protoconid and/or metaconid 3 Adult Dentine exposed on paracone and/or metacone (as well as protocone and metaconule); dentine exposed on hypoconid and/or entoconid MACROPODID MACROPODID 4 Adult Dentine exposed continuously along protoloph/protolophid 5 Aged Dentine exposed continuously along lophs/lophids 1 Juvenile No wear on cusps; lingual and buccal halves of mures separated by enamel 2 Sub- Cusps worn slightly; one or two mures show dentine between the buccal adult and lingual halves 3 Adult Dentine exposed between buccal and lingual halves of all mures

RODENTS RODENTS 4 Adult Dentine exposed between first and second mures 5 Aged Dentine exposed between all mures; no enamel between mures; thin rim of enamel on periphery of crown

Table 2.3.3. Tooth wear categories. Peramelemorphian tooth wear categories for lower first molar, adapted from Morrell (2002); Macropodid tooth wear categories for upper and lower M1or M2, adapted from McArthur and Sanson (1988) and Lentle et al. (1998) but note that application of these categories to both upper and lower dentition assumes that the buccal cusps of the lower teeth wear down at the same time as the lingual cusps of the upper teeth – [see Sanson 1989 paper to check if this is the case]; Rodent tooth wear categories adapted from De Oliveira et al. (1998).

CHAPTER 2

first lower molars were used in assigning fossil peramelemorphians to an age class so as to overcome the problem of differential wear of teeth in the tooth row of individuals.

For macropodids, upper and lower first molars were used (second molars were used where first molars were not available so as to maintain maximum sample size).

Catastrophic and attritional death curves for extant bandicoots and macropodids were constructed for comparison, using the data of Dufty (1994) and Dawson (1995) respectively (Figs 2.4.9. B and 2.4.10. A). Also for comparison, dasyurids and rodents from Brown Bone Cave (Jurien Caves, Western Australia; AR 19145, collection of

UNSW) owl pellet deposit, of probable Quaternary age, were examined, as examples of fossil catastrophic mortality profiles (Figs 2.4.9. A and 2.4.11).

2.4. Results and discussion

There appears to be a bias in the size of the specimens preserved at most sites. Most numerous are the specimens in the small size category. This is most likely a result of animals with smaller body sizes generally having faster turnover rates and larger populations. However, the preservation of a high proportion of small specimens is unusual because small material is very fragile. This preservation is therefore interpreted to be the result of a low energy depositional regime combined with a lack of aerial exposure (see below for discussion of weathering patterns). Mike’s Menagerie is the only site of the six to have produced greater numbers of specimens in the medium size category than in the small size category. It is possible that this is a result from the fact that Mike’s Menagerie has not been sampled as extensively as the other five sites.

Examination of weathering patterns reveals that all sites have a predominance of smooth bone for the small size category (Fig. 2.4.1. A), indicating minimal aerial exposure. The

CHAPTER 2

120

100

80 Bitesantennary (n=2653) Ringtail (n=167) Encore (n=736) 60 Quantum Leap (n=149) Percent Mike's Menagerie (n=84) Camel Sputum (n=1237)

0 smooth smooth-cracked cracked cracked-flaking flaking eroded

W eathering category

B. 80

Ringtail (n=35) Encore (n=216) 60 Quantum Leap (n=73) Mike's Menagerie (n=83) Camel Sputum (n=332) 50

40 Percent

0 smooth smooth-cracked cracked cracked-flaking flaking eroded Weathering category

Figure 2.4.1. Percentages of various levels of bone weathering for: A, small size category and B, medium size category. Behrensmeyer’s (1978) weathering categories 0, 1, 2, and 2-3 correspond to smooth, smooth-cracked and cracked, cracked-flaking, and flaking, respectively but are possibly not equivalent in exposure time. Some weathering stages shown by photos.

CHAPTER 2

medium size category shows a greater variance of weathering states (Fig. 2.4.1. B).

Encore Site has a majority of medium-sized specimens in the ‘eroded’ category. The interpretation of the ‘eroded’ state is uncertain but it may possibly represent the effect of chemical reactions. The large size category, for sites with adequate sample sizes among the six studied (not shown here), follows the trend of increasing variance of weathering states. It appears that medium sized and large elements have had slightly longer aerial exposure prior to fossilisation than small elements. This may be because larger bones last longer unprotected and, of the small elements, only those that were quickly buried survived.

Breakage patterns (Fig. 2.4.2; raw data in Appendix A1) reveal that the majority of limb bones, from all sites, were broken either fresh (spiral breaks) or after fossilisation

(tranverse breaks). Relatively few were broken after long aerial exposure (stepped breaks). This is consistent with the extent of weathering. In Riversleigh deposits, post- fossilisation breaks may occur in situ when erosion creates fissures through the limestone, or as a result of the collection process.

The extent of fragmentation of particular skeletal element types is shown for small and medium size categories for the sites (Figs 2.4.3. A and B). Small elements from all sites follow a similar pattern, from periotics, teeth and foot/hand bones being mostly unbroken or entire, to pelves, dentaries, maxillae and crania being mostly fragmented

(ribs are ignored because of the small sample size). The medium size category follows a similar pattern in most cases (ignoring elements with a sample size under five), but with a smaller proportion of entire elements. This again may be due to longer exposure of larger elements prior to burial.

CHAPTER 2

60 Bitesantennary (n=85) Ringtail (n=12) 50 Encore (n=14) Quantum Leap (n=51) 40 Mike's Menagerie (n=11) Percent Camel Sputum (n=117) 30

0 spiral transverse longitudinal stepped Type of breakage

Figure 2.4.2. Percentages of various break shapes on limb bones for all size categories (break shape pictured above corresponding breakage type).

Figure 2.4.4 shows the extent of fragmentation for all elements and all size categories combined. There is a predominance of elements that are unbroken in all sites. This is due mainly to the fact that isolated teeth were included in the counts (except in the case of Bitesantennary, due to the enormous number of isolated teeth). The total proportion of broken elements is approaching 50% for all sites, except Encore, suggesting a considerable degree of localised disturbance, such as trampling.

Fossils from none of the sites show sufficient surface abrasion to indicate significant fluvial transport. For example, from Quantum Leap only 3 specimens out of 1107 exhibit abrasion, with this being slight in two of the specimens and moderate in one.

CHAPTER 2

100%

80%

60% QL 40%

20%

0% ) ) ) ) ) ) 8) 3) 7 8) 2) 4 5 5 4) 4 0) ( 8 ( 1 14 s (2 (3 e ( ( 100% c e i um ( bs bs a ot n th (2 ri ries (5 e im pula ill 90% ter te l a x pelvis peri s rebrae (18 sc cranial (6 denta ma foot/hand ve(20 80% Element 70% 60% CS 50% MM 40% 30% 100% 20%

10% 80% 0%

) ) ) ) ) ) ) ) ) ) ) 1 04 e (4 (27 60% s (70 la nial (3 eth (642 ribs (2 ra te limbs (117 pelvis (16 c scapu periotics (99 dentarie maxillae foot/hand (4 vertebrae (328 40% Element

20%

0% 100% ) ) 0 2) (1 s (5) (15 is (2) v nd (29) mbs (9) iotic li r teeth (4 ha ebrae (36) cranial (1) pel 80% rt ntaries pe e maxillae foot/ v de Element 60% Ring

40% Bite Ring 100% 20%

90%

80% 0% ) ) (1) 2) (43) 63 (41) 70% is (4) v ribs h (35 ics ( ae (27 ies (74) et br mbs iot li pel tar 60% te cranial (1) ot/hand maxillae (31) fo per verte den 50% Element

40%

30%

20%

10%

0% 100% periotics sacra (17) ribs (151) teeth vertebrae foot/hand limbs (683) scapulae cranial (64) dentaries maxillae (3164) (2307) (952) (267) (83) (440) (55) 90% Element 80%

LEGEND 70% entire 60% partial (<50% broken) Ec 50% fragmented (>50% broken) 40% 30% 20% 10%

) ) ) ) ) 3) 7) (7) (1 (6 (43 58) (10) 901 (27 (23 al (1 s (1 ibs (4 bs i ae r m lae h nd ics li an ies il ebr cr pelvi /ha iot t ar ax teet er nt m er v foot p de Element Figure 2.4.3. A. Fragmentation of particular elements for small size category for Quantum Leap (QL), Camel Sputum (CS), Mike’s Menagerie (MM), Ringtail (Ring), Bitesantennary (Bite), and Encore (Ec) Sites

CHAPTER 2

100% LEGEND 90% entire 80% partial (<50% broken) 70% fragmented (>50% broken) 60%

50% QL 40% 30%

20% 10% 0% ) ) ) ) 2 1 0 0 ( ( 6 1 (4) s l ( e (35) (37) (17) e ia d ( h e s (23) s i t la a b is r n n e il i lv ta ra e x br mb r e n c /ha t e li p ma rt de ot scapulae (2) e fo v Element 100% 90% 80%

70%

60%

CS 50%

40% 30% 20%

10% 0% ) ) ) ) 1 (4 l (2) (3) (9) 10) (174) s (30) ia ribs cs (25 b lae illae ( /hand (49) teeth (4 brae taries (17 lim cran pelvis en scapu max foot perioti d MM verte Element 100% 90% 80% 70% 60% 50%

40%

30%

20%

10%

) 1) s (8) (11) ribs (1) riotics (3 teeth (23) limbs pelvis (4) t/hand (44) maxillae (4) pe dentarie ertebrae (1 foo v 100% Element 90%

80%

70%

60%

50%

Ring 40%

30%

20%

10%

0% Ec cranial (1) foot/hand teeth (10) ribs (3) vertebrae limbs (3) dentaries maxillae (17) (9) (2) (1) 100% Element 90% 80% Figure 2.4.3. B. Fragmentation of

70% particular elements for medium size

60% category from Quantum Leap (QL),

50% Camel Sputum (CS), Mike’s

40% Menagerie (MM), Ringtail (Ring)

30% and Encore (Ec) Sites. Bitesantennary

20% not included due to small samle size.

10%

0% foot/hand periotics limbs (16) vertebrae teeth (96) pelvis (3) dentaries scapulae (91) (4) (57) (10) (1) Element 35

CHAPTER 2

40 Percent 30

) ) ) ) 8 4 6 84) 68 430) 0 2 n=6 n=1 ( ( il (n= a p (n=1 rie ary (n=2580 a n ngt ge Ri Encore na Me 's e k Bitesanten Quantum Le Camel Sputum (n=211 Mi

Entire Partial (<50% broken) Fragmented (>50% broken)

Figure 2.4.4. Extent of fragmentation for all elements and all size categories combined (n = sample size)

Vertebrate remains from the six sites are disarticulated, except for one snake vertebral column from Camel Sputum. The separation of all vertebrae normally indicates exposure prior to burial (Behrensmeyer 1975, Hill 1979). However, in combination with the predominance of smooth bones, the extent of disarticulation suggests moist conditions created by microhabitats or standing water. Such conditions have been found to accelerate disarticulation and lead to rapid burial and therefore, good preservation

(Coe 1980). The bat deposit at Bitesantennary Site probably occurred in water rather than guano because ostracod and fish fossils are found there (Hand 1997). From

Quantum Leap there is one almost complete kangaroo skeleton (Fig. 2.4.5) but the bones are associated, not articulated. Quantum Leap also preserves fragments of turtle shell which have been reconstructed into a nearly complete carapace (Fig. 2.4.6). Also,

Encore Site has produced associated skull and postcranials of Wanburoo sp.

(Kear et al. 2001) and Rhizosthenurus flanneryi (Kear 2002, Kirkham 2004).

CHAPTER 2

Figure 2.4.5. Nambaroo sp. 3 from Quantum Leap Site (ruler = 30 cm)

Figure 2.4.6. Turtle carapace from Quantum Leap Site

This is further evidence that, for these sites, bones were not exposed to significant fluvial transport. Aside from these associated elements, most of the sites have produced

CHAPTER 2

disassociated elements indicating again some non-transport post-mortem disturbance, such as bioturbation.

Elements in the medium and large size categories, on the whole, are represented in similar proportions to the expected for an unbiased sample (Fig. 2.4.7. B and Appendix

A2 for raw data; only medium category is shown in Fig 2.4.7). The expected proportions of elements are the relative numbers of each element type in an average marsupial skeleton. Medium elements from Mike’s Menagerie and small elements from most sites have a higher than expected proportion of teeth and dentaries and fewer foot/hand bones, ribs and vertebrae than expected (Fig. 2.4.7. A and Appendix A2). A possible explanation is that ribs, vertebrae and foot/hand bones are not as robust as dental elements and therefore have a greater chance of being broken beyond recognition, especially if they are in the small category and thus very fragile. Also, a collecting/sorting bias may result from the fact that dental elements, even if broken, are more easily recognisable and more taxonomically informative than broken postcranial pieces.

Hydraulic transport potential groupings are shown only for the small and medium size categories (Fig. 2.4.8); the sample size for the large category was too small for most sites. The Quantum Leap medium category exhibits almost identical distribution among the groups to the expected for no fluvial transport (0.5

Leap small category also approaches to this pattern. Ringtail medium, Encore medium,

Bitesantennary small, Mike’s Menagerie small and Camel Sputum small categories deviate slightly from the expected in having fewer group 1 elements. Dental elements

CHAPTER 2

Quantum Leap Camel Sputum Mike's Menagerie Encore Site Ringtail Bitesantennary Expected pelvis sacrum sternum ribs maxilla limbs teeth cranium periotics scapulae dentaries Element vertebrae foot/hand

Quantum Leap Camel Sputum Mike's Menagerie Site Encore Ringtail Expected pelvis sacrum sternum maxilla ribs limbs

cranium teeth periotics scapulae dentaries

vertebrae Element foot/hand

Figure 2.4.7. Skeletal representation for: A, small size category and B, medium size category

Encore (small; n=2398)

Ringtail (small; n=587) Camel Sputum (small; n=2630)

Mike's Menagerie (small; n=183)

Bitesantennary (small; n=4599)

Quantum Leap (small; n=780)

1 2 Expected 3

Site (size; number in sample) number in (size; Site Camel Sputum (medium; n=472)

Quantum Leap (medium; n=126)

Ringtail (medium; n=41) Encore (medium; n=336)

Mike's Menagerie (medium; n=322) 0% 20% 40% 60% 80% 100%

Figure 2.4.8. Hydraulic transport potential groupings for elements from fossil sites

CHAPTER 2

make up the greater numbers in groups 2 and 3 for those categories. The majority of elements from Bitesantennary are from bats. Certain bat vertebrae are fused, so the proportion of vertebrae in a bat skeleton would be slightly less than in the average marsupial skeleton (upon which the expected numbers are based). Also, bat vertebrae are very small and taxonomically uninformative so there may be a collection bias against them. This may explain the slight underrepresentation of group 1 elements from

Bitesantennary. For the other sites, this may be due to a collection/sorting bias again, or may reflect the varying susceptibilities of skeletal elements to destruction (caused by bioturbation or exposure) rather than indicating fluvial transport (see below for further discussion of the extent of fragmentation of different elements). If, however, the lower- than-expected number of group 1 elements is taken as evidence of fluvial transport, then the fossil samples from these sites represent the untransported, or lag, component which is authochtonous.

Encore small, Ringtail small and Mike’s Menagerie medium categories show a considerable underrepresentation of group 1 elements. In the case of Ringtail Site, interpreted to be pool deposits, this may be explained by the depositional environment.

This deposit potentially had a weak flow of water, affecting mainly small sized elements. If the number of group 1 elements is taken as evidence of fluvial transport, then fossil samples from Ringtail and Encore site deposits represent the less transported, potentially untransported, lag component which is likely to be autochthonous. The

Mike’s Menagerie situation is harder to explain because evidence from the medium category is not consistent with the small category. If water moves group 1 elements of the medium size category, it would surely move group 1 elements of the small size

CHAPTER 2

category to a greater extent. Perhaps there is a link with Camel Sputum’s medium category which has more elements in group 1 than expected. As previously noted,

Mike’s Menagerie and Camel Sputum are in close proximity and are linked by a continuous fossiliferous deposit. Work in progress is investigating whether these two sites are in fact parts of the same deposit. If this is the case, deviations exhibited by

Camel Sputum and Mike's Menagerie medium size categories will not be so large when data for the two sites are combined. Further sampling of Mike’s Menagerie site may also clarify the issue.

Riversleigh Oligocene-Pliocene terrestrial predators include mammalian and reptilian taxa (Wroe 2002) and birds of prey (Boles 1993). There are considerably fewer terrestrial reptilian predator species than there are mammalian predator species (Wroe

2002) and reptilian predators are less likely to leave evidence of their predation. Prey assemblages may be recognised by the size of prey, species composition, and modification of the bones of prey, which may all be predator-specific (Andrews 1990).

At this stage, the six fossil sites are not interpreted to be the result of predator accumulations based on lack of bone modification (such as rounding, digestion and tooth marks), skeletal representation, and taxonomic and size variation. There is no evidence of digestion nor obvious tooth marks on the majority of specimens from the six sites (Table 2.4.1). Taxonomic diversity and size range at each site is broad (for example, Camel Sputum marsupials vary in size from the tiny Burramys brutyi with an approximate body mass of 21g, through to Neohelos tirarensis with a body mass of approximately 124kg, estimated using the body mass equations of Myers, 2001; also, bats from Camel Sputum would have weighed approximately 5 g, S. Hand pers. comm.

2005). Boles (2000) has found no evidence of owl predator accumulations at

CHAPTER 2

Site % digestion % tooth marks Total sample size Camel Sputum 0.23 0.51 3550 Mike’s Menagerie 1.72 0.86 581 Ringtail 0.12 0.5 806 Quantum Leap 1.36 1.36 1107 Encore 0.4 0.43 3001 Bitesantennary 0 0.01 8086

Table 2.4.1. Percentage of specimens (of all size categories) exhibiting evidence of possible digestion or tooth marks (with corresponding examples of each pictured; scale bar = 2 cm)

Riversleigh yet. Also the age-class distribution of bandicoots at Camel Sputum

(Fig. 2.4.9. A) does not follow the pattern for owl prey accumulation shown by the

Jurien Caves dasyurids. For medium-sized to large mammals, the assemblages preserve many long bones with their shafts intact and with epiphyses present; skeletal representation is not consistent with that found at the kill sites of medium-sized to large mammalian predators, which leave mostly phalanges, metapodials/tarsals, teeth and fragments of shafts of long bones (Haynes 1982); and skeletal representation is not consistent with that found at carnivore feeding places and dens, which preserve a majority of limb and girdle bones (Arribas and Palmqvist 1998; Palmqvist and Arribas

2001) and display a high frequency of gnawing marks on cranial, limb and girdle elements, and on preserved epiphyses (Palmqvist et al. 1996).

Further work is necessary to determine if Macroderma (genus of predatory bat) was involved in accumulation of small taxa at the six sites. The modern ghost bat,

Macroderma gigas, is opportunistic in its hunting habits, taking as many as 40 species of bird (Boles 1999), a broad range of small mammals (Douglas 1967), and other

CHAPTER 2

A 60

40 Jurien Caves dasyurids, n=74 30 Camel Sputum peramelids, n=82

123456 Tooth wear category

60 B

Catastrophic mortality 40 profile, n=63 (equivalent to numbers of living bandicoots; from Dufty 1994) 30 Attritional mortality profile (derived from Dufty 1994)

1234567

Age class (3 months)

Figure 2.4.9. Mortality profiles for: A, dasyurids from Jurien Caves owl pellet material and bandicoots from Camel Sputum; and B, Perameles gunni from Hamilton, Victoria

CHAPTER 2

vertebrates and invertebrates to be eaten in roost caves (Boles 1999). Very few specimens of Macroderma have been identified from the six Riversleigh sites

(maximum of three specimens at each site), suggesting the sites do not represent

Macroderma roosts. To determine if Macroderma is in fact responsible for accumulation of any remains at the sites, fossils of birds, frogs and reptiles - potential prey for Macroderma - will need to examined in addition to mammalian fossils.

Results of the age-class distribution analysis for Camel Sputum bandicoots (Fig. 2.4.9.

A) are shown along with the age-class distribution for dasyurids (several species of varying sizes from the genera Phascogale, Planigale, Sminthopsis, Parantechinus and

Pseudantechinus) from the ?late Cainozoic/?Quaternary Jurien Caves owl pellet accumulation. The dasyurids exhibit a catastrophic mortality profile because their predator (potentially Tyto alba, Tyto novaehollandiae or Ninox connivens, based on modern owl distributions) would have selected for size, not for age of individuals (as in the case of T. novaehollandiae, Mooney 1993), thus taking a random sample from the living population. Catastrophic death curves reflect the numbers of living individuals for each age class in the population. Attritional death curves, on the other hand, reflect the number of individuals for each age class of a population, that are dying - i.e. there is high mortality of juveniles, mortality decreases for adults and then increases for aged individuals - (Voorhies 1969, Lyman 1994). Camel Sputum bandicoots seem to follow the pattern of attritional mortality, with a high proportion of juveniles (tooth wear category 1), a relatively low proportion of adults (categories 2-4), an increase from adults to aged individuals (category 5), and total die off (category 6 ); also see

Fig. 2.4.9. B for comparison, but note that age classes do not correspond directly to tooth wear categories for Fig. 2.4.9. A.

CHAPTER 2

Age distribution for euro/wallaby population from Fowlers Gap, NSW

30 Catastrophic mortality profile 25 (equivalent to numbers of living euros/wallabies; from Dawson 1995) 20 Attritional mortality profile

Percent (derived from Dawson 1995) 15

0 0-2 2-4 4-8 8-12 12-16 16-20+ Age class (years)

B 35

Percent 15

0 123456 Tooth wear category

Figure 2.4.10. Mortality profiles for: A, extant euro/wallaby populations from Fowlers Gap, NSW; B, fossil species Bulungamaya delicata and Wabularoo naughtoni from Camel Sputum

CHAPTER 2

An age-class distribution analysis of the kangaroos from Camel Sputum revealed a similar pattern. Fig. 2.4.10. B shows a distinct increase in frequency of individuals from tooth wear category 5 to category 6. This is consistent with an attritional mortality pattern as seen in Fig. 2.4.10. A for modern macropodids. The attritional curve in

Fig. 2.4.10. A begins at the 8-12 year period because 0-4 year olds have not been caught in the correct proportion (see Dawson 1995). For further contrast with catastrophic mortality profiles, four species of rodents (identified by H. Godthelp, 2002) from Jurien

Caves owl pellet deposit are shown (Fig. 2.4.11) – individually and lumped together; the pattern is similar regardless. The very low frequency of Rattus fuscipes individuals in tooth wear categories 1 and 2 is probably a result of different habitat use by young individuals compared to adults. Lunney (1998) notes that the young of R. fuscipes forage briefly and spend most of their time in burrows.

80 Pseudomys albocinereus (n=71) 70 Notomys cf. mitchelli (n=42) 60 Pseudomys shortridgei 50 and Rattus tunney (n=324) Rattus fuscipes (n=47) Percent 40

30 total of all species (n=484) 20

0 12345 Tooth wear category

Figure 2.4.11. Jurien Caves rodent age distributions

CHAPTER 2

Assemblages accumulated over long time periods (attritional mortality profile), that are not the result of predator activity, typically exhibit high taxonomic diversity

(Varricchio and Horner 1993); assemblages with a catastrophic mortality profile may or may not exhibit high taxonomic diversity, depending on whether accumulation was instantaneous, over a period of time such as for pitfall trap deposits, or the result of certain predators. Instantaneous, mass mortality events are often taxon-specific in their preservation (Behrensmeyer et al. 2000); pitfall trap accumulations may exhibit high taxonomic diversity; and owl pellet accumulations, for certain species, do not exhibit high taxonomic diversity (Andrews 1990; Dodson and Wexlar 1979). Due to high taxonomic diversity, Ringtail, Encore, Bitesantennary, Quantum Leap and Mike’s

Menagerie deposits are presumed to have accumulated over time and not to be due to instantaneous mortality events. It is expected that Ringtail Site, being pool deposits, will not exhibit the catastrophic mortality profile of a pitfall trap, but a greater sample size is necessary for this to be demonstrated. The fauna from Camel Sputum Site exhibits an attritional mortality profile and suggests Camel Sputum is not a pitfall trap deposit and this perhaps also will apply to Mike’s Menagerie if it is shown to be a part of the same deposit. Creaser (1997) states that the lithology of the sediments and fossils at Camel

Sputum and Mike’s Menagerie suggest that these deposits accumulated over a period of time.

Time-averaging at the six sites is not expected to be long-term because animals at different evolutionary stages are not mixed and preservation quality is consistent within each site. Kidwell and Flessa (1996) describe extensively time-averaged samples as having ‘cumulative destruction’, where older remains, mixed with more recent remains,

CHAPTER 2

exhibit greater modification due to longer exposure to postmortem destructive processes. Short-term time-averaging, in the order of up to several thousand years

(Fürsich and Aberhan 1990), is possible for the sites examined here. Surface terrestrial environments tend to exhibit higher temporal resolutions than cave deposits which have low sediment input, however, it seems that in the majority of cases looked at, fossil mammal assemblages do not contain more than one million years of time-averaging

(Kidwell and Flessa 1996). For Riversleigh sites, it is difficult to determine exactly to what extent the sites are time-averaged. Arena (2004) notes that fluvial barrage infill deposits, such as the Ringtail Site deposit, may have depositional time spans ranging between 100 and 1 000 years; also cave deposits may range from 1 000 to over

1 000 000 years in depositional time.

2.5. Summary

Palaeoecological studies of the local faunas from Camel Sputum, Mike’s Menagerie,

Ringtail, Encore, Quantum Leap and Bitesantennary Sites can be carried out in future with consideration of the taphonomic biases affecting remains at each site.

The Bitesantennary local fauna is taxonomically biased, containing mostly bats, so further palaeoecological study will have to be restricted to bat communities.

Skeletal element associations at Quantum Leap and Encore, and lack of abrasion of specimens from all sites suggest that water is not a significant transport agent for these samples of specimens. The general over-representation of dental elements for all sites may be the result of a collecting or sorting bias and/or may reflect the lower susceptibility of these elements to destruction; if in any case it is due to water flow, the

CHAPTER 2

sample represents the non-transported component so palaeoecological studies will not be affected.

The lack of surface deterioration and the break patterns of limb bones from all sites indicate minimal aerial exposure of bones prior to burial and therefore reduced risk of bones being moved considerable distances. Burial of bones at all the sites is likely to have occurred in wet or moist conditions (either underwater or in wet microenvironments), as suggested by the extent of disarticulation combined with minimal weathering. Disassociation of the majority of elements at all sites suggests bioturbation of pre-fossilised burial sediment by churning or mixing caused by the movement of live organisms (Behrensmeyer et al. 1989).

The involvement of Macroderma as an accumulator of small vertebrates seems unlikely due to the small number of Macroderma specimens recovered from the sites, however, further analysis is required to completely rule out this possibility. At this stage there appear to be no significant predator/scavenger-caused biases.

Bandicoots from Camel Sputum Site exhibit an attritional mortality profile and the remaining sites are presumed to have accumulated over time and not to represent instantaneous mortality events. Accumulation at all sites is considered to be autochthonous, but exact duration of accumulation can not be ascertained.

CHAPTER 2

2.6. References

Andrews, P., 1990. Owls, Caves and Fossils. Natural History Museum Publications. London.

Archer, M., 1979. Wabularoo naughtoni gen. et sp. nov., an enigmatic kangaroo (Marsupialia) from the middle Tertiary Carl Creek limestone of northwestern Queensland. Memoirs of the Queensland Museum 19 (3): 299-307.

Archer, M. and Flannery, T., 1985. Revision of the extinct gigantic rat kangaroos (Potoroidae: Marsupialia), with description of a new Miocene genus and species and a new Pleistocene species of Propleopus. Journal of Paleontology 59 (6): 1331-1349.

Archer, M., Hand, S. J. and Godthelp, H., 1988. A new Order of Tertiary zalambdodont marsupials. Science 239: 1528-1531.

Archer, M., Jenkins, F., Hand, S. J., Murray, P. and Godthelp, H., 1992. Description of the skull and non-vestigial dentition of a Miocene platypus (Obdurodon dicksoni n.sp.) from Riversleigh, Australia, and the problem of monotreme origins. Pp. 15-27 in Platypus and echidnas. Ed. M. Augee. The Royal Zoological Society of NSW. Sydney.

Arena, D. A., 2004. The geological history and development of the terrain at the Riversleigh World Heritage Area during the middle Tertiary. PhD thesis. University of New South Wales, Sydney.

Arribas, A. and Palmqvist, P., 1998. Taphonomy and palaeoecology of an assemblage of large mammals: hyaenid activity in the lower Pleistocene site at Venta Micena (Orce, Guadix-Baza Basin, Granada, Spain). Geobios 31(3) supplement: 3-47.

Badgley, C., 1986. Counting individuals in mammalian fossil assemblages from fluvial environments. Palaios 1: 328-38.

Bassarova, M., Archer, M. and Hand, S.J., 2001. New species of ringtail possums (Pseudocheiridae, Marsupialia) referable to the Oligo-Miocene genus Paljara from deposits of the Riversleigh World Heritage property, northwestern Queensland. Memoirs of the Association of AustralasianPalaeontologists 25: 61-75.

Behrensmeyer, A. K., 1975. The taphonomy and paleoecology of Plio-Pleistocene vertebrate assemblages of Lake Rudolf, Kenya. Bulletin of the Museum of Comparative Zoology 146: 473-578.

Behrensmeyer, A. K., 1978. Taphonomic and ecological information from bone weathering. Paleobiology 4: 150-62.

Behrensmeyer, A. K., 1984. Taphonomy and the fossil record. American Scientist 72: 558-566.

Behrensmeyer, A. K., 1991. Terrestrial vertebrate accumulations. Pp. 291-335 in Taphonomy – Releasing the Data Locked in the Fossil Record. Eds P. A. Allison and D. E. G. Briggs. Plenum Press. New York.

CHAPTER 2

Behrensmeyer, A. K., Gordon, K. D. and Yanagi, G. T., 1989. Nonhuman bone modification in Miocene fossils from Pakistan. Pp. 99-120 in Bone Modification. Eds R. Bonnichsen, and M. H. Sorg. Centre for the Study of the First Americans. Orono, Maine.

Behrensmeyer, A. K. and Kidwell, S. M., 1985. Taphonomy’s contributions to paleobiology. Paleobiology 11: 105-119.

Behrensmeyer, A. K., Kidwell, S. M. and Gastaldo, R. A., 2000. Taphonomy and paleobiology. Paleobiology 26(4) supplement: 103-147.

Black, K., 1997. A new species of Palorchestidae (Marsupialia) from the late middle to early late Miocene Encore Local Fauna, Riversleigh, northwestern Queesland. Memoirs of the Queensland Museum 41(2): 181-185.

Black, K. and Archer, M., 1997. Nimiokoala gen. nov. (Marsupialia, Phascolarctidae) from Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41(2): 209-228.

Boles, W. E., 1993. Pengana robertbolesii, a peculiar bird of prey from the Tertiary of Riversleigh, northwestern Queensland, Australia. Alcheringa 17: 19-25.

Boles, W. E., 1999. Avian prey of the Australian Ghost Bat, Macroderma gigas (Microchiroptera: Megadermatidae): prey characteristics and damage from predation. Australian Zoologist 31: 82-91.

Boles, W. E., 2000. Investigations on Australian Tertiary avifauna with an emphasis on the fossil birds of Riversleigh, northwestern Queensland. PhD thesis. University of New South Wales. Sydney.

Brammall, J. R., 1998. A new petauroid possum from the Oligo-Miocene of Riversleigh, northwestern Queensland. Alcheringa 23: 31-50.

Brammall, J. and Archer, M., 1997. A new Oligo-Miocene species of Burramys (Marsupialia, Burramyidae) from Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41(2): 247-268.

Cadee, G. C., 1991. The history of taphonomy. Pp. 3-21 in The processes of fossilization. Ed. S. K. Donovan. Belhaven Press, London.

Clemens, W. A. and Plane, M., 1974. Mid-Tertiary Thylacoleonidae (Marsupialia, Mammalia). Journal of Paleontology 48: 652-660.

Coe, M., 1980. The role of modern ecological studies in the reconstruction of paleoenvironments in sub-Saharan Africa. Pp. 55-71 in Fossils in the Making. Eds A. K. Behrensmeyer and A. P. Hill. University of Chicago. Chicago.

Cooke, B. N., 1992. Primitive macropodids from Riversleigh, northwestern Queensland. Alcheringa 16: 201-217.

CHAPTER 2

Cooke, B. N., 1997. New Miocene bulungamayine kangaroos (Marsupialia, Potoroidae) from Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41(2): 281-294.

Cooke, B.N., 1999. Wanburoo hilarus gen. et sp. nov., a lophodont bulungamayine kangaroo (Marsuplialia: Macropodoidea: Bulungamayinae) from the Miocene records of Riversleigh, northwestern Queensland. Records of the Western Australian Museum Supplement 57: 239-253.

Coombs, M. C. and Coombs, W. P., 1997. Analysis of the geology, fauna, and taphonomy of Morava Ranch Quarry, early Miocene of Northwest Nebraska. Palaios 12: 165-187.

Creaser, P., 1997. Oligocene-Miocene sediments of Riversleigh: the potential significance of topography. Memoirs of the Queensland Museum 41(2): 303-314.

Crosby, K., Nagy, M. and Archer M., 2001. Wyulda asherjoeli, a new phalangerid (Diprotodontia: Marsupialia) from the early Miocene of Riversleigh, northwestern Queensland. Memoirs of the Association of Australasian Palaeontologists 25: 77-82.

Cutler, A. H., Behrensmeyer, A. K. and Chapman, R. E., 1999. Environmental information in a recent bone assemblage: roles of taphonomic processes and ecological change. Palaeogeography, Palaeoclimatology, Palaeoecology 149: 359-372.

Dawson, T. J. 1995. Kangaroos – Biology of the Largest Marsupial. University of New South Wales Press Ltd., Sydney.

De Oliveira, J. A., Strauss, R. E. and dos Reis, S. F. 1998. Assessing relative age and age structure in natural populations of Bolomys lasiurus (Rodentia: Sigmodontinae) in northeastern Brazil. Journal of Mammalogy 74(4): 1170-1183.

Dodson, P., 1973. The significance of small bones in paleoecological interpretation. Contributions to Geology 12: 15-19.

Dodson, P. and Wexlar, D., 1979. Taphonomic investigations of owl pellets. Paleobiology 5: 275-284.

Douglas, A. M., 1967. The natural history of the Ghost Bat, Macroderma gigas (Microchiroptera, Megadermatidae), in Western Australia. Western Australian Naturalist 10: 125-138.

Dufty, A. C., 1994. Population demography of the Eastern Barred Bandicoot (Perameles gunni) at Hamilton, Victoria. Wildlife Research 21: 445-457.

Fieller, N. R. J. and Turner, A., 1982. Number estimation in vertebrate samples. Journal of Archaeological Science 9: 49-62.

CHAPTER 2

Flannery, T.F. and Archer, M., 1987. Strigocuscus reidi and Trichosurus dicksoni, two new fossil phalangerids (Marsupialia: Phalangeridae) from the Miocene of northwestern Queensland. Pp. 527-536 in Possums and Opossums: Studies in Evolution. Ed. M. Archer. Surrey Beatty & Sons and Royal Zoological Society of NSW. Sydney.

Flannery, T., Archer, M. and Plane, M., 1982. Middle Miocene kangaroos (Macropodoidea: Marsupialia) from three localities in northern Australia, with a description of two new subfamilies. BMR. Journal of Australian Geology & Geophysics 7: 287-302.

Fürsich, F. T. and Aberhan, M., 1990. Significance of time-averaging for palaeocommunity analysis. Lethaia 23: 143-152.

Grayson, D. K., 1978. Minimum numbers and sample size in vertebrate faunal analysis. American Antiquity 43: 53-65.

Grayson, D. K., 1984. Quantitative Zooarchaeology. Academic Press Inc., Orlando.

Hadly, E. A., 1999. Fidelity of terrestrial vertebrate fossils to a modern ecosystem. Palaeogeography, Palaeoclimatology, Palaeoecology 149: 389-409.

Hand, S., 1997. New Miocene leaf-nosed bats (Michrochiroptera: Hipposideridae) from Riversleigh, Northwestern Queensland. Memoirs of the Queensland Museum 41(2): 335-349.

Hand, S., 1998a. Xenorhinos, a new genus of old world leaf-nosed bats (Microchiroptera: Hipposideridae) from the Australian Miocene. Journal of Vertebrate Paleontology 18 (2): 430-439.

Hand, S., 1998b. Riversleigha williamsi gen. et sp. nov., a large Miocene hipposiderid (Microchiroptera) from Riversleigh, Queensland. Alcheringa 22: 259-276.

Hand, S. J., Archer, M., Godthelp, H., Rich, T. and Pledge, N., 1993. Nimbadon, a new genus and three new species of tertiary zygomaturines (Marsupialia: Diprotodontidae) from northern Australia, with a reassessment of Neohelos. Memoirs of the Queensland Museum 33(1): 193-210.

Haynes, G., 1982. Utilization and skeletal disturbance of North American Prey Carcasses. Arctic 35: 266-281.

Hill, A. P., 1979. Disarticulation and scattering of mammal skeletons. Paleobiology 3: 261-274.

Holtzman, R. C., 1979. Maximum likelihood estimation of fossil assemblage composition. Paleobiology 5: 77-89.

Johnston, P. G., 1998. Long-nosed potoroo.Pp. 301-302 in The Mammals of Australia. Ed. R. Strahan. New Holland Publishers Pty Ltd. Sydney.

CHAPTER 2

Kear, B. P., Archer, M. and Flannery, T. F., 2001. Bulungamayine (Marsupialia: Macropodidae) postcranial elements from the late Miocene of Riversleigh, northwestern Queensland. Memoirs of the Association of Australasian Palaeontologists 25: 103-122.

Kear, B. P. 2002. Phylogenetic implications of Macropodid (Marsupialia: Macropodoidea) postcranial remains from Miocene deposits of Riversleigh, northwestern Queensland. Alcheringa 26: 299-318.

Kidwell, S. M. and Flessa, K. W., 1996. The quality of the fossil record: populations, species and communities. Annual Review of Earth and Planet Science 24: 433-464.

Kirkham, Z. 2004. The Cranial Description of the Primitive Macropodid Rhizosthenurus flanneryi and its Phylogeny Based on Cranial and Postcranial Characters. Honours Thesis. University of New South Wales, Sydney.

Korth, W. W., 1979. Taphonomy of microvertebrate fossil assemblages. Annals of the Carnegie Museum 48: 235-285.

Kowalewski, M., 1996. Time-averaging, overcompleteness, and the geological record. The Journal of Geology 104: 317-326.

Kowalewski, M., 1997. The reciprocal taphonomic model. Lethaia 30: 86-88.

Krantz, G. S., 1968. A new method of counting mammal bones. American Journal of Archaeology 72: 286-288.

Lentle, R. G., Stafford, K. J., Potter, M. A., Springett, B. P. and Haslett, S. 1998. Incisor and molar wear in the tammar wallaby (Macropus eugenii Desmarest). Australian Journal of Zoology 46: 509-527.

Lunney, D. 1998. Bush rat, Rattus fuscipes. Pp. 651-653 in The mammals of Australia. Ed. R. Strahan. New Holland Publishers Pty Ltd, Sydney.

Lyman, R. L., 1994. Vertebrate Taphonomy. Cambridge University Press. Great Britain.

McArthur, C. and Sanson, G. D. 1988. Tooth wear in eastern grey kangaroos (Macropus giganteus) and western grey kangaroos (Macropus fuliginosus), and its potential influence on diet selection, digestion and population parameters. Journal of Zoology London 215: 491-504.

Martill, D. M., 1991. Bones as stones: the contribution of ertebrate remains to the lithologic record. Pp. 270-292 in The processes of fossilization. Ed. S. K. Donovan. Belhaven Press, London.

Martin, R. E. 1999. Taphonomy. A process approach. Cambridge University Press, Cambridge.

Merchant, J. C. 1998. Swamp wallaby. Pp. 404-405 in The Mammals of Australia. Ed. R. Strahan. New Holland Publishers Pty Ltd. Sydney.

CHAPTER 2

Mooney, N. 1993. Diet of the masked owl in Tasmania: past and present. Pp. 160-174 in Australian raptor studies. Ed. P. Olsen. Australasian Raptor Association, R.A.O.U., Melbourne.

Muirhead, J. 1992. A specialised thylacinid, Thylacinus macknessi, (Marsupialia: Thylacinidae) from Miocene deposits of Riversleigh, northwestern Queensland. Australian Mammalogy 15: 67-76.

Muirhead, J. 1997. Two new early Miocene thylacines from Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41(2): 367-377.

Muirhead, J. and Filan, S., 1995. Yarala burchfieldi, a plesiomorphic bandicoot (Marsupialia, Peramelemorphia) from Oligo-Miocene deposits of Riversleigh, northwestern Queensland. Journal of Paleontology 69(1): 127-134.

Murray, P. 1990. Primitive marsupial tapirs (Propalorchestes novaculacephalus Murray and P. ponticulus sp. nov.) from the mid-Miocene of north Australia (Marsupialia: Palorchestidae). The Beagle, Records of the Northern Territory Museum of Arts and Sciences 7: 39-51.

Myers, T. J. 2001. Prediction of marsupial body mass. Australian Journal of Zoology 49: 99-118.

Myers, T. J., Crosby, K., Archer, M. and Tyler, M. 2001. The Encore Local Fauna, a late Miocene assemblage from Riversleigh, northwestern Queensland. Memoirs of the Association of Australasian Palaeontologists 25: 147-154.

Olszewski, T. 1999. Taking advantage of time-averaging. Paleobiology 25: 226-238.

Palmqvist, P. and Arribas, A. 2001. Taphonomic decoding of the paleobiological information locked in a lower Pleistocene assemblage of large mammals. Paleobiology 27(3): 512-530.

Palmqvist, P., Martinez-Navarro, B. and Arribas, A., 1996. Prey selection by terrestrial carnivores in a lower Pleistocene paleocommunity. Paleobiology 22(4): 514-534.

Peterson, C. H., 1977. The paleoecological significance of undetected short-term temporal variability. Journal of Paleontology 51: 976-981.

Saunders, J. J., 1977. Late Pleistocene vertebrates of the western Ozark Highland, Missouri. Illinois State Museum Reports of Investigations No. 33, Springfield.

Shipman, P., 1981. Life History of a Fossil - an Introduction to Taphonomy and Paleoecology. Harvard University. Cambridge.

CHAPTER 2

Shotwell, J. A. 1955. An approach to the paleoecology of mammals. Ecology 36: 327- 337.

Springer, M. S., 1987. Lower molars of Litokoala (Marsupialia: Phascolarctidae) and their bearing on phascolarctid evolution. Pp. 319-325 in Possums and Opossums: studies in evolution. Ed. M. Archer. Surrey Beatty and Sons and Royal Zoological Society of NSW. Sydney.

Varricchio, D. J. and Horner, J. R., 1993. Hadrosaurid and lambeosaurid bone beds from the Upper Cretaceous Two Medicine Formation of Montana: taphonomic and biologic implications. Canadian Journal of Earth Science 30: 997-1006.

Voorhies, M. R., 1969. Taphonomy and population dynamics of an early Pliocene vertebrate fauna, Knox County, Nebraska. University of Wyoming, Contributions to Geology Special paper No. 1: 1-69.

Woodburne, M. O., Tedford, R. H. and Archer, M., 1987. New Miocene ringtail possums (Marsupialia: Pseudocheiridae) from South Australia. Pp. 639-679 in Possums and Opossums: studies in evolution. Ed. M. Archer. Surrey Beatty and Sons and Royal Zoological Society of NSW. Sydney.

Wroe, S., 1996. An investigation of phylogeny in the giant extinct rat kangaroo Ekaltadeta (Propleopinae, Potoroidae, Marsupialia). Journal of Paleontology 70 (4): 681-690.

Wroe, S., 1997. Mayigriphus orbus gen. et sp. nov., a Miocene dasyuromorphian from Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41 (2): 439-448.

Wroe, S., 1998. A new ‘bone – cracking’ dasyurid (Marsupialia), from the Miocene of Riversleigh, northwestern Queensland. Alcheringa 22: 277-284.

Wroe, S., 1999. The geologically oldest dasyurid (Marsupialia) from the Miocene of Riversleigh, nothwestern Queensland. Palaeontology 42: 501-527.

Wroe, S., 2001. Maximucinus muirheadae, gen. et. sp. nov. (Thylacinidae: Marsupialia), from the Miocene of Riversleigh, northwestern Queensland, with estimates of body weights for fossil thylacinids. Australian Journal of Zoology 49: 603-614.

Wroe, S., 2002. A review of the relative significance of mammalian and reptilian carnivores in late Oligocene-Pleistocene Australia: the myth of reptilian domination. Australian Journal of Zoology 50: 1-24.

CHAPTER 2

Appendix A1. Raw data for percentage of various break shapes on limb bones for all size categories for the sites studied. Bites = Bitesantennary Site; MM = Mike’s Menagerie Site.

Breakage Bites Ringtail Encore Quantum Leap MM Camel Sputum spiral 41.18 66.67 28.57 33.33 45.45 40.2 transverse 29.41 25 50 39.22 45.45 44.4 longitudinal 16.47 0 7.14 7.84 0 4.3 stepped 12.94 8.33 14.29 19.61 9.09 11.1

CHAPTER 2

Expected Bite (small) Ring (small) Ring (med) Ec (small) Ec (med) MM (small) MM (med) CS (small) CS (med) CS (large) QL (small) QL (med) foot/hand 34.31 3.31 8.09 36.96 10.42 34.12 15 13.02 14 8.2 40.85 23.06 29.73 vertebrae 22.55 11.66 4.19 19.57 1.64 20 17.5 2.96 11.37 29.2 28.17 20.62 18.92 teeth 19.12 27.44 54.48 21.74 73.06 36.18 41.5 69.23 50.95 20.3 16.9 31.93 3.78 ribs 12.75 1.87 0.14 6.52 0.15 0 0 0.3 0.73 0.7 0 2.66 20 limbs 5.88 8.64 6.94 6.52 0.27 4.71 5.5 3.25 4.06 5 2.82 3.88 12.43 dentaries 1 5.45 10.98 4.35 6.1 3.53 12 6.51 10.61 16 1.41 6.1 1.08 scapulae 1 0.41 0.14 0 0 0.29 0 0 0.14 0.5 1.41 0.55 1.62 periotics 1 39.17 9.83 0 9.04 1.18 2.5 0.89 3.88 4.4 2.82 0.89 0 cranium 0.5 1.31 0.14 2.17 0.04 0 0.5 0 0.31 0.7 0 6.65 0.5 maxilla 0.5 0.71 4.48 2.17 0.42 0 4.5 2.37 3.36 13.4 2.82 1.77 2.16 pelvis 0.5 0.01 0.58 0 0.23 0 1 1.18 0.55 1.5 2.82 1.55 9.19 sacrum 0.5 0.21 0 0 0 0 0 0.3 0.03 0 0 0 0.5 sternum 0.5 0 0 0 0 0 0 0 0 0 0 0.33 0

APPENDIX A2. Raw data (percentages) for skeletal representation for small, medium and large size categories for the fossil sites studied and for the expected proportions. Abbreviations are as follows: Bite = Bitesantennary Site; Ring = Ringtail Site; Ec = Encore Site; MM = Mike’s Menagerie Site; CS = Camel Sputum Site; QL = Quantum Leap Site. Total sample sizes (n) used for each site are as follows:

Bite (small), n = 8078 Ring (small), n = 692 Ring (med), n = 46 Ec (small), n = 2621 Ec (med), n = 340 MM (small), n = 200 MM (med), n = 338 CS (small), n = 2885 CS (med), n = 595 CS (large), n = 71 QL (small), n = 902 QL (med), n = 185

CHAPTER 2

APPENDIX A3. Specimens used from Camel Sputum site for age distribution analysis

Specimen Tooth/teeth Taxon Tooth wear category registration used Macropodids Bandicoots number AR 9804 lowers Bulungamaya delicata 1 AR 9934 uppers Bulungamaya delicata 1 AR 11460 uppers Bulungamaya delicata 2 AR 14480 uppers Bulungamaya delicata 2 AR 15315 lowers Bulungamaya delicata 3 AR 16495 uppers Bulungamaya delicata 0 AR 16498 uppers Bulungamaya delicata 1 AR 16670 uppers Bulungamaya delicata 5 AR 16807 uppers Bulungamaya delicata 2 AR 16911 uppers Bulungamaya delicata 5 AR 16914 lowers Bulungamaya delicata 1 AR 17066 uppers Bulungamaya delicata 1 AR 17075 uppers Bulungamaya delicata 2 AR 17273 lowers Bulungamaya delicata 0 AR 9939 lowers Wabularoo naughtoni 1 AR 9954 lowers Wabularoo naughtoni 3 AR 10359 lowers Wabularoo naughtoni 4 AR 10983 lowers Wabularoo naughtoni 2 AR 15748 uppers Wabularoo naughtoni 5 AR 16644 lowers Wabularoo naughtoni 2 AR 16762 uppers Wabularoo naughtoni 2 AR 16910 uppers Wabularoo naughtoni 3 AR 17002 lowers Wabularoo naughtoni 5 AR 17065 lowers Wabularoo naughtoni 1 AR 17457 lowers Wabularoo naughtoni 2 QMF 20496 uppers Wabularoo naughtoni 1 QMF 23487 lowers Wabularoo naughtoni 3 AR 9008 M1 Peramelemorphia 3 AR 10070 M1 Peramelemorphia 5 AR 10077 M1 Peramelemorphia 5 AR 10078 M1 Peramelemorphia 2 AR 10199 M1 Peramelemorphia 2 AR 10549 M1 Peramelemorphia 2 AR 10878 M1 Peramelemorphia 3 AR 10879 M1 Peramelemorphia 1 AR 10880 M1 Peramelemorphia 5 AR 10883 M1 Peramelemorphia 3 AR 10884 M1 Peramelemorphia 5 AR 10885 M1 Peramelemorphia 1 AR 10897 M1 Peramelemorphia 2 AR 10960 M1 Peramelemorphia 1 AR 11456 M1 Peramelemorphia 1 AR 11458 M1 Peramelemorphia 1 AR 11623 M1 Peramelemorphia 1 AR 11633 M1 Peramelemorphia 1 AR 11636 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 6 AR 11662 M1 Peramelemorphia 5 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 2 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 1

CHAPTER 2

Specimen Tooth/teeth Taxon Tooth wear category registration used Macropodids Bandicoots number AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 2 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 2 AR 11662 M1 Peramelemorphia 3 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 2 AR 11662 M1 Peramelemorphia 2 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 1 AR 11662 M1 Peramelemorphia 2 AR 11662 M1 Peramelemorphia 1 AR 11772 M1 Peramelemorphia 5 AR 11861 M1 Peramelemorphia 1 AR 12794 M1 Peramelemorphia 3 AR 12861 M1 Peramelemorphia 1 AR 12862 M1 Peramelemorphia 3 AR 12863 M1 Peramelemorphia 4 AR 12882 M1 Peramelemorphia 4 AR 12950 M1 Peramelemorphia 4 AR 13018 M1 Peramelemorphia 4 AR 13909 M1 Peramelemorphia 3 AR 13910 M1 Peramelemorphia 3 AR 13917 M1 Peramelemorphia 5 AR 13918 M1 Peramelemorphia 3 AR 13923 M1 Peramelemorphia 1 AR 13924 M1 Peramelemorphia 5 AR 13927 M1 Peramelemorphia 1 AR 15707 M1 Peramelemorphia 5 AR 15740 M1 Peramelemorphia 4 AR 15742 M1 Peramelemorphia 1 AR 15743 M1 Peramelemorphia 6 AR 16459 M1 Peramelemorphia 1 AR 16525 M1 Peramelemorphia 3 AR 16622 M1 Peramelemorphia 2 AR 16796 M1 Peramelemorphia 2 AR 16976 M1 Peramelemorphia 4 AR 17004 M1 Peramelemorphia 1 AR 17005 M1 Peramelemorphia 5 AR 17006 M1 Peramelemorphia 4 AR 17089 M1 Peramelemorphia 3 AR 17090 M1 Peramelemorphia 1 AR 17260 M1 Peramelemorphia 1 QMF 20578 M1 Peramelemorphia 1 QMF 20579 M1 Peramelemorphia 1 QMF 24227 M1 Peramelemorphia 1 QMF 29705 M1 Peramelemorphia 2

APPENDIX A3. Continued

CHAPTER 3

THE CALCANEUM… ON THE HEELS OF MARSUPIAL LOCOMOTION

CHAPTER 3

THE CALCANEUM – ON THE HEELS OF MARSUPIAL LOCOMOTION

3.1. Introduction

3.1.1. Locomotor behaviour

Locomotor behaviour is an ecological adaptation and as such, is useful in ecological and palaeontological animal community studies. The basic media or substrates available to mammals for locomotion are water (for aquatic mammals such as the platypus), air (for chiropterans), the ground (for mammals referred to here as ‘terrestrial’) or vegetation

(for arboreal and semi-arboreal, or ‘scansorial’, mammals). Species may be specialised for locomotion on only one substrate, or they may utilise more than one to varying degrees. The locomotor behaviour of mammalian species reflects their use of these substrates and therefore reflects habitat use and thus, at the animal community level, locomotor behaviour reflects general habitat/vegetation structure (Damuth 1992). For example, a large proportion of the mammalian fauna of tropical forests with a closed canopy is arboreal because of the reduced primary productivity on the shaded forest floor (Bourlière 1973, Eisenberg 1990).

A variety of ways exist for locomoting on each type of substrate. For ground and vegetation substrates (the focus of this study) some of the ways include: ‘cursorial’ locomotion, or quadrupedal running, and ‘saltatorial’ locomotion (bipedal hopping on hind-feet, as described by Bishop, 1997) on the ground; ‘fossorial’ locomotion, or digging through the ground; ‘grasp-climbing’ (as in phalangeroids) and ‘claw-climbing’

(as in dasyurids) on vegetation (collectively referred to here as ‘arboreal’, but discussed individually by Szalay, 1994). The extent of detail necessary, in terms of describing

61 CHAPTER 3

locomotor behaviour, varies with the particular aims of a study and is subject to the limitations of the available evidence.

For the study of palaeocommunities, morphology of fossil remains must be used to predict ecological aspects, such as locomotor behaviour, of the constituent fauna (Van

Valkenburgh 1987). This is indirect evidence (in contrast with behavioural observation) and as such limits the detail within the locomotor categories to an extent. One advantage of using broad categories is that they are widely applicable to many different communities thereby enabling comparisons to be made between these communities.

Skeletal features are related to, and can be used to infer, ecological adaptations such as locomotor behaviour. The direction and types of forces placed on bones vary with the type of locomotion. So, the form of skeletal elements reflects, to varying degrees, the loading placed on the skeleton by locomotion (Szalay 1994, Bishop 1997). Among the studies that demonstrate this are: Oxnard’s (1968) canonical analyses of forelimb and shoulder measurements, which distinguish arboreal and non-arboreal forms of locomotion in mammals; Walker’s (1974) investigation of correlation between skeletal anatomy and locomotion in primates; Jenkins and McClearn’s (1984) exploration of the relation between foot structure and locomotor use in climbing animals; Van

Valkenburgh’s (1987) multivariate analyses of locomotor behaviour of carnivore species; the biomechanical and multivariate analyses of macaque calcanea by Fa-Hong et al. (1993); Szalay’s (1994) research into morphological and functional relationships in marsupial pedal and tarsal elements; Janis et al.’s (2002) statistical comparisons of camelid and ruminant artiodactyl limb anatomy as it relates to locomotion; Argot’s

(2001) and Lemelin’s (1999) functional analyses of didelphid forelimbs and feet; Szalay

62 CHAPTER 3

and Sargis’ (2001) study of marsupial postcranial form-function relationships; and

Youlatos’ (2003) examination of primate calcaneal features.

3.1.2. Aims

The aims of this study were to find a relationship between the known locomotor behaviour of extant Australo-Papuan marsupial species and measurements of their calcanea and, if found, to use this relationship to determine the locomotor behaviour of fossil marsupial taxa from measurements of their calcanea. The fossil taxa are from

Camel Sputum, Encore, Mike’s Menagerie and Quantum Leap Sites. The marsupial calcaneum sample sizes from Bitesantennary and Ringtail sites are too small to consider.

Analogy is important in making hypotheses about the functional potential of extinct animals (Hickman 1988, Walker 1974). Because there is no direct information about the locomotor habits of extinct Riversleigh marsupials, this had to be deduced from analogy to modern relatives.

3.1.3. Why calcanea?

The calcaneum was chosen for this study for a number of reasons. Firstly, it appears to be the most robust and easily identifiable foot element and therefore provides the largest sample size for each fossil site compared to other elements of the foot. Secondly, the fact that it is a foot element is important because there is a relatively more rigid control on the hindlimbs, through necessary adaptations to particular substrates, than on forelimbs which are additionally used for habitat exploration, food manipulation and social activities (Szalay 1994). Thirdly, the fact that the calcaneum is so robust not only

63 CHAPTER 3

provides for a better sample size but also indicates that it has important load-bearing functions. The load-bearing mechanics of the hindfoot are associated with locomotor behaviour (Szalay 1994). The calcaneum is suited to high tensile, bending and compressive forces, thus providing a support for the body weight that is firm yet elastic

(Hall and Shereff 1993).

The calcaneum is involved, along with adjoining foot elements, in rotation, flexion/extension and adduction/abduction (sideways inward and outward) movements of the foot. The contour and orientation of the articulating surfaces of the calcaneum are particularly informative in terms of indicating the extent of these movements (Sarrafian

1993). For example, cursorial and saltatorial locomotions require stability during high- speed movement over ground, which results from suppressed rotation of the foot through restricted lateral movement at the calcaneocuboid articulation and close contact between the astragalus and calcaneum (Bishop 1997). Arboreal locomotion, on the other hand, involves specific adaptations for greater freedom of movement that allow climbing on the uneven and potentially unstable surfaces of vegetation. Thus, measurements of these articulating surfaces are potentially useful in determining locomotor behaviour.

Ideally, the entire foot complex/unit should be used for such a study because articulating bones are closely adapted functionally for particular bioroles (Szalay 1994). However, due to the nature of most fossil deposits, associated or articulated elements rarely occur, making it impossible to determine which elements belong together as part of one individual’s functional unit. This is also the reason that the fossil calcanea used in this study could not be assigned to species already described – most Riversleigh fossil

64 CHAPTER 3

species have been described on the basis of their dentition, which in most cases is not associated with postcranials.

3.1.4 Ancestry as a confounding factor

It has to be acknowledged that functional role is not the only factor dictating the form of skeletal structure – ancestry has a substantial input (Szalay 1982, Bishop 1997). Form is phylogenetically constrained within taxa and functional adaptations must develop within these constraints, in addition to constructional and architectural constraints imposed by physics and the properties of the ‘building materials’ available for construction (Hickman 1988, Szalay 1982). This is apparent from casual observations of the heel bones of closely related taxa, which exhibit obvious similarities. The influence of phylogeny on the morphology of species cannot always be distinguished from that of function, so no attempt will be made here to differentiate between the two a priori. The results of the analysis undertaken will reveal whether or not function (or locomotor behaviour) can be inferred from morphometrics, irrespective of the constraints of phylogeny. If species group together within the same locomotor category, despite being only distantly related, or converesly, if closely related taxa are distinguished on the basis of locomotor behaviour then the influence of function on morphology may be considered sufficient for the purposes of this study.

65 CHAPTER 3

3.2. Methods

3.2.1. Specimens

Specimens of modern marsupials examined are from the collections of the Australian

Museum (‘A’, ‘M’, ‘P’ or ‘S’ prefix), the South Australian Museum (‘SAM’ prefix) and the University of New South Wales (‘UNSWZ’ and ‘AR’ prefixes). A list of the specimens measured and the raw measurement data (each measurement taken three times and averaged) are provided in Appendices B1 and B2. Appendix B4 lists fossil specimens and their raw measurements. Fossil specimens are from the Queensland

Museum collection (‘QMF’ prefix).

Fossil calcanea were assigned to groups based on similarity in size and morphology; each ‘group’ is an unnamed morphotype. As mentioned above, most postcranial elements from Riversleigh sites cannot be assigned to described species because descriptions are based on dentitions not generally associated with postcranials in the fossil deposits. Terminology and abbreviations for calcaneum morphology follow

Szalay (1982, 1994).

3.2.2. Measurements

Listed in Table 3.2.1 are the raw calcaneum measurements and abbreviations (as they appear in the Appendices). Figure 3.2.1 illustrates the raw measurements which were taken with Brown and Sharpe SO 9001 digital calipers (TESA Cal 1P65) to the nearest

0.01 mm. The choice of measurements covers all surfaces of the calcaneum, with a focus on the dimensions of articulating surfaces. The fibula does not articulate with the calcaneum in all species (Szalay 1994), but a general region of the calcaneofibular is apparent. Measurements of this region or of a calcaneofibular articulation itself are referred to here as ‘region of the calcaneofibular’. The fourteen raw measurements were

66 CHAPTER 3

Raw measurement description [abbreviation] Tuber length (from proximal end of calcaneum to proximal end of CLAJP in dorsal view) [ONE] Tuber width (maximum width in dorsal view) [TWO] Dorsal calcaneum head length (from proximal end of CLAJP to distal end of CaCu in dorsal view) [THREE] Dorsal calcaneum head width (maximum medio-lateral width in dorsal view) [FOUR] Calcaneocuboid maximum width (medio-lateral direction in distal view) [FIVE] Region of the calcaneofibular maximum length (in proximal-distal direction) [SIX] Region of the calcaneofibular maximum width (in dorso-plantar direction) [SEVEN] Lateral calcanum head length (from proximal end of CaFi to distal end of CaCu in lateral view) [EIGHT] Lateral calcaneum head width (maximum width of calcaneum head in dorso-plantar direction) [NINE] Entire calcaneum length (in lateral view) [TEN] Medial tuber length (from proximal end of CLAJP to proximal end of calcaneum in medial view) [ELEVEN] Medial calcaneum head length (from proximal end of CLAJP to distal end of calcaneum in medial view) [TWELVE] Calcaneocuboid maximum height (dorso-plantar direction in distal view) [THIRTEEN] Region of the calcaneofibular protrusion (distance from tuber in proximal view) [FOURTEEN]

Table 3.2.1. Descriptions of raw measurements for calcanea. Abbreviations (as in Szalay 1994): CLAJP, continuous lower ankle joint pattern; CaCu, calcaneocuboid; CaFi, calcaneofibular.

FOUR B NINE A TWELVE

SEVEN EIGHT

THREE SIX

FOURTEEN TEN CLAJP ELEVEN

ONE

C CaFi TWO

CaCu THIRTEEN

FIVE

Figure 3.2.1. Illustration of raw measurements for calcanea. A is dorsal view, B is lateral view, C is distal view (see Table 3.2.1 for measurement description)

67 CHAPTER 3

RATIOS OF RAW VARIABLE DESCRIPTION OF VARIABLE MEASUREMENTS NAME TWO/ONE TALONID talonid shape THREE/FOUR HEAD shape of dorsal surface of calcaneum head (most of which articulates with astragalus) THREE/ONE HEADTAL calcaneum head length relative to talonid length THIRTEEN/FIVE CACU calcaneocuboid shape SIX/SEVEN CAFI shape of calcaneofibular region EIGHT/NINE LATHEAD dorso-plantar thickness of calcaneum head relative to lateral length TWELVE/EIGHT MEDLAT medio-lateral orientation of calcaneocuboid FOURTEEN/TEN CAFI/TALONID extent of outward protrusion of region of the calcaneofibular

Table 3.2.2. Variables used in canonical variates analysis (variables are derived from raw measurements)

converted to ratios, which were subsequently used as the variables in the statistical analysis (Table 3.2.2). The reasons for this were, firstly, to clarify shape aspects of the bones; secondly, comparisons between groups are easier when values are not absolute; and thirdly, to reduce the number of variables relative to species, as the analysis requires. Allometric relationships may confound morphometric analyses. Ratio variables were regressed against body mass for extant marsupials to determine whether there is a significant correlation (which indicates allometric relationship). No correlation was found, thus allometry is not expected to confound the results.

3.2.3. Locomotor categories

The present study distinguishes four categories of locomotion for Australian and New

Guinean marsupials: 1) arboreal, includes species that spend the majority of their time on vegetation (feeding and sleeping); 2) scansorial, includes species that spend a considerable proportion of time on the ground but are capable climbers; 3) terrestrial, includes quadruped species restricted mostly to the ground or underground; 4) hopping or bipedal terrestrial includes species using mostly bipedal hopping to locomote. These categories serve to highlight differences between species in their use of available substrates and thus to illustrate their distribution among the structural components of the

68 CHAPTER 3

habitat. Aerial and aquatic locomotor behaviours are not included in the following analysis because this study focuses only on Australo-Papuan marsupial species.

Small mammals, due to their small body size, are often confronted with uneven substrates and (for them) steeply inclined surfaces of tree roots, fallen logs and rocks

(Jenkins 1974). For this reason, even those that spend most of their time on the ground are actually able to climb. Small mammals (under approximately 500 g body mass, for the purposes of this study) therefore have a versatile locomotor pattern and in this study are classified as ‘scansorial’, because they have the functional potential to locomote on the ground or in vegetation.

3.2.4. Notes on the modern marsupial sample

This study looks at the species within each locomotor group as single entities, so individuals within each species group are not included separately, instead, the average of their measurements is used. Measurements were taken of as many specimens as were available for each modern marsupial species. Within-species morphological variation is expected to be smaller than between-species variation (except in some cases of extreme sexual dimorphism within a species of which no examples were included). In this study, sexual dimorphism is unimportant because the male and female members of the species used have the same locomotor behaviour. If different species group together according to their mode of locomotion, then individuals of the same species should as well. This was tested for the modern Isoodon macrourus (Northern Brown Bandicoot) by entering individuals of one sex into the analysis as ‘known’ and individuals of the other sex as

‘unknowns’ (see Appendix B3 for specimen list and raw data) and verifying that unknowns are placed in the appropriate category by the analysis. I. macrourus was

69 CHAPTER 3

chosen for its relatively larger sample size and because the average body mass of males is almost twice that of females (Gordon 1998), thus exhibiting some degree of sexual dimorphism.

3.2.5. Statistical analysis

A canonical variates analysis (multiple discriminant analysis) was carried out to see if the calcaneum measurements distinguish species on the basis of their locomotor behaviour. Canonical variates analysis is a multi-group discriminant analysis that highlights differences between groups (locomotor categories in this case) by spacing out the means of the groups to the maximum extent through construction of linear functions

(Oxnard 1968). The separation between the groups is maximised relative to the variation within each group and the analysis calculates the likelihood of an unknown (in this case a fossil) belonging to one or other of the groups (Reyment et al. 1984).

The analysis was performed using SYSTAT (for Windows) version 7.0 (1997) and graphs were produced with PAST (Palaeontological Statistics) version 1.27 (Hammer et al. 2004).

70 CHAPTER 3

3.3. Results

Assumptions of canonical variates analysis include normality of data and equivalence of variances/covariances across groups. A list of the modern species used in the analysis, the locomotor categories they were assigned to, and the corresponding values for each variable used for the analysis is provided in Table 3.3.1. Fossil morphotypes entered into the analysis and corresponding values for each variable are listed in Table 3.3.2.

There is no direct test for multivariate normality, but if all variables exhibit univariate normality then departures from multivariate normality are usually inconsequential (Hair et al. 1998). Each variable was examined graphically and statistically for the modern sample. A normal probability plot shows the data points plotted over the straight diagonal line of the normal distribution (Hair et al. 1998). The plotted data follow the straight diagonal closely for most variables in this analysis (Appendix B5), indicating that these variables are normally distributed. The statistical test of normality is based on the value of skewness for the data (Hair et al. 1998). All variables, except TALONID and CAFI, are normally distributed at the 0.01 probability level.

A statistical test for equality of covariance matrices is Box’s M test, but it is very sensitive to departures from normality. A violation of the assumption of equivalence of covariance matrices across groups has minimal impact if the groups are of approximately equal size - if the largest group size divided by the smallest group size is less than 1.5 (Hair et al. 1998). The ‘hopping’ category consists of almost twice as many species as the ‘terrestrial’ category in this analysis. The analysis was re-run with the ‘hopping’ category reduced to a group size equal to that for the ‘terrestrial’ category and results were very similar to those presented here for the larger group of hoppers.

71 CHAPTER 3

ORDER SPECIES LOCOMOTION BODYMASS (g) TALONID HEAD HEADTAL CACU CAFI LATHEAD MEDLAT CAFI/TOTAL

DAS Dasyuroides byrnei* terrestrial # 110 0.783 1.335 1.568 0.827 1.212 1.552 0.912 0.133 Dasyurus maculatus* scansorial 5500 0.635 1.336 1.177 0.858 1.176 1.552 0.885 0.126 PER Isoodon macrourus* terrestrial 1600 0.497 1.190 0.779 1.083 1.034 1.330 1.061 0.080 DIP Phascolarctos cinereus* arboreal 5800 0.697 1.102 1.389 1.019 1.066 1.372 0.883 0.165 Petaurus norfolcens* arboreal # 230 0.661 1.550 1.947 0.992 1.339 1.959 0.786 0.127 Pseudocheirus peregrinus* arboreal 900 0.572 1.358 1.326 1.076 1.471 1.848 0.832 0.114 Pseudochirops archeri* arboreal 2183 0.473 1.468 1.250 1.217 1.306 1.821 0.818 0.148 Spilocuscus maculatus* arboreal $ 3813.5 0.573 1.321 1.279 1.102 1.318 1.820 0.915 0.108 Trichosurus vulpecula* arboreal 2850 0.858 1.295 1.653 1.039 1.184 1.870 0.891 0.124 Onychogalea fraenata* hopper 5500 0.547 0.910 0.802 0.903 1.479 1.030 1.402 0.139 Aepyprymnus rufescens* hopper 3250 0.559 0.874 0.812 0.833 1.129 0.988 1.226 0.149 Burramys parvus terrestrial # 41.5 0.714 1.413 1.929 1.278 1.463 1.804 0.851 0.104 Hemibelideus lemuroides arboreal 952.5 0.688 1.566 1.711 1.041 1.206 1.642 0.750 0.051 Petauroides volans arboreal 1300 0.758 1.323 1.533 0.996 1.145 1.358 0.796 0.088 Petaurus breviceps arboreal # 127.5 0.375 1.858 1.527 1.050 1.468 2.245 0.821 0.129 Phalanger gymnotis scansorial $ 2705 0.739 1.295 1.385 0.935 1.313 1.694 0.882 0.118 Phalanger orientalis arboreal $ 2487.5 0.579 1.443 1.651 1.043 1.328 1.797 0.931 0.094 Pseudochirulus cf. arboreal $ # 300 0.622 1.428 1.509 0.895 1.332 1.866 0.726 0.167 canescens Lasiorhinus latifrons terrestrial 25500 0.847 1.242 1.275 0.802 1.361 1.227 0.881 0.100 Vombatus ursinus terrestrial 26000 0.812 1.138 1.170 1.342 1.133 1.209 0.741 0.090 Bettongia gaimardi hopper 1660 0.582 0.929 0.789 0.869 1.070 1.000 1.244 0.126 cuniculus Bettongia penicillata hopper 1300 0.508 0.863 0.748 0.873 0.826 0.906 0.982 0.123 Dendrolagus dorianus arboreal $ 10068 1.165 0.851 1.392 0.720 0.978 1.145 1.013 0.113 Dendrolagus goodfellowi arboreal $ 8025 0.801 0.771 1.032 0.705 1.016 1.345 1.088 0.132 Dendrolagus inustus arboreal $ 10554 0.770 0.663 0.842 0.663 0.891 1.239 1.035 0.156 Dendrolagus lumholtzi arboreal 6750 0.819 0.781 1.129 0.721 0.922 1.199 0.994 0.170 Dendrolagus matschii arboreal $ 9228 1.160 0.762 1.327 0.649 1.158 1.263 0.883 0.180 Dendrolgus scottae arboreal $ 9250 0.875 0.692 0.860 0.719 0.853 1.132 0.878 0.133 Dorcopsis afrata hopper 0.676 0.859 0.812 0.672 0.994 0.956 1.273 0.173 Dorcopsis muelleri hopper $ 5000 0.552 0.702 0.598 0.748 0.975 1.012 1.358 0.146 Hypsiprymnodon scansorial 520 0.716 1.106 1.102 0.799 3.172 1.449 0.738 0.095 moschatus Lagorchestes hirsutus hopper 1265 0.535 0.973 0.889 0.796 1.503 1.088 1.286 0.139 Macropus agilis hopper 15000 0.468 0.991 0.737 0.868 1.046 1.117 1.317 0.120 Macropus eugenii hopper 6500 0.633 1.211 0.909 0.828 1.316 1.115 1.354 0.109 Macropus fuliginosus hopper 40500 0.433 0.973 0.621 0.852 1.466 0.934 1.362 0.130 Macropus giganteus hopper 49000 0.452 0.944 0.597 0.865 1.526 1.063 1.347 0.112 Macropus parma hopper 5350 0.577 0.866 0.854 0.868 1.208 1.111 1.254 0.136 Macropus parryi hopper 13500 0.589 0.858 0.818 0.863 1.469 1.036 1.255 0.112 Macropus rufus hopper 46250 0.492 1.151 0.681 0.898 1.510 0.967 1.287 0.094 Onychogalea unguifera hopper 6650 0.437 0.886 0.598 0.818 1.566 0.926 1.354 0.100 Petrogale brachyotis hopper 4050 0.611 0.897 0.866 0.770 0.788 1.062 0.924 0.130 Petrogale mareeba hopper 8300 0.589 0.990 0.931 0.799 1.397 1.205 0.921 0.103 Petrogale penicillata hopper 7100 0.626 1.078 0.986 0.761 1.402 1.126 0.999 0.101 Petrogale persephone hopper 6200 0.564 0.986 0.889 0.853 1.171 1.012 1.046 0.100 Potorous tridactylus hopper 1100 0.567 0.771 0.796 0.756 0.986 0.934 1.093 0.163 Wallabia bicolor hopper 15000 0.558 0.974 0.878 0.853 1.140 0.885 1.374 0.134 DAS Antechinus agilis scansorial # 28 0.869 1.389 1.844 0.660 1.111 1.767 0.821 0.083 Antechinus stuartii scansorial # 44 0.784 1.664 1.813 0.802 1.333 1.707 0.766 0.138 Dasyurus viverrinus scansorial 1090 0.651 1.358 1.239 0.695 1.280 1.612 0.936 0.117 Parantechinus apicalis terrestrial # 27.5 0.586 1.487 1.586 0.615 1.403 2.050 0.847 0.144 Phascogale tapoatafa arboreal # 193.5 0.595 1.561 1.432 0.953 1.438 1.603 0.913 0.108 Sarcophilus harrisii scansorial 8000 0.548 1.161 0.991 0.783 1.131 1.565 0.855 0.142 Sminthopsis murina terrestrial # 17 0.562 1.667 1.731 0.787 2.269 2.529 0.886 0.121 Thylacinus cynocephalus terrestrial 25000 0.508 1.037 0.803 0.911 1.312 1.182 1.008 0.126 PER Echymipera kalubu terrestrial $ 954 0.563 1.277 0.877 1.130 1.208 1.517 0.914 0.127 Echymipera rufescens terrestrial 0.556 1.283 0.979 1.150 1.197 1.818 1.061 0.076 1000(%) Isoodon obesulus terrestrial 775 0.582 1.169 0.938 1.118 1.421 1.640 0.990 0.082 Macrotis lagotis terrestrial 1350 0.755 1.509 1.155 0.949 1.418 1.709 0.951 0.130 Perameles nasuta terrestrial 975 0.565 1.330 0.929 1.294 1.017 1.391 1.024 0.039 Peroryctes raffrayana terrestrial 834 0.603 1.337 0.978 1.190 1.020 1.725 1.006 0.111 NOTO Notoryctes typhlops terrestrial # 55 0.282 1.633 0.934 0.452 0.369 2.433 1.031 0.109

Table 3.3.1. Data for modern species used in canonical variates analysis. Abbreviations: DAS, Dasyuromorphia; PER, Peramelemorphia; DIP, Diprotodontia; NOTO, Notoryctemorphia. * indicates average of more than one specimen # indicates species placed in ‘scansorial’ category in analysis due to having <500g body mass $ indicates Flannery (1995) as reference; reference for the remainder is Strahan (ed. 1998) % indicates body mass inferred from body length

72 CHAPTER 3

SITE ORDER MORPHOTYPE TALONID HEAD HEADTAL CACU CAFI LATHEAD MEDLAT CAFI/TOTAL CS DIP CSB 0.439 1.065 0.932 0.924 1.198 1.135 1.206 0.100 CSI 0.447 1.229 1.040 0.688 1.357 1.317 1.063 0.097 CSS1 0.467 0.799 0.808 0.771 1.105 1.133 1.135 0.084 CSS2 0.631 0.769 0.857 0.795 1.178 1.102 1.125 0.103 CST 0.496 1.151 0.843 0.869 1.105 1.169 1.133 0.077 CSV 0.360 0.916 0.716 0.766 1.289 1.112 1.042 0.084 CSC 0.551 1.104 0.982 0.801 0.991 1.185 1.064 0.077 CSJ 1.014 1.004 1.681 0.856 1.093 1.202 1.071 0.197 CSU 0.810 0.983 1.298 0.767 1.057 1.330 1.088 0.125 CSQ 0.564 1.425 1.693 1.012 1.364 1.798 0.964 0.097 CSR 0.661 1.573 1.421 0.702 0.981 1.704 0.783 0.060 DAS CSP 0.517 1.454 1.216 1.163 1.607 2.250 0.965 0.093 PER CSa 0.541 1.243 0.973 1.022 1.026 1.347 1.046 0.164 CSd 0.561 1.227 0.948 1.118 1.348 1.278 0.996 0.117 CSe 0.671 1.340 1.190 1.202 1.278 1.568 1.175 0.063 CSf 0.445 1.461 1.245 0.952 1.192 1.828 1.002 0.116 CSg 0.459 1.490 0.992 1.079 1.681 1.560 1.307 0.087 CSh 0.609 1.322 1.322 1.144 1.566 1.728 1.155 0.095 CSm 0.485 1.326 1.381 1.466 1.144 2.211 1.056 0.104 CSn 0.514 1.514 1.500 1.372 1.063 2.571 0.889 0.082 CSo 0.362 1.500 1.087 1.333 0.842 1.789 1.147 0.076 CSma 0.484 1.721 1.540 1.320 1.101 2.294 0.993 0.104 CSx 0.657 1.271 1.648 1.143 1.523 2.488 0.956 0.129 CSy 0.568 1.189 1.217 1.309 1.716 2.140 1.057 0.061 CSz 0.831 1.325 1.657 1.429 1.085 1.800 1.204 0.094 MM DIP MM1 0.559 1.039 0.948 0.749 1.217 1.206 1.138 0.118 MM2 0.631 0.857 0.823 0.852 1.089 0.873 1.091 0.131 MM3 0.548 0.826 0.843 0.836 1.333 1.247 1.060 0.117 PER MM4 0.650 1.227 1.188 1.393 1.600 1.870 0.971 0.065 MM5 0.583 1.492 1.311 2.233 1.833 2.560 1.012 0.112 MM6 0.508 1.131 0.885 1.213 1.031 1.587 0.914 0.064 MM7 0.561 1.254 0.933 1.267 1.116 1.421 1.065 0.107 MM8 0.589 1.322 1.102 1.144 1.566 1.728 1.155 0.095 Ec DIP R.flanneryi 0.659 0.922 0.919 0.914 1.374 0.864 1.431 0.154 EcG 0.505 0.986 0.812 0.658 0.889 1.008 1.193 0.135 EcH 0.530 0.890 0.825 0.846 1.220 0.941 1.164 0.132 EcJ 0.631 0.825 0.884 0.756 1.297 1.136 1.107 0.084 EcK 0.384 0.837 0.738 0.704 0.884 1.094 1.024 0.087 EcL 0.433 1.090 0.873 0.814 1.213 1.254 1.122 0.111 EcM 0.548 0.828 0.794 0.736 1.137 1.128 1.089 0.097 EcN 0.440 1.247 0.854 0.861 1.039 1.153 1.190 0.099 EcO 0.641 1.084 0.895 0.851 1.349 1.144 1.084 0.113 Trichosurus 0.649 1.404 1.560 0.927 1.269 1.429 0.806 0.058 DAS EcE 0.530 1.189 1.111 0.950 1.649 1.861 1.164 0.084 EcF 0.624 1.559 1.262 1.091 1.182 1.923 0.960 0.110 PER EcA 0.404 1.465 0.942 1.120 1.354 1.968 1.065 0.008 EcB 0.481 1.260 1.058 1.465 1.367 1.539 1.044 0.061 EcC 0.572 1.219 1.322 1.135 1.474 2.214 1.155 0.073 EcD 0.647 1.429 1.176 1.400 1.250 2.375 0.821 0.090 QL DIP Nambaroo 0.624 0.866 0.799 0.828 1.050 1.083 0.950 0.200 sp.3 PER QLA 0.498 1.170 1.058 1.508 1.051 2.257 0.948 0.095 QLB 0.551 1.185 1.282 1.301 1.366 1.971 0.958 0.107 QLC 0.457 1.170 0.910 1.567 0.950 1.483 0.897 0.079 QLD 0.808 1.367 1.314 1.246 1.507 2.300 0.891 0.148 QLE 0.825 1.145 1.762 1.416 1.224 2.233 1.027 0.126 QLF 0.508 1.183 0.930 1.551 1.102 1.581 0.941 0.124 QLG 0.444 1.818 1.111 1.152 1.250 2.273 0.950 0.080

Table 3.3.2. Data for fossil morphotypes used in canonical variates analysis. Abbreviations: DAS, Dasyuromorphia; PER, Peramelemorphia; DIP, Diprotodontia; CS, Camel Sputum; MM, Mike’s Menagerie; Ec, Encore; QL, Quantum Leap.

73 CHAPTER 3

Canonical variates analysis is a robust technique, in any case, that can tolerate deviations from these assumptions to some extent (Turbón et al. 1997).

A further assumption is minimal multicollinearity of the variables. One way to assess the impact of multicollinearity is to look at tolerance values for the variables, where tolerance is 1 minus the proportion of the variable’s variance explained by the other predictor variables (Hair et al. 1998). Tolerance values approaching one indicate little collinearity, but variables with tolerance values approaching zero are accounted for by other variables in the analysis to a great extent (ibid.). Tolerance levels for variables in this analysis are all over 0.2, so at least 20% of the variation explained by each variable is unique to that variable.

Table 3.3.3 shows summary statistics for the locomotor categories. For the TALONID variable, means differ most between ‘arboreal’ and ‘hopper’ categories. The HEAD variable means differ between ‘scansorial’ and ‘hopper’ to the greatest extent. Means for the HEADTAL variable differ most between species with climbing ability

(‘arboreal’ and ’scansorial’) and ground-dwelling species (‘terrestrial’ and ‘hopper’).

The CACU and CAFI/TOTAL variable means distinguish ‘terrestrial’ species. CAFI and LATHEAD variable means distinguish ‘scansorial’ species, and the MEDLAT and

LATHEAD variables distinguish ‘hopper’ species.

74 CHAPTER 3

Group Statistic TALONID HEAD HEADTAL CACU CAFI LATHEAD MEDLAT CAFI/TOTAL X 0.77 1.1 1.312 0.908 1.131 1.489 0.908 0.127 Arboreal 0.691 0.903 0.869 0.568 0.618 0.738 0.338 0.129 (N = 14) OR SD 0.205 0.33 0.277 0.198 0.186 0.292 0.097 0.036 X 0.656 1.441 1.519 0.862 1.496 1.797 0.842 0.123 Scansorial 0.494 0.753 0.956 0.663 2.06 1.08 0.21 0.083 (N = 15) OR SD 0.12 0.198 0.301 0.167 0.537 0.295 0.065 0.021 X 0.597 1.286 0.983 1.038 1.135 1.562 0.97 0.097 Terrestrial 0.566 0.596 0.495 0.89 1.053 1.251 0.32 0.091 (N = 11) OR SD 0.16 0.168 0.156 0.252 0.296 0.367 0.095 0.027 X 0.55 0.937 0.791 0.826 1.237 1.022 1.222 0.126 Hopping 0.242 0.509 0.389 0.23 0.778 0.32 0.481 0.079 (N = 21) OR SD 0.066 0.116 0.116 0.058 0.246 0.085 0.158 0.022

Table 3.3.3. Summary statistics for modern species. X, group means; OR, observed ranges; SD standard deviations; N, number of species within each group

Figs 3.3.1 A, B and C show the pattern of separation on the canonical variates plot for modern marsupial species. Canonical variate 1 distinguishes the hopping category from all others; canonical variate 2 distinguishes terrestrial from scansorial; variates 2 and 3 distinguish terrestrial from arboreal; and variate 2 provides the best separation between arboreal and scansorial, though there is considerable overlap on visual examination.

Eigenvalues (Table 3.3.4) indicate the variation explained by the canonical axes of the plot. The first canonical variate accounts for 65.8% of the variation in calcaneum morphometrics; 26.9% of the variation is explained by the second canonical variate; and

7.3% by the third.

75 CHAPTER 3

A. 5

60 4

61 3 3 2057 56 TERRESTRIAL 58 2 59 62 28 1 40 35 ARBOREAL 3734 2122 55 78 30 42 2 Score2 0 45 25 5319 23 6 4136 43 16 3938 44 24 9 10 1129 1450 13 47 52 15 -1 46 HOPPERS 27 26 32 4 171 18 -2 SCANSORIAL 311249 33 48

-3 Long talonid 515 54 Thick calcaenum head

-3 -2 -1 0 1 2 3 4

Score1

B. 3

27 23 25 28 24 2 26 ARBOREAL

9 12 20 1 33 4 46 3042 22 48 38 HOPPERS 1439 8 58 114316 57 0 5 17 29 45 21 1 6 TERRESTRIAL Score3 53 61 60 1013 3455 3 18 47 32 44 19 56 41 2 7 -1 51 37 50 36 Calcaenum head 54 4931 35 52 62 59 shorter than talonid -2 40 SCANSORIAL Increased protrusion of CaFi 15 from calcaneum body

-2 -1 0 1 2 3 4 5

Score2

Figure 3.3.1. (Continued next page…) Canonical variates plot for modern marsupial species. A, score 1 against score 2; B, score 2 against score 3. Numbers indicate species as listed in D, with category colours in graphs corresponding to those in D. Morphological features that score positively or negatively high along the axes (and thus distinguish locomotor categories) are indicated along respective axes.

76 CHAPTER 3

3 C. 23 27 25 28 24 2 26 ARBOREAL 9 12 20 1 33 4 3046 42 22 38 48SCANSORIAL 39 14 588 11 43 1657 0 2921 45 17 5 1 6

Score3 53 61 10 34 60 13 47 55 3 18 32 44 19 56 41 2 7 -1 37 51 TERRESTRIAL 50 36 HOPPERS 35 31 49 54 52 62 59 -2 40

-3 -2 -1 0 1 2 3 4

Score1

D. Number Species Number Species 1 Dasyurus byrnei 32 Lagorchestes hirsutus 2 Dasyurus maculatus 33 Macropus 3 Isoodon macrourus 34 Macropus agilis 4 Phascolarctos cinereus 35 Macropus eugenii 5 Petaurus norfolcens 36 Macropus fuliginosu 6 Pseudocheirus peregrinus 37 Macropus giganteus 7 Pseudochirops archeri 38 Macropus parma 8 Spilocuscus maculatus 39 Macropus parryi 9 Trichosurus vulpecula 40 Macropus rufus 10 Onychogalea fraenata 41 Onychogalea unguifera 11 Aepyprymnus rufescens 42 Petrogale brachyotis 12 Burramys parvus 43 Petrogale mareeba 13 Hemibelideus lemuroides 44 Petrogale penicillata 14 Petauroides volans 45 Petrogale persephone 15 Petaurus breviceps 46 Potorous tridactylus 16 Phalanger gymnotis 47 Wallabia bicolor 17 Phalanger orientalis 48 Antechinus agilis 18 Pseudochirulus canescens 49 Antechinus stuartii 19 Lasiorhinus latifrons 50 Dasyurus viverrinus 20 Vombatus ursinus 51 Parantechinus apicalis 21 Bettongia gaimardi cuniculus 52 Phascogale tapoatafa 22 Betongia penicillata 53 Sarcophilus harrisii 23 Dendrolagus dorianus 54 Sminthopsis murina 24 Dendrolagus goodfellow 55 Thylacinus cynocephalus 25 Dendrolagus inustus 56 Echymipera kalubu 26 Dendrolagus lumholtzi 57 Echymipera rufescens 27 Dendrolagus matschii 58 Isoodon obesulus 28 Dendrolagus scottae 59 Macrotis lagotis 29 Dorcopsis afrata 60 Perameles nasuta 30 Dorcopsis muelleri 61 Peroryctes raffrayana 31 Hypsiprymnodon moschatus 62 Notoryctes typhlops Figure 3.3.1. Continued… C, score 1 against score 3 and D, species corresponding to numbers in graphs.

77 CHAPTER 3

A global test of differences between multivariate means showed a significant difference among locomotor categories (Pillai’s trace = 1.824, d.f. = 24, 156, P<0.001). Table 3.3.5 shows the classification results.

CANONICAL VARIATE VARIABLE 1 2 3 CONSTANT -5.523 -3.799 1.491 TALONID 6.710 (0.927) 8.149 (1.126) 0.418 (0.058) HEAD 0.551 (0.116) 2.958 (0.622) -6.830 (-1.435) HEADTAL -1.639 (-0.362) -7.331 (-1.617) 2.597 (0.573) CACU 1.499 (0.251) 4.350 (0.729) 2.328 (0.390) CAFI -0.085 (-0.029) -1.541 (-0.524) -1.121 (-0.381) LATHEAD 4.111 (1.063) 2.449 (0.633) 1.635 (0.423) MEDLAT -4.738 (-0.551) -0.211 (-0.025) 0.562 (0.065) CAFI/TOTAL 1.635 (0.043) -14.707 (-0.387) -2.463 (-0.065) Eigenvalue 4.690 1.924 0.519 Variance (%) 65.8 26.9 7.3

Table 3.3.4. Canonical discriminant functions (values in brackets are standardised coefficients for canonical variates)

Allocated locomotion category Arboreal Scansorial Terrestrial Hopping Arboreal (14) 86/71 14/22 0/7 0/0 Scansorial (15) 20/27 80/73 0/0 0/0 Terrestrial (11) 0/0 9/18 82/73 9/9 Actual Actual

locomotion locomotion Hopping (21) 0/10 0/0 0/0 100/90

Table 3.3.5. Classification results from canonical variates analysis (values indicate the percentage of allocations of individuals to each group; values in italics indicate jacknife classification results; values in parentheses are the numbers of species for each group)

This study aimed to develop a model with which to allocate subjects (fossil calcanea) to groups (locomotor categories). Table 3.3.6 gives the fossil morphotypes and their predicted group allocation with associated probabilities, based on the distance for the particular specimen from the locomotor group centroid on the canonical variates plot.

Some of the morphotypes were estimated as roughly under 500 g body mass and in these cases the locomotor classification is uninformative in terms of habitat structure

78 CHAPTER 3

Morphotype Taxon Arboreal Scansorial Terrestrial Hopper

CSa Peramelemorphian 21 4 44 31 CSB Macropodoid 0 0 0 100 CSC Macropodoid 1 0 0 99 CSd Peramelemorphian 5 0 91 4 Cse # Peramelemorphian 1 0 99 0 CSf # Peramelemorphian 31 59 8 1 CSg # Peramelemorphian 0 0 43 57 CSh # Peramelemorphian 78 6 15 1 CSI Macropodoid 0 0 0 100 CSJ Vombatiform 80 20 0 0 CSM Peramelemorphian 76 0 24 0 CSN # Peramelemorphian 21 0 79 0 CSO # Peramelemorphian 0 0 100 0 CSP Dasyuromorphian 5 0 95 0 CSQ Possum 24 76 0 0 CSR # Possum 9 64 26 0 CSS1 Macropodoid 0 0 0 100 CSS2 Macropodoid 3 0 0 97 CST Macropodoid 0 0 1 99 CSU Vombatiform 94 4 0 2 CSV Macropodoid 0 0 0 100 CSMA Peramelemorphian 13 1 86 0 CSX # Peramelemorphian 98 2 0 0 CSY # Peramelemorphian 35 0 65 0 CSZ # Peramelemorphian 87 0 13 0 R. flanneryi Macropodoid 0 0 0 100 Trichosurus Possum 23 76 0 1 EA Peramelemorphian 0 0 100 0 EB Peramelemorphian 1 0 99 0 EC # Peramelemorphian 90 0 10 0 ED # Peramelemorphian 0 0 100 0 EE # Dasyuromorphian 78 5 16 1 EF # Dasyuromorphian 1 0 99 0 EG Macropodoid 0 0 0 100 EH Macropodoid 0 0 0 100 EJ Macropodoid 3 0 0 97 EK Macropodoid 0 0 0 100 EL Macropodoid 0 0 0 100 EM Macropodoid 0 0 0 100 EN Macropodoid 0 0 0 100 EO Macropodoid 3 1 7 89 MM1 Macropodoid 0 0 0 100 MM2 Macropodoid 0 0 0 100 MM3 Macropodoid 12 0 0 88 MM4 # Peramelemorphian 3 0 97 0 MM5 # Peramelemorphian 0 0 100 0 MM6 Peramelemorphian 0 0 100 0 MM7 Peramelemorphian 0 0 100 0 MM8 # Peramelemorphian 4 0 96 0 Nambaroo sp. 3 Macropodoid 12 1 0 86 QLA # Peramelemorphian 1 0 99 0 QLB Peramelemorphian 93 0 6 0 QLC Peramelemorphian 0 0 100 0 QLD # Peramelemorphian 3 0 97 0 QLE # Peramelemorphian 100 0 0 0 QLF Peramelemorphian 0 0 100 0 QLG # Peramelemorphian 0 0 100 0 Table 3.3.6. Percent probabilities for locomotor group allocations for fossil morphotypes (highest value highlighted). # indicates calcaneum length under 1 mm, therefore body mass presumed to be under 500 g and consequently, the morphotype is classified as ‘small mixed’, regardless of the predicted locomotion.

79 CHAPTER 3

due to the small size of the animals (as explained above). These small fossil taxa were therefore referred to a new category, ‘small mixed’, so as not to skew proportions of taxa among locomotor categories that are representative of actual habitat use. Body mass was estimated as approximately under 500g based on calcaneum length measurements. Calcaneum length versus body mass for the modern sample (Fig. 3.3.2) shows that those with a length less than about 10 mm weigh less than 500g. Notoryctes

(Marsupial Mole) does not follow this trend possibly because it is highly specialised for digging, being a rapid scratch-digger (i.e. digs with both fore and hind feet simultaneously).

2 M.rufusM.giganteus M.fuliginosu

M.parryiM.agilis W.bicolorT.cynocephal Dor.muelleriO.unguiferaD.inustus Pet.persephoPet.penicill M.parmaM.eugeniiD.goodfellowL.latifronsV.ursinus O.fraenataD.lumholtziPet.mareebaD.scottaeD.dorianus A.rufescensPet.brachyotS.harrisiiD.matschii P.nasutaM.lagotisI.macrourusBet.gaimardiS.maculatusD.maculatusP.cinereus Lag.hirsutus N.typhlops P.raffrayanaE.rufescensBet.penicillP.gymnotisT.vulpecula E.kalubuP.tridactyluP.orientalis I.obesulusD.viverrinusP.volansP.archeri P.peregrinus 1 P.canescensH.moschatusH.lemuroides

P.tapoatafa D.byrneiP.norfolcens P.breviceps

log Calcaneum Length P.apicalis A.stuartii S.murinaAnt.agilisB.parvus

3 4 5 6 7 8 9 10 11 12 13

log Body Mass

Figure 3.3.2. Log 10 body mass against log 10 calcaneum length for the modern marsupial sample. Red points indicate body mass under 500 g.

80 CHAPTER 3

3.4. Discussion

The results of this study indicate that calcaneum morphometrics are informative in terms of locomotor function for marsupials. Arboreal, scansorial, terrestrial and hopping

(or saltatorial) locomotor behaviours were distinguished on the basis of calcaneum articular surface shapes and orientations and relative sizes of structural components of the calcaneum. The clearest distinctions are between the hopping category and the remaining three categories, followed by distinction of the terrestrial category from the two climbing categories, arboreal and scansorial. The arboreal and scansorial categories are not completely distinct from one another. Van Valkenburgh (1987) also found less confidence in the separation of these modes of locomotion compared to terrestrial and semi-fossorial modes for carnivores. Unfortunately fossorial behaviour was not investigated in the current study, despite the fact that species with fossorial and semi- fossorial adaptations were included. The reason for this is that digging adaptations are more pronounced in forelimbs as opposed to hindlimbs, but this study only takes the calcaneum into account.

The clear separation of the hopping category from other locomotor categories may partly be due to phylogenetic influences. All species in this category are macropodoids.

Other categories, however, do contain a mix of species from different orders, so function does have a measurable effect on morphology. For example, the terrestrial category includes members of the orders Diprotodontia, Peramelemorphia and

Dasyuromorphia; the scansorial category includes members of Dasyuromorphia and

Diprotodontia; and the arboreal category includes possums and kangaroos. Faith (1989) and Faith and Belbin (1994) propose a method for removing phylogenetic effects in functional studies of morphology. This method could not be applied to the numerical data used here because it requires character states with polarity.

81 CHAPTER 3

As indicated by group means, hoppers have medio-laterally thinner and longer talonids

(lower TALONID values), wider calcaneum heads (lower HEAD values), and dorso- plantarly thicker calcaneum heads (lower LATHEAD value) relative to other locomotor categories. Of these variables, LATHEAD and TALONID are shown to be the most important in distinguishing hoppers from the other locomotor types on the canonical variates plot (Table 3.3.4).

For species capable of climbing (arboreal and scansorial), group means indicate calcaneum heads are longer than talonids, and the calcaneocuboid is more steeply angled antero-posteriorly compared to ground dwelling species (terrestrial and hopping)

- seen in HEADTAL and MEDLAT values. The calcaneucuboid of terrestrial species is bigger dorso-plantarly and in terrestrial species the calcaneofibular protrudes further from the calcaneum body relative to arboreal and scansorial species (higher CACU value and lower CAFI/TOTAL value respectively). The HEADLAT, CACU and

CAFI/TOTAL variables are most important for separation of groups on the second canonical variate. The HEAD variable, in combination with HEADLAT, provides the best separation of arboreal species from scansorial and terrestrial species on the second and third axes of the canonical variates plot.

These results are consistent with previous studies. A long talonid has been associated with cursorial habits (see Youlatos 2003 and references within) and a square-shaped calcaneo-astragalar facet with subtalar stability necessary for terrestrial locomotion

(Szalay and Decker 1974). Talonid shape is largely determined by loading (Szalay

1994). The results of Fa-Hong et al. (1993) indicate that the morphological structure of the talonid separates macaque species into functional groups. The current analysis

82 CHAPTER 3

shows hopping species to have longer, thinner talonids and hopping and terrestrial taxa to have wider, shorter calcaneum heads compared to arboreal and scansorial taxa. This is not unexpected given that fast hopping is not unlike cursorial locomotion in the requirement for stability. Stability is provided by a wide calcaneum head region and a secure calcaneo-astragalar articulation (Szalay 1994, Bishop 1997). Climbing, on the other hand, requires inversion and eversion of the foot that is a result of a long, narrow calcaneo-astragalar facet (Youlatos 2003). The greater dorso-plantar thickness of the calcaneum head in hoppers is possibly related to the force exerted on the region of the calcaneoastragalar articulation during hopping.

The size and orientation of the calcaneocuboid articulation are important in determining pedal and tarsal movements in mammals. Flexion-extension, rotation and adduction/abduction of the forefoot are a result of force transmission from the calcaneum to the cuboid at this articulation (Szalay 1994). Stability in the feet of terrestrial species is indicated by the dorso-plantarly thicker calcaneocuboid articulation

(relative to climbing species) that restricts such movements to some extent. Orientation of the calcaneocuboid also restricts such movements to a greater extent when it is perpendicular to the antero-posterior axis, as seen in hoppers. The more steeply angled this articulation is relative to the antero-posterior axis, the more adduction/abduction, and perhaps rotation, are possible, as seen in species with climbing ability.

The calcaneofibular region is informative in terms of extent of mobility between the crus (tibia and fibula) and the pes (Bishop 1997). This region is more pronounced in terrestrial species relative to climbing species; reduction of the articulating region

83 CHAPTER 3

between the fibula and calcaneum, as seen in Dendrolagus (tree kangaroos), suggests greater mobility at the articulation (Bishop 1997)

Modern species entered into the analysis as unknowns were grouped correctly. They included specimens from female Isoodon macrourus (Northern Brown Bandicoot), to test for possible effects of sexual dimorphism; and juveniles of Pseudocheirus peregrinus (Common Ringtail Possum), Spilocuscus maculatus (Common Spotted

Cuscus), Trichosurus vulpecula (Brushtail Possum) and Onychogalea fraenata (Bridled

Nailtail Wallaby). Juveniles were possibly grouped correctly because the articular patterns in the bones of marsupials are established prior to adult size being reached – the bone is ossified from an epiphysis at the posterior end of the tuber (Szalay 1994).

Some modern species used as ‘knowns’ in the analysis were reallocated to different locomotor categories to those specified for them. For example, Lasiorhinus latifrons

(Southern Hairy-nosed Wombat), a terrestrial species, is predicted to be scansorial in the analysis. This may potentially have to do with wombats being slower movers than, for example, dasyurids and bandicoots. Slow terrestrial locomotion requires less stability, and thus a less restrictive form-function relationship, than that necessary for terrestrial cursors and jumpers (Szalay 1994). Of the arboreal species used in the analysis,

Hemibelideus lemuroides (Lemuroid Ringtail Possum) and Phalanger orientalis

(Northern Common Cuscus) were missclassified as scansorial because the specimens have very slightly smaller Mahalanobis distances from the scansorial group centroid than from the arboreal group centroid. Another point to consider is that some animals may be intermediate between locomotor categories, but for the purposes of the analysis have had to be allocated to one group only. The feeding, postural and manipulative

84 CHAPTER 3

behaviour of an animal may influence the extent to which overlap into different categories of locomotor behaviour occurs (Argot 2001).

3.5. Conclusions

The results of this study indicate that heel bone measurements are informative in terms of function. This method of inferring locomotion only requires a single element thus could be applied to any fossil site that contains marsupial calcanea. The allocation of fossil taxa from Riversleigh sites to categories of locomotor behaviour will form a component of the data set to be used in the paleoecological study of these sites.

Inferences about the habitats of past communities will be made on this basis and implications for vegetation will be discussed in chapter 5.

85 CHAPTER 3

3.6. References

Argot, C. 2001. Functional-adaptive anatomy of the forelimb in the Didelphidae, and the paleobiology of the Paleocene marsupials Mayulestes ferox and Pucadelphys andinus. Journal of Morphology 247: 51-79.

Bishop, N. 1997. Functional anatomy of the macropodid pes. Proceedings of the Linnean Society of N.S.W. 117: 17-50.

Damuth, J. D. 1992. Taxon-free characterization of animal communities. Pp. 183-203 in Terrestrial Ecosystems Through Time. Eds A. K. Behrensmeyer, J. D. Damuth, W. A. Di Michele, R. Polts, H. D. Sues and S. L. Wing. The University of Chicago Press, Chicago.

Eisenberg, J. F. 1990. The behavioural/ecological significance of body size in the Mammalia. Pp. 25-37 in Body Size in Mammalian Paleobiology: Estimation of Biological Significance. Eds J. Damuth and B. J. MacFadden. Cambridge University Press, Cambridge.

Fa-Hong, Y., Yan-Zhang, P., Ru-Liang, P, Zhi-Zhang, Y. and Hong, W. 1993. Comparative studies on ankle joints in quadriceps. Acta Zoologica Sinica 39: 424-430.

Faith, D. P. 1989. Homoplasy as pattern: multivariate analysis of morphological convergence in Anseriformes. Cladistics 5: 235-258.

Faith, D. P. and Belbin, L. 1994. Distinguishing phylogenetic effects in multivariate models relating Eucalyptus convergent morphology to environment. Pp. 169-188 in Phylogenetics and Ecology. Eds P. Eggleton and R. I. Vane-Wright. Academic Press, London.

Flannery, T. F. 1982. Hindlimb structure and evolution in the kangaroos (Marsupialia: Macropodoidea). Pp. 507-524 in The Fossil Vertebrate Record of Australasia. Eds P. V. Rich and E. M. Thompson. Monash University Offset Printing Unit, Victoria.

Flannery, T. F. 1995. Mammals of New Guinea. Reed Books, Sydney.

Gordon, G. 1998. Northern brown bandicoot, Isoodon macrourus (Gould, 1842). Pp. 174-175 in The Mammals of Australia. Ed. R. Strahan. New Holland Publishers Pty Ltd, Sydney.

Hair, J. F., Anderson, R. E., Tatham, R. L., Black, W. C. 1998. Multivariate Data Analysis. Prentice-Hall Inc., U.S.A.

Hall, R. L. and Shereff, M. J. 1993. Anatomy of the calcaneus. Clinical Orthopaedics and Related Research 290: 27-35.

Hammer, Ø., Harper, D. A. T. and Ryan, P. D. 2004. PAST – PAlaeontological STatistics, Version 1.27. Computer program available online at http://folk.uio.no/ohammer/past

86 CHAPTER 3

Hickman, C. S. 1988. Analysis of form and function in fossils. American Zoologist 28: 775-793.

Janis, C. M., Theodor, J. M. and Boisvert, B. 2002. Locomotor evolution in camels revisited: a quantitative analysis of pedal anatomy and the acquisition of the pacing gait. Journal of Vertebrate Paleontology 22 (1): 110-121.

Jenkins, F. A. 1974. Tree shrew locomotion and the origins of primate arborealism. Pp. 85-115 in Primate Locomotion. Ed. F. A. Jenkins, Jr. Academic Press Inc., New York.

Jenkins, F. A. and MacLearn, D. 1984. Mechanisms of hind foot reversal in climbing mammals. Journal of Morphology 182: 197-219.

Lemelin, P. 1999. Morphological correlates of substrate use in didelphid marsupials: implications for primate origins. Journal of Zoology London 247: 165-175.

Oxnard, C. E. 1968. The architecture of the shoulder in some mammals. Journal of Morphology 126: 249-290.

Reyment, R. A., Blackith, R. E., Campbell, N. A. 1984. Multivariate Morphometrics. Academic Press, London.

Sarrafian, S. K. 1993. Biomechanics of the subtalar joint complex. Clinical Orthopaedics and Related Research 290: 17-26.

Strahan, R. (ed.) 1998. The Mammals of Australia. New Holland Publishers Pty Ltd, Sydney.

SYSTAT – Version 7.0 for Windows. 1997. SPSS Inc., Chicago (Computer program).

Szalay, F. S. 1982. A new appraisal of marsupial phylogeny and classification. Pp. 621- 640 in Carnivorous Marsupials. Ed. M. Archer. Royal Zoological Society of New South Wales, Australia.

Szalay, F. S. 1994. Evolutionary History of the Marsupials and an analysis of Osteological Characters. Cambridge University Press, Cambridge.

Szalay, F. S. and Decker, R. L. 1974. Origins, evolution and function of the tarsus in late Cretaceous eutherians and Paleocene primates. Pp. 233-259 in Primate Locomotion. Ed. F. A. Jenkins, Jr. Academic Press, New York.

Szalay, F. S. and Sargis, E. J. 2001. Model-based analysis of postcranial osteology of marsupials from the Palaeocene of Itaborai (Brazil) and the phylogenetics and biogeography of Metatheria. Geodiversitas 23: 139-302.

Turbón, D., Pérez- Pérez, A. and Springer, C. B. 1997. A multivariate analysis of Pleistocene hominids: testing hypotheses of European origins. Journal of Human Evolution 32: 449-468.

87 CHAPTER 3

Van Valkenburgh, B. 1987. Skeletal indicators of locomotor behavior in living and extinct acrnivores. Journal of Vertebrate Paleontology 7: 162-182.

Walker, A. 1974. Locomotor adaptations in past and present prosimian primates. Pp. 349-381 in Primate Locomotion. Ed. F. A. Jenkins. Academic Press, New York.

Youlatos, D. 2003. Calcaneal features of the Greek Miocene primate Mesopithecus pentelicus (Cercopithecoidea: Colobinae). Geobios 36 (2): 229-239.

88 CHAPTER 3

APPENDIX B1. Calcaneum measurements for modern marsupial species represented by more than one specimen

SPECIES SPECIMEN NSEX ONE TWO THREE FOUR FIVE SIX SEVEN EIGHT NINE TEN ELEVEN TWELVE THIRTEEN FOURTEEN Dasyuroides byrnei M 26213 F 2.78 2.13 4.13 3.1 2.45 1.18 0.97 4.08 2.85 6.76 3.23 3.83 2.33 0.86 Dasyuroides byrnei M 24651 ? 2.67 2.11 4.28 3.1 2.47 1.27 0.78 3.78 2.2 6.39 3.06 3.59 1.9 0.83 Dasyuroides byrnei M 24037 M 2.83 2.24 4.57 3.52 2.82 1.38 1.41 4.46 2.89 7.58 4.36 3.81 2.17 1.06 Average for D. byrnei 2.76 2.16 4.3267 3.24 2.58 1.277 1.05333 4.1067 2.6467 6.91 3.55 3.743333 2.1333333 0.91666667 Dasyurus maculatus M 11137 M 9.1 5.78 10.82 7.83 6.73 3.45 2.72 10.03 6.7 18.47 11.1 9.05 5.09 2.67 Dasyurus maculatus M 725 ? 10.63 6.64 11.67 9.55 7.54 3.49 2.89 11.46 7.32 21.33 11.39 9.85 6.21 2.29 Dasyurus maculatus A 6379 ? 10.39 6.21 11 8.29 7.19 3.17 2.25 10.78 7.67 20.14 11.7 9.55 6.03 2.85 Dasyurus maculatus B 5449 ? 9.75 6.29 12.04 9.22 7.26 3.93 3.4 12.23 7.5 21.98 11.31 10.53 6.43 2.69 Dasyurus maculatus M 35912 M 10.06 5.77 11.38 8.89 6.08 3.42 3.42 12.05 7.69 20.49 11.36 10.25 6.7 2.8 Dasyurus maculatus M 35612 M 9.29 6.37 11.92 8.75 7.57 3.71 3.6 11.91 7.14 20.55 11.1 10.34 6.62 2.82 Dasyurus maculatus M 35469 M 9.06 6.3 11.53 8.05 6.54 3.8 3.2 11.17 6.98 19.91 11.02 10.2 5.6 1.96 Dasyurus maculatus M 35470 M 9.63 5.88 11.65 7.54 6.45 3.6 2.57 11.31 7.24 19.64 10.56 10.31 5.84 2.42 Dasyurus maculatus M 37205 ? 9.32 6.3 11.71 8.91 7.5 3.69 3.45 11.59 7.46 20.23 11.8 10.28 6.02 2.32 Dasyurus maculatus S 1146 ? 10.13 6.82 11.01 8.81 7.42 3.68 3.04 11.3 7.79 20.66 11.34 10.06 6.5 2.37 Dasyurus maculatus S 1560 ? 9.83 5.68 11.42 8.61 7.2 4.09 3.51 11 6.94 19.73 12.07 10.02 5.44 2.83 Average for D. maculatus 9.7445 6.1855 11.468 8.5864 7.044 3.639 3.09545 11.348 7.3118 20.28 11.34091 10.04 6.0436364 2.54727273 Isoodon macrourus M 35890 M 10.26 4.67 8.98 7.52 4.02 3.1 2.65 8.1 6.7 18.82 10.56 8.32 4.23 1.6 Isoodon macrourus M 35410 M 10.86 5.42 7.99 7.6 3.6 2.49 3.09 9.18 6.4 19.15 11.47 9.78 5.01 1.6 Isoodon macrourus M 33980 M 11.54 6.36 9.69 7.43 4.86 3.09 2.54 9.25 6.69 20.65 12.08 9.86 4.76 1.9 Isoodon macrourus M 33981 M 14.18 6.84 9.85 8.14 5.28 3.78 3.77 9.47 7.28 23.5 13.92 10.24 5.23 1.46 Average for I. Macrourus 11.71 5.8225 9.1275 7.6725 4.44 3.115 3.0125 9 6.7675 20.53 12.0075 9.55 4.8075 1.64 Phascolarctos cinereus P 481 ? 9.16 6.81 13.21 12.83 7.18 6.02 5.99 12.29 8.91 19.84 8.88 9.84 7.86 3.27 Phascolarctos cinereus M 23629 F 9.67 6.13 13.33 10.51 8.04 5.87 6.08 12.86 9.92 20.99 9.37 12.12 8.12 2.74 Phascolarctos cinereus M 32063 ? 9.65 6.47 13.3 11.39 8.3 5.96 5.59 13.39 9.26 21.07 9.63 12.67 8.25 3.13 Phascolarctos cinereus M 35008 M 9.82 7.27 13.34 13.51 9.25 7.36 6 13.33 9.71 21.78 11.43 11.18 9.17 4.69 Average for P. cinereus 9.575 6.67 13.295 12.06 8.193 6.303 5.915 12.968 9.45 20.92 9.8275 11.4525 8.35 3.4575 Petaurus norfolcensis M 37571 ? 2.3 1.65 5.29 3.35 2.49 2.05 0.89 5 2.74 7 3.45 3.82 2.54 1.02 Petaurus norfolcensis M 36038 ? 3.1 1.26 4.51 2.79 1.84 1.32 0.98 4.71 2.32 6.29 3.03 3.32 2.02 0.91 Petaurus norfolcensis M 32998 F 3.31 2.14 5.18 3.22 2.23 1.32 1.14 4.93 2.26 7.57 3.45 4.01 2.42 1.09 Petaurus norfolcensis M 32748 F 2.53 1.52 5.04 3.17 2.66 1.9 1.3 5.46 2.38 7.52 3.04 4.04 2.01 0.71 Petaurus norfolcensis M 31834 F 2.14 1.98 5.22 3.38 2.36 1.77 1.39 5.18 2.61 7.21 3.5 4.23 2.13 0.84 Petaurus norfolcensis M 36041 F 2.7 1.69 5.92 3.55 2.39 1.63 1.38 5.71 2.55 8.25 4.39 4.62 2.59 0.76 Petaurus norfolcensis M 35158 ? 2.09 1.77 4.22 3.36 2.16 1.5 1.5 4.09 3.05 5.95 3.29 3.52 2.29 0.97 Average for P. norfolcensis 2.5957 1.7157 5.0543 3.26 2.304 1.641 1.22571 5.0114 2.5586 7.113 3.45 3.937143 2.2857143 0.9 Pseudocheirus peregrinus M 31210 ? 5.16 3.05 7.64 6.1 3.69 2.53 1.32 7.39 4.48 11.66 5.84 6.15 4.29 1.15 Pseudocheirus peregrinus M 33753 F 5.54 3.28 6.85 5.36 3.86 2.81 2.02 7.52 3.89 11.52 5.99 6.04 3.84 1.22 Pseudocheirus peregrinus M 35472 M 5.33 3.29 7.03 4.85 3.6 2.7 1.86 7.55 3.62 12.3 5.57 6.34 3.88 1.29 Pseudocheirus peregrinus M 34735 ? 5.38 2.62 6.88 4.61 3.37 2.14 1.72 6.38 3.62 10.77 5.91 5.46 3.61 1.62 Average for P. peregrinus 5.3525 3.06 7.1 5.23 3.63 2.545 1.73 7.21 3.9025 11.56 5.8275 5.9975 3.905 1.32 Pseudochirops archeri M 148 F 6.28 2.94 7.94 5.29 3.87 3.16 2.02 7.91 4.44 12.45 6.54 6.08 4.61 1.83 Pseudochirops archeri M 8950 M 6.29 3.01 7.77 5.41 3.74 3.16 2.82 7.86 4.22 13.03 6.11 6.82 4.65 1.94 Average for P. archeri 6.285 2.975 7.855 5.35 3.805 3.16 2.42 7.885 4.33 12.74 6.325 6.45 4.63 1.885 Spilocuscus maculatus M 23597 F 10.93 5.43 12.94 10.4 7.19 4.81 4.25 13.05 7.2 22.97 11.33 12.3 7.39 3.13 Spilocuscus maculatus M 11634 M 8.67 5.96 12.44 8.71 6.05 5.6 4.23 12.42 6.72 19.85 9.94 10.87 7.3 2.22 Spilocuscus maculatus M 32783 ? 9.33 5.18 11.62 8.9 6.43 4.83 3.08 11.5 6.39 19.95 10.13 10.66 6.99 1.42 Average for S. maculatus 9.6433 5.5233 12.333 9.3367 6.557 5.08 3.85333 12.323 6.77 20.92 10.46667 11.27667 7.2266667 2.25666667 Trichosurus fuliginosus S 946 ? 7.42 6.54 12.24 9.34 6.47 4.85 3.47 12.95 5.67 18.92 9.41 10.48 6.8 2.9 Trichosurus vulpecula M 12686 ? 6.77 4.17 10.31 7.7 5.72 2.85 3.03 10.86 4.73 16.71 7.2 9.64 6.04 2.36 Trichosurus vulpecula S 495 M 5.55 4.54 9.49 6.35 5.23 3.5 3.2 9.64 5.32 14.9 6.13 9.05 5.62 1.45 Trichosurus vulpecula M 34417 ? 6.68 7.42 11.64 10.33 7.21 3.56 2.77 11.7 8.42 18.47 9.37 11.06 7.12 1.86 Average for T. vulpecula 6.605 5.6675 10.92 8.43 6.158 3.69 3.1175 11.288 6.035 17.25 8.0275 10.0575 6.395 2.1425 Onychogalea fraenata S 1201 ? 17.25 9.48 12.08 15.23 11.1 8.15 5.65 11.42 10.42 29.73 15.22 15.93 10.95 4.26 Onychogalea fraenata A 18590 ? 15.3 8.1 10.2 12.29 9.14 6.61 3.47 9.48 10.17 24.8 14.9 12.98 7.52 3.16 Onychogalea fraenata M 12663 ? 15.38 8.65 16.15 14.72 10.86 5.16 4.35 11.21 10.57 29.08 16.18 16.1 9.6 4.18 Average for O. fraenata 15.977 8.7433 12.81 14.08 10.37 6.64 4.49 10.703 10.387 27.87 15.43333 15.00333 9.3566667 3.86666667 Aepyprymnus rufescens M 34608 ? 14.74 7.6 10.62 13.33 10.19 6.22 5.12 9.46 10.65 24.15 15.04 11.81 8.26 3.74 Aepyprymnus rufescens M 37142 M 15.67 9.52 14.43 15.85 10.34 7.38 4.59 11.29 10.08 27.42 15.26 13.95 7.93 3.59 Aepyprymnus rufescens M 22183 ? 12.68 7.28 9.77 10.66 8.75 4.93 4.38 8.54 9.26 21.58 13.24 10.5 7.31 2.78 Aepyprymnus rufescens M 9104 M 12.6 7.1 10.8 12.64 9.32 4.49 4.67 9.37 8.82 21.57 12.5 10.72 7.6 3.05 Aepyprymnus rufescens M 36622 F 12.43 6.32 10.16 10.35 8.53 4.35 4.44 8.78 7.97 21.21 11.89 11.11 7.93 3.5 Aepyprymnus rufescens S 1308 ? 12.1 7.49 10.85 13.17 9.57 4.63 4.32 9.23 9.18 22.03 12.72 11.72 8.51 3.79 Aepyprymnus rufescens M 9697 M 13.34 7.6 10.88 12.83 10.19 5.5 5.11 10.02 10.65 23.57 13.87 12.22 8.49 3.61 Aepyprymnus rufescens A 1900 ? 13.6 6.99 9.48 11.01 9.18 5.06 4.5 9.19 8.86 23.23 13.02 10.99 7.13 3.69 Aepyprymnus rufescens AR 19239 ? 13.3 7.48 10.78 12.03 9.48 4.92 4.93 9.18 10.63 23.49 14.38 11.23 8.13 3.34 Average for A. rufescens 13.384 7.4867 10.863 12.43 9.506 5.276 4.67333 9.4511 9.5667 23.14 13.54667 11.58333 7.9211111 3.45444444

89 CHAPTER 3

APPENDIX B2. Raw calcaneum measurements for modern marsupial species represented by single specimens

SPECIES SPECIMEN 0NE TWO THREE FOUR FIVE SIX SEVEN EIGHT NINE TEN ELEVEN TWELVE THIRTEENFOURTEEN Burramys parvus M 9942 1.26 0.9 2.43 1.72 1.08 0.79 0.54 2.49 1.38 3.46 1.6 2.12 1.38 0.36 Hemibelideus lemuroides M 525 3.98 2.74 6.81 4.35 4.36 3.34 2.77 7.52 4.58 10.7 5.37 5.64 4.54 0.55 Petauroides volans M 34653 5.74 4.35 8.8 6.65 5.08 4.11 3.59 9.36 6.89 13.02 6.5 7.45 5.06 1.14 Petaurus breviceps M 35708 2.75 1.03 4.2 2.26 1.8 0.91 0.62 4.13 1.84 5.83 2.81 3.39 1.89 0.75 Phalanger gymnotis M 21907 7.36 5.44 10.19 7.87 6.28 4.24 3.23 10.18 6.01 16.81 8.38 8.98 5.87 1.99 Phalanger orientalis M 27197 5.84 3.38 9.64 6.68 4.91 3.64 2.74 9.67 5.38 14.69 6.18 9 5.12 1.38 Pseudochirulus cf. canescens S 1077 4.95 3.08 7.47 5.23 4.67 2.73 2.05 7.26 3.89 10.61 4.84 5.27 4.18 1.77 Lasiorhinus latifrons M 13515 14.15 11.99 18.04 14.52 13.2 6.14 4.51 18.09 14.74 30.49 17.97 15.94 10.59 3.04 Vombatus ursinus P 793 15.08 12.25 17.64 15.5 10.2 4.42 3.9 17.82 14.74 31.06 20.3 13.21 13.74 2.79 Bettongia gaimardi cuniculus S 1682 11.54 6.72 9.1 9.8 7.87 3.97 3.71 7.47 7.47 19.91 11.7 9.29 6.84 2.51 Bettongia penicillata M 36569 10.69 5.43 8 9.27 6.46 3.33 4.03 6.64 7.33 17.38 10.68 6.52 5.64 2.14 Dendrolagus dorianus M 32134 11.73 13.66 16.33 19.19 12.3 8.35 8.54 14.73 12.86 26.82 13.49 14.92 8.82 3.02 Dendrolagus goodfellowi M 9145 14.83 11.88 15.31 19.85 12.4 7.4 7.28 13.77 10.24 30.18 15.05 14.98 8.73 3.99 Dendrolagus inustus M 25390 21.43 16.5 18.04 27.21 17.8 11.3 12.7 17.27 13.94 36.85 22.22 17.87 11.81 5.76 Dendrolagus lumholtzi M 9102 13.34 10.92 15.06 19.29 12.7 8.15 8.84 14.16 11.81 27.76 14.3 14.07 9.18 4.72 Dendrolagus matschii M 9146 10.84 12.57 14.38 18.86 12.6 8.94 7.72 13.46 10.66 23.44 11.04 11.88 8.15 4.22 Dendrolgus scottae M 25397 16.02 14.01 13.77 19.9 11.7 7.58 8.89 13.2 11.66 27.75 16.99 11.59 8.38 3.69 Dorcopsis afrata M 19462 17.14 11.58 13.92 16.2 13.3 6.48 6.52 10.85 11.35 29.42 17.57 13.81 8.95 5.1 Dorcopsis muelleri M 32341 23.06 12.73 13.78 19.63 14 7.54 7.73 13.92 13.75 35.88 19.82 18.91 10.48 5.23 Hypsiprymnodon moschatus SAM M11940 5.32 3.81 5.86 5.3 4.63 3.14 0.99 6.1 4.21 11.05 5.38 4.5 3.7 1.05 Lagorchestes hirsutus M 37520 10.63 5.69 9.45 9.71 7.17 4.66 3.1 7.4 6.8 19.09 9.99 9.52 5.71 2.66 Macropus M 37206 17.23 13.13 26.45 24 19.7 9.16 8.51 28 23.77 42.58 22.74 21.65 18.82 7.19 Macropus agilis M 9164 29.96 14.01 22.08 22.27 15.6 9.81 9.38 17.15 15.36 49.49 27.99 22.59 13.57 5.96 Macropus eugenii M 33667 16.75 10.6 15.22 12.57 11 6.46 4.91 11.47 10.29 30.56 16.18 15.53 9.11 3.32 Macropus fuliginosus M 33374 44.62 19.33 27.73 28.49 23.5 15.2 10.38 24.11 25.8 71.07 42.55 32.83 20.05 9.21 Macropus giganteus M 37256 53.86 24.34 32.14 34.06 26.6 19.9 13.04 27.87 26.22 84.96 48.13 37.55 23 9.5 Macropus parma M 37503 16.93 9.77 14.46 16.69 10 6.38 5.28 11.54 10.39 29.23 16.79 14.47 8.71 3.98 Macropus parryi S 1144 26.05 15.34 21.31 24.83 16.1 10.8 7.35 17.33 16.73 49.17 25.03 21.75 13.91 5.53 Macropus rufus M 33372 53.6 26.36 36.49 31.69 27.5 20.4 13.48 31.44 32.52 88.63 47.52 40.47 24.72 8.33 Onychogalea unguifera M 21593 23.38 10.21 13.97 15.77 12.5 7.94 5.07 12.68 13.7 36.59 20.99 17.17 10.22 3.66 Petrogale brachyotis M 33804 13.03 7.96 11.29 12.58 8.77 6.14 7.79 9.11 8.58 22.23 14.35 8.42 6.75 2.88 Petrogale mareeba M 9187 14.68 8.65 13.67 13.81 9.12 7.25 5.19 10.65 8.84 26.13 15.49 9.81 7.29 2.7 Petrogale penicillata M 36619 18.24 11.41 17.98 16.68 12.4 9.21 6.57 14.2 12.61 33.76 19.64 14.18 9.44 3.41 Petrogale persephone M 22833 18.63 10.51 16.57 16.81 11.9 9.57 8.17 13.75 13.59 33.93 19.3 14.38 10.15 3.39 Potorous tridactylus UNSWZ 399 9.36 5.31 7.45 9.66 6.76 2.84 2.88 6.36 6.81 15.61 9.21 6.95 5.11 2.54 Wallabia bicolor AR 4830 27.57 15.39 24.22 24.87 16.7 10.7 9.37 17.36 19.62 48.36 27.53 23.86 14.25 6.48 Antechinus agilis M 33544 1.22 1.06 2.25 1.62 1.44 0.6 0.54 2.12 1.2 3.36 1.88 1.74 0.95 0.28 Antechinus stuartii M 35638 1.34 1.05 2.43 1.46 1.31 0.76 0.57 2.39 1.4 3.56 1.79 1.83 1.05 0.49 Dasyurus viverrinus P 750 6.31 4.11 7.82 5.76 5.47 2.1 1.64 7.51 4.66 12.99 6.41 7.03 3.8 1.52 Parantechinus apicalis M 22256 1.81 1.06 2.87 1.93 1.74 0.87 0.62 2.87 1.4 4.3 2.28 2.43 1.07 0.62 Phascogale tapoatafa M 33612 3.38 2.01 4.84 3.1 2.13 1.51 1.05 4.73 2.95 7.59 3.87 4.32 2.03 0.82 Sarcophilus harrisii P 714 12.28 6.73 12.17 10.48 8.37 3.03 2.68 12.43 7.94 22.37 13.63 10.63 6.55 3.18 Sminthopsis murina M 24662 1.3 0.73 2.25 1.35 1.08 0.59 0.26 2.2 0.87 3.23 1.44 1.95 0.85 0.39 Thylacinus cynocephalus M 763 26.54 13.49 21.3 20.54 16.7 10.4 7.95 20.31 17.18 45.06 28.9 20.48 15.19 5.68 Echymipera kalubu M 24591 8.1 4.56 7.1 5.56 3.7 2.56 2.12 7.48 4.93 14.73 8.71 6.84 4.18 1.87 Echymipera rufescens M 30817?? 8.11 4.51 7.94 6.19 4 2.98 2.49 7.89 4.34 16.35 8.13 8.37 4.6 1.25 Isoodon obesulus M 11135 6.62 3.85 6.21 5.31 3.14 2.26 1.59 6.87 4.19 12.79 7.29 6.8 3.51 1.05 Macrotis lagotis M 22184 10.34 7.81 11.94 7.91 5.85 2.68 1.89 10.87 6.36 19.8 10.09 10.34 5.55 2.57 Perameles nasuta M 11238 10.67 6.03 9.91 7.45 4.12 3.67 3.61 9.04 6.5 19.89 11.34 9.26 5.33 0.77 Peroryctes raffrayana M 30817?? 8.53 5.14 8.34 6.24 4.11 3.03 2.97 8.35 4.84 16.12 8.76 8.4 4.89 1.79 Notoryctes typhlops SAM 637 9.34 2.63 8.72 5.34 2.08 1.22 3.31 9.61 3.95 17.11 8.86 9.91 0.94 1.86

90 APPENDIX B3. Modern marsupial calcaneal measurements for species used as 'unknowns' in analysis

SPECIES SPECIMEN SEX ONE TWO THREE FOUR FIVE SIX SEVEN EIGHT NINE TEN ELEVEN TWELVE THIRTEEN FOURTEEN Isoodon macrourus M 35567 ? 12.76 5.28 10.66 8.2 4.95 3.79 3.41 10.11 7.61 22 12.45 10.48 5.61 2.2 Isoodon macrourus M 35906 F 10.18 4.79 8.44 6.7 3.33 2.11 2.96 8.5 6.15 18.48 10.56 9.16 4.82 1.16 Isoodon macrourus M 35411 F 9.75 4.23 8.07 7.01 2.94 2.69 2.28 8.05 5.97 17.98 10.26 8.07 4.21 1.52 Isoodon macrourus M 36017 ? 11.1 5.53 9.28 7.42 4.13 3.25 2.87 9.39 6.43 20.28 10.99 9.39 4.66 1.88 Isoodon macrourus M 36016 F 9.86 4.31 9.48 6.99 4.02 3.22 2.7 9.22 6.14 18.16 9.9 8.63 4.23 1.58 Isoodon macrourus AR 7639 ? 12.17 5.78 9.34 7.4 4.69 3.16 3.35 8.55 7.2 20.45 12.08 8.95 4.99 2.41 Pseudocheirus peregrinus M 35238, juvenile F 3.39 2.42 5.41 4.55 3.2 1.96 2 6.15 3.13 8.49 3.88 5.3 3.6 1.14 Spilocuscus maculatus M 23627, juvenile ? 8.17 4.17 10.94 8.14 6.02 4.32 3.1 10.88 6.18 18.15 9.04 10.07 6 1.51 Trichosurus vulpecula M 35471, juvenile M 5.15 4 9.04 7.06 6 3.45 2.75 9.68 4.39 13.5 6.77 8.06 5.26 1.27 Onychogalea fraenata M 37128, juvenile M 16.11 8.66 15.16 15.1 11.01 7.42 4.76 11.17 10.76 30.05 16.27 16.27 9.04 3.48 Onychogalea fraenata A 18591, juvenile? ? 16.11 8.61 12.53 13.86 9.43 6.35 3.82 9.68 9.63 25.77 14.78 13.48 8.4 3.17 Aepyprymnus rufescens M 23601, no epyph F 11.11 6.15 11.21 12.86 10.09 5.9 4.7 9.57 9.45 21.92 11.32 11.92 7.38 4.18

91 CHAPTER 3

APPENDIX B4. Raw calcaneum measurements for fossil marsupials (where more than one specimen represents a morphotype, average values are given)

MORPHOTYPE SPECIMEN ONE TWO THREE FOUR FIVE SIX SEVEN EIGHT NINE TEN ELEVEN TWELVE THIRTEEN FOURTEEN (number of specimens) CS B (2) QMF 51413 CS B 10.94 4.8 10.2 9.58 6.98 3.75 3.13 8.4 7.4 21.24 13.04 10.13 6.45 2.13 CS I (1) QMF 51413 CS I 8.27 3.7 8.6 7 4.8 2.85 2.1 7.9 6 16.56 9.36 8.4 3.3 1.6 CS S1 (1) QMF 51413 CS S1 16.74 7.81 13.52 16.93 12.1 6.63 6 13.2 11.65 30.99 17.54 14.98 9.33 2.6 CS S2 (1) QMF 51413 CS S2 17.82 11.24 15.28 19.86 13.26 7.94 6.74 13.61 12.35 33.72 17.73 15.31 10.54 3.47 CS T (1) QMF 51413 CS T 12.75 6.33 10.75 9.34 6.88 3.8 3.44 9 7.7 21.9 11.9 10.2 5.98 1.68 CS V (1) QMF 51413 CS V 21.5 7.75 15.39 16.81 13.86 7.45 5.78 15.3 13.76 36.3 22 15.95 10.62 3.04 CS C (3) QMF 51413 CS C 17.77 9.8 17.45 15.8 11.93 5.7 5.75 16.05 13.55 38.87 20.17 17.08 9.55 2.98 CS J (1) QMF 51413 CS J 7.2 7.3 12.1 12.05 9 4.7 4.3 11.9 9.9 19.04 7.65 12.75 7.7 3.75 CS U (1) QMF 51413 CS U 10.25 8.3 13.3 13.53 10.3 5.55 5.25 12.5 9.4 22.7 9.53 13.6 7.9 2.83 CS Q (1) QMF 51413 CS Q 5.05 2.85 8.55 6 4.3 2.25 1.65 8.45 4.7 13.85 5.6 8.15 4.35 1.34 CS R (1) QMF 51413 CS R 3.63 2.4 5.16 3.28 2.85 1.05 1.07 4.6 2.7 8.32 4.26 3.6 2 0.5 CS P (1) QMF 51413 CS P 5.8 3 7.05 4.85 2.45 2.25 1.4 7.2 3.2 12.95 6.45 6.95 2.85 1.2 CS a (15) QMF 51413 CSa 7.35 3.98 7.15 5.75 3.6 2 1.95 6.98 5.18 14 8.13 7.3 3.68 2.3 CS d (6) QMF 51413 CSd 9 5.05 8.53 6.95 3.8 3.1 2.3 8.18 6.4 16.1 9.05 8.15 4.25 1.88 CS e (36) QMF 51413 CSe 4.2 2.82 5 3.73 1.83 1.38 1.08 4.28 2.73 9.18 4.81 5.03 2.2 0.58 CS f (4) QMF 51413 CSf 4.4 1.96 5.48 3.75 2.73 1.55 1.3 5.3 2.9 9.75 4.62 5.31 2.6 1.13 CS g (1) QMF 51413 CSg 5.03 2.31 4.99 3.35 2.28 1.58 0.94 4.82 3.09 9.54 5 6.3 2.46 0.83 CS h (6) QMF 51413 CSh 3.2 1.95 4.23 3.2 1.6 1.3 0.83 3.68 2.13 7.36 3.08 4.25 1.83 0.7 CS m (5) QMF 51413 CSm 4.8 2.33 6.63 5 1.93 1.83 1.6 6.3 2.85 11.53 4.75 6.65 2.83 1.2 CS n (1) QMF 51413 CSn 2.2 1.13 3.3 2.18 1.13 0.85 0.8 3.6 1.4 6.1 2.65 3.2 1.55 0.5 CS o (1) QMF 51413 CSo 3.45 1.25 3.75 2.5 1.35 0.8 0.95 3.4 1.9 7.2 3.35 3.9 1.8 0.55 CS ma (1) QMF 51413 CSma 4.09 1.98 6.3 3.66 2 1.74 1.58 6.08 2.65 10.82 4.48 6.04 2.64 1.12 CS x (1) QMF 51413 CSx 3.44 2.26 5.67 4.46 2.1 1.66 1.09 6.12 2.46 9.14 3.5 5.85 2.4 1.18 CS y (1) QMF 51413 CSy 3.87 2.2 4.71 3.96 1.81 1.39 0.81 4.6 2.15 9.24 4.17 4.86 2.37 0.56 CS z (1) QMF 51413 CSz 2.36 1.96 3.91 2.95 1.4 0.89 0.82 3.24 1.8 6.35 2.5 3.9 2 0.6 MM 1 (1) QMF 51414 MM1 11.5 6.43 10.9 10.49 7.09 4.49 3.69 9.06 7.51 21.12 11.46 10.31 5.31 2.5 MM 2 (1) QMF 51414 MM2 17.82 11.24 14.66 17.1 12.4 7.84 7.2 13.11 15.01 33.72 17.73 14.3 10.56 4.42 MM 3 (1) QMF 51414 MM3 11.79 6.46 9.94 12.04 8.21 5 3.75 8.68 6.96 20.72 11.2 9.2 6.86 2.43 MM 4 (3) QMF 51414 MM4 4.14 2.69 4.92 4.01 1.5 1.6 1 4.88 2.61 9.26 4.65 4.74 2.09 0.6 MM 5 (1) QMF 51414 MM5 2.06 1.2 2.7 1.81 1.16 0.44 0.24 2.56 1 5 3.05 2.59 2.59 0.56 MM 6 (1) QMF 51414 MM6 7.8 3.96 6.9 6.1 2.96 2 1.94 7 4.41 14.13 8.21 6.4 3.59 0.9 MM 7 (1) QMF 51414 MM7 9 5.05 8.4 6.7 2.85 2.5 2.24 7.8 5.49 16.1 9.05 8.31 3.61 1.72 MM 8 (1) QMF 51414 MM8 3.84 2.26 4.23 3.2 1.6 1.3 0.83 3.68 2.13 7.36 3.9 4.25 1.83 0.7 R. flanneryi (1) QMF 31456 31.28 20.61 28.75 31.18 21.49 13.42 9.77 20.51 23.75 58.3 33.16 29.34 19.64 8.95 Ec G (1) QMF 51415 Ec G 19.28 9.74 15.65 15.88 14.31 6.18 6.95 12.71 12.61 33.92 18.88 15.16 9.41 4.57 Ec H (2) QMF 51415 Ec H 18.62 9.87 15.36 17.26 13.24 8.2 6.72 13.44 14.28 34.96 19.06 15.65 11.2 4.61 Ec J (3) QMF 51415 Ec J 17.82 11.24 15.75 19.1 13.67 8.9 6.86 14.41 12.69 33.72 17.73 15.95 10.33 2.84 Ec K (4) QMF 51415 Ec K 20.2 7.75 14.9 17.81 14.59 6.4 7.24 14.79 13.52 34.29 20.93 15.14 10.27 3 Ec L (3) QMF 51415 Ec L 21.02 9.11 18.36 16.84 13.9 7.79 6.42 15.93 12.7 39.21 21.29 17.88 11.31 4.34 Ec M (1) QMF 51415 Ec M 11.79 6.46 9.36 11.31 9.71 4.91 4.32 8.91 7.9 20.72 11.2 9.7 7.15 2.01 Ec N (1) QMF 51415 Ec N 11.24 4.94 9.6 7.7 7.56 3.74 3.6 8.69 7.54 20.89 10.43 10.34 6.51 2.06 Ec O (2) QMF 51415 Ec O 17.95 11.5 16.06 14.81 12.18 6.81 5.05 13.66 11.94 30.89 19.9 14.81 10.36 3.5 Trichosurus (1) QMF 36438 6.55 4.25 10.22 7.28 5.65 3.11 2.45 10.29 7.2 16.95 7.7 8.29 5.24 0.99 Ec E (1) QMF 51415 Ec E 1.98 1.05 2.2 1.85 1 0.61 0.37 2.14 1.15 4.18 1.86 2.49 0.95 0.35 Ec F (1) QMF 51415 Ec F 2.1 1.31 2.65 1.7 1.1 0.65 0.55 2.5 1.3 4.54 2.25 2.4 1.2 0.5 Ec A (5) QMF 51415 Ec A 6.69 2.7 6.3 4.3 2.5 1.3 0.96 6.2 3.15 13.13 6.73 6.6 2.8 0.1 Ec B (2) QMF 51415 Ec B 6.24 3 6.6 5.24 2.26 1.64 1.2 6.17 4.01 13.04 6.49 6.44 3.31 0.8 Ec C (2) QMF 51415 Ec C 3.67 2.1 4.85 3.98 1.85 1.4 0.95 4.76 2.15 9.02 4.03 5.5 2.1 0.66 Ec D (3) QMF 51415 Ec D 1.7 1.1 2 1.4 0.7 0.5 0.4 1.9 0.8 3.35 1.7 1.56 0.98 0.3 QL A (1) QMF 50856 A 4.82 2.4 5.1 4.36 1.93 1.65 1.57 5.19 2.3 9.77 4.84 4.92 2.91 0.93 QL B (2) QMF 50856 B 4.54 2.5 5.82 4.91 2.39 2.09 1.53 6.21 3.15 10.21 4.72 5.95 3.11 1.09 QL C (1) QMF 50856 C 5.91 2.7 5.38 4.6 1.94 1.92 2.02 5.34 3.6 10.99 5.78 4.79 3.04 0.87 QL D (1) QMF 50856 D 2.61 2.11 3.43 2.51 1.38 1.01 0.67 3.22 1.4 5.55 2.63 2.87 1.72 0.82 QL E (1) QMF 50856 E 1.43 1.18 2.52 2.2 0.89 0.82 0.67 2.59 1.16 4.14 1.8 2.66 1.26 0.52 QL F (1) QMF 50856 F 8.26 4.2 7.68 6.49 2.54 2.92 2.65 8 5.06 15.05 7.92 7.53 3.94 1.86 QL G (1) QMF 50865 1.8 0.8 2 1.1 0.79 0.5 0.4 2 0.88 3.9 1.8 1.9 0.91 0.31 Nambaroo sp. 3 (1) QMF 34532 18.2 11.36 14.55 16.81 13.06 5.62 2.77 12.2 11.26 31.65 17.82 11.59 10.81 18.77

92 CHAPTER 3

APPENDIX B5. Normal probability plots for all variables (Zskewness values indicated above; Zskewness values under +/-2.58 indicate normal distribution at the 0.01 probability level).

TALONID: 3.46 CAFI: 8.30

4 1. 2 1. 1 1 3 0.9 0.8 0.7 2 0.6 Sample values Sample values 0.5 1 0.4 0.3 0.2 -2 -1 0 1 2 3 -2 -1 0 1 2 3

Normal order statistic medians Normal order statistic medians

HEAD: 0.69 LATHEAD: 2.35

2 3

Sa mpl e valu 2 es 1

Sample values 1

-2 -1 0 1 2 3 -2 -1 0 1 2 3 Normal order statistic medians Normal order statistic medians

HEADTAL: 1.81 MEDLAT: 2.07

2 1. 5 1. 4 1. 3 1. 2 1. 1 1 1 0.9 Sample values values Sample 0.8 0.7 0.6 0.5 -2 -1 0 1 2 3 -2 -1 0 1 2 3 Normal order statistic medians No rmal order statistic medians

CACU: 1.43 TOTAL: -0.88

0.2 1. 4 1. 3 1. 2 1. 1 1 0.9 0.1 0.8 Sample values

Sample values 0.7 0.6 0.5 0.4

-2 -1 0 1 2 3 -2 -1 0 1 2 3 Normal order statistic medians Normal order statistic medians

CHAPTER 4

DIETS OF RIVERSLEIGH FOSSIL MAMMALS

CHAPTER 4

THE DIETS OF RIVERSLEIGH FOSSIL MAMMALS

4.1. Introduction

Animals adapt to utilising specific food resources in order to reduce competition from other species. Food resources are related to types of vegetation available and vegetation structure in general and thus to habitat. The dietary adaptations of the species in a community may be an indication of the variety of niches available to the animals, so it follows that if the diets of fossil species can be determined from their remains then aspects of their palaeoecology such as habitat structure can be inferred.

This chapter reviews literature on the use of dental and masticatory systems and cranial features to infer the diets of mammals. Dietary, or trophic, categories are constructed for the purposes of classifying described fossil marsupial species from Camel Sputum,

Mike’s Menagerie, Encore, Ringtail, Bitesantennary and Quantum Leap Sites according to possible food intake. The inherent detail of these categories reflects the amount of reliable information obtainable from the fossils and suffices for the general purpose of making comparisons between mammal communities based on diet. The relatively broad categories relate to general habitat structure as opposed to specific details of habitat. In most cases the diet of each fossil species from the selected sites is suggested, based on dental or cranial morphology, by the author of the species description.

94 CHAPTER 4

4.2. Mammalian cranio-dental functional morphology

Mammalian teeth are differentiated into incisors, canines and premolars, for ingestion of food, and molars, for mastication (Kay and Hylander 1978, Sanson 1991). The morphological structure of the teeth is associated with the properties of the food resource used by an animal (Eisenberg 1978, Kay and Hylander 1978, Sanson 1991) and it is widely accepted that diet is reflected to some extent in dental structure and jaw morphology (Sanson 1991). In general, it is easy to distinguish a meat-eater from a plant-eater, but more difficult to distinguish diets within each of these major types.

Enlarged molar and premolar grinding surfaces, adapted for horizontal shearing, indicate herbivory. Carnivory, on the other hand, involves a simplified tooth pattern, long cutting blades on the premolars and molars (Van Valkenburgh 1988) used for vertical shearing (Van Valen 1969), and loss of crushing function (Savage 1977).

Trends towards herbivory are also apparent in cranial adaptations, for example, increased size and extent of attachment of pterygoid muscles for greater medial jaw movements, which results in more effective grinding (Eisenberg 1978, Davison and

Young 1990). Janis (1995) presents schematically the main differences between the skulls of carnivores and herbivores. These differences include: a larger momentum arm for the masseter and a larger area for insertion of the masseter in herbivores; a larger momentum arm for the temporalis and a larger area for insertion of the temporalis in carnivores; the position of the jaw joint with respect to the tooth row differs between herbivores and carnivores; the presence of a postglenoid process in carnivores and the absence of this structure in herbivores; the presence of the postorbital bar in herbivores and its absence in carnivores; and a larger and more elevated nuchal area in carnivores relative to herbivores.

95 CHAPTER 4

Eisenberg (1978) defines a herbivore as an animal that feeds on vegetation, specifically stems and foliage but not including nectar, fruit, seeds, flowers, sap or roots. Herbivores require certain adaptations in dental morphology, jaw musculature and gut morphology in order to process cellulose-rich leaves and stems as a source of energy (Moir 1968 and

Greaves 1974, cited in Eisenberg 1978). Cellulose is difficult to digest and requires chemical breakdown by cellulase enzymes (Withers 1992) and more mechanical processing than the components of a non-herbivorous diet (Kay and Hylander 1978).

Plants have evolved chemical and structural defences to being consumed and consequently, the animals that consume these plants have also evolved adaptations to overcome such defences (Eisenberg 1978). Crown height is important for determining the extent of herbivory of some animals (Sanson 1991, Mendoza et al. 2002 and references within) as it allows for tooth wear with the mechanical processing of abrasive material (Eisenberg 1978). Crown height may also relate to the abrasive nature of the food, such as underground roots and tubers that are consumed with grit (see Janis 1995, p.86 for further discussion of this).

Herbivores include folivores (or browsers) and grazers. Browsers have a diet of plant material low in fibre content, and grazers consume high fibre plant material such as grasses, with intermediates also occurring (Sanson 1982). Distinctions between these herbivore strategies are apparent in molar cusps, which are often retained in browsers for greater pulping compared to grinding (Kay and Hylander 1978); increased molar surface areas and complexity of grinding surfaces in grazers; and relatively higher molar crowns in grazers (Eisenberg 1978). Mendoza et al. (2002: 224) list craniodental features that are important in distinguishing browsing from grazing ungulates. Further distinctions are seen in macropodines, with browsers having occlusally straight tooth

96 CHAPTER 4

rows, mainly antero-posterior direction of movement between molars, and no molar progression, in contrast to grazers (Sanson 1982). Premolar and incisor structure also distinguishes browsers from grazers with, for example, the premolar dentition of folivorous prosimians and marsupials having shearing capabilities (Kay and Hylander

1978).

Animals often rely on a combination of food sources and identifying each of these may be difficult. Often the only way to distinguish species that consume a variety of food types is on the basis of the dominant dietary adaptation if it is apparent. For example, frugivory is often associated with folivory; the extent of folivory may be determined by the degree of molar complexity and addition of accessory crests, which emphasise a cutting-shearing action and are conducive to increase in leaf consumption (Lanyon and

Sanson 1986).

Body size is of some relevance in determining diet. Insectivorous arboreal mammals are consistently smaller than folivorous arboreal mammals (Kay and Hylander 1978), and due to energetic restrictions and problems of detoxification, arboreal folivores have a lower limit on size (Eisenberg 1978). Arboreal folivores also have an upper limit on size, imposed on them by the weight-bearing capacities of trees and the mechanics of movement in trees (Eisenberg 1978).

Carnivory can occur to varying extents, mixed with insectivory or herbivory.

Determining the degree of carnivory is a matter of weighing up the different functional components of the dentition. Wroe (1998) discusses this with respect to propleopine macropodoids. Vertical and horizontal shearing capacity in dentition is apparent in

97 CHAPTER 4

carnivores with the relative significance of each of these shear types varying with the importance of vertebrate prey versus plant matter in the diet (Van Valkenburgh 1989,

Wroe 1998). Further to this, Wroe (1998) points out that each molar of dasyuromorphian carnivores has both vertical and horizontal shearing capacity and species that consume greater amounts of invertebrates relative to vertebrates show relatively equal vertical and horizontal shearing complexes, whereas in obligate carnivores, vertical shear dominates. The killing behaviour of placental carnivorans can be determined from incisor and dental arcade shape (Biknevicius et al. 1996) and from canine shape and strength (Van Valkenburgh and Ruff 1987). Some carnivores include bone in their diets. Enlarged premolars are characteristic of a bone crushing function

(Van Valkenburgh 1988). Wroe (1998) discusses the premolars of a fossil species from

Encore Site, Ganbulanyi djadjinguli, with respect to this. A summary of cranial features, also useful in identifying carnivory in mammals, is presented by Wroe (1998).

It is accepted that dental and cranial morphologies reflect diet to some extent, however,

Sanson (1991) cautions that teeth alone are not always an unquestionable predictor of diet. Wroe (1998) notes that the inference of diet should not be based on any single feature in isolation, but must take into consideration characters in combination. There are many interacting factors involved in food intake that include, aside from dentition, gut morphology and behaviour of an animal (Sanson 1991). However, on a broad level, it is possible to make trophic distinctions between animals based on dentition. Janis

(1990) finds a number of convergences between kangaroos and ungulates that are correlated with dietary preference but are obviously not taxon-specific. This implies that these particular characters are determined by function rather than by phylogeny, and thus there are cases of animals displaying analogous mechanical solutions to utilising

98 CHAPTER 4

similar food types. Conversely, Seligsohn and Szalay (1978) show that closely related, herbivorous lemurids display very different masticatory adaptations related to their specific diets, so that a selective alteration of the mechanical design is apparent in the dentition as a result of dietary preference.

4.3. Trophic categories

Predictions of diet based on fossil remains (dental and/or cranial) are most reliable when they are general and in cases where modern representatives from the same clade, with homologous morphology are used as analogues (Lauder 1995).

Table 4.3.1 describes trophic categories in terms of food type consumed. These categories are broad and indicate the major food type intake, but in all cases, other food types may also be consumed. To some extent, the categories reflect the structural position of feeding among habitat substrates (e.g. herbs, roots, tubers, fungi and fruiting bodies are all consumed from the ground; leaves, fruit and flowers are consumed on the vegetation itself in most cases). These categories are used for the classification of extant mammals in the palaeoecological analyses in Chapter 5. Fossil taxa are also classified among these categories, however, the browser/frugivore category is combined with the frugivore/pollen/nectar category because animals often consume these food types in combination and, in most cases, are active in the same broad structural level of the vegetation for consumption of these food types.

99 CHAPTER 4

TROPHIC CATEGORY DIET 1 ROOTS/FUNGI herbs, roots, tubers, fungi and fruiting bodies shoots and stems of grasses and herb vegetation and 2 GRAZER ground fruit 3 GRANIVORE majority seeds, nuts, some grasses, fungi or fruit majority leaves or flowers; or majority fruit, 4 BROWSER/FRUGIVORE supplemented with leaves predominantly fruit, pollen or nectar, in any order of 5 FRUGIVORE/POLLEN/NECTAR preference; no invertebrates or leaves combination of invertebrates including snails with 6 OMNIVORE either nectar, fruit, exudates, seed/plant matter, or roots/tubers majority invertebrates; some fruit, nectar, seeds, plant 7 INSECTIVORE fibre may be included hunters of large and small vertebrates; scavengers; 8 CARNIVORE some invertebrates may be included

Table 4.3.1. Description of trophic categories. Categories 4 and 5 have been combined for the analyses in Chapter 5 because they are difficult to distinguish for fossil taxa. The combination of these two categories is referred to as ‘browser/frugivore’.

Tables 4.3.2, 4.3.3, 4.3.4, 4.3.5, 4.3.6, and 4.3.7 list species or higher level taxa from the six Riversleigh sites and trophic categories they have been assigned to based on inferred diets as indicated by authors of species descriptions, or in the case of higher level taxa, from closest relatives or modern analogues. Species lists have been compiled using

Arena’s (2004) lists of published species and species described in unpublished theses for each site. Species described in theses but not published are indicated by square brackets in the tables. Higher-level taxa have been identified by preparators at the

University of New South Wales and recorded in the catalogue of Riversleigh specimens.

100 CHAPTER 4

FAMILY ANIMAL DIET REFERENCE Yalkaparidontidae Yalkaparidon coheni insectivore Long et al. (2002) Notoryctidae ? insectivore inferred from diet of extant Notoryctes typhlops, Johnson (1998) Thylacinidae Ngamalacinus carnivore Muirhead (1997) timmulvaneyi Wabulacinus ridei carnivore Muirhead (1997) Yaralidae Yarala burchfieldi insectivore Muirhead (1994), Muirhead and Filan (1995) Insertae sedis [Bulunga palara] insectivore Muirhead (1994) (Peramelemorph- [Galadi grandis] omnivore extrapolated from Muirhead (1994) ians) [Gunawidgi tubus] omnivore extrapolated from Muirhead (1994) [Madju ignotae] omnivore extrapolated from Muirhead (1994) [Madju variae] omnivore extrapolated from Muirhead (1994) Wynyardiidae Namilamadeta browser/frugivore inferred from traces of selenodont crest pattern on Namilamadeta snideri dentition (Long et al. 2002) Phascolarctidae Nimiokoala greystanesi browser/frugivore Black and Archer (1997) Diprotodontidae Neohelos sp. browser/frugivore inferred from diet given for Zygomaturinae in Long et al. (2002) Neohelos tirarensis browser/frugivore inferred from diet given for Zygomaturinae in Long et al. (2002) Palorchestidae Propalorchestes browser/frugivore K. Black pers. comm. (2005) ponticulus Palorchestes browser/frugivore Flannery and Archer (1985) Phalangeridae [Ilungalya aletes] browser/frugivore Crosby (2002) [Ilungalya lygaeus] browser/frugivore Crosby (2002) [Onirocuscus inversus] browser/frugivore Crosby (2002) [Onirocuscus silvacultrix] browser/frugivore Crosby (2002) Wyulda asherjoeli browser/frugivore Crosby et al. (2001) Pilkipildridae ? omnivore Archer et al. (1987) Acrobatidae ? omnivore based on modern acrobatid species, Woodside (1998) Burramyidae Burramys brutyi browser/frugivore inferred from bunodont molars described by Brammal and Archer (1997) Pseudocheiridae Marlu cf. Kutjamarpensis browser/frugivore inferred from selenodont dentition Paljara maxbourkei browser/frugivore Bassarova et al. (2001) Paljara nancyhawardae browser/frugivore Bassarova et al. (2001) Pildra [sp. 2] browser/frugivore Carbry (2001) Insertae sedis Djaludjangi yadjana omnivore Brammall (1999) Macropodidae Bulungamaya delicata browser/frugivore Flannery et al.(1983) Gangaroo bilamina browser/frugivore Cooke (1996), Flannery et al.(1983) Nowidgee matrix omnivore Cooke (1996) Wabularoo naughtoni browser/frugivore Flannery et al.(1983) Potoroidae ? omnivore based on Potoroinae, Sanson (1982) Hypsiprymnodont- Hypsiprymnodon omnivore Wroe et al. (1998), Long et al. (2002) and based on idae modern species (Strahan, ed. 1998) Ekaltadeta ima omnivore Wroe (1997), Wroe et al. (1998), Wroe (1996), Archer and Flannery (1985) Balbaridae Balbaroo gregoriensis browser/frugivore inferred from Flannery et al.(1983) Ganawamaya acris browser/frugivore extrapolated from Cooke (1992, thesis) Nambaroo [sp.5] browser/frugivore Cooke (1996) Wururoo sp. browser/frugivore Cooke (1996) Hipposideridae Brachipposideros insectivore S. Hand pers. comm. (2005) Megadermatidae Macroderma carnivore S. Hand pers. comm. (2005)

Table 4.3.2. Diets of species and higher-level taxa as indicated by references for Camel Sputum Site

101 CHAPTER 4

FAMILY ANIMAL DIET REFERENCE Yalkaparidont- Yalkaparidon insectivore Long et al. (2002) idae Notoryctidae ? insectivore inferred from diet of extant Notoryctes typhlops, Johnson (1998) Thylacinidae Thylacinus carnivore Muirhead (1992, 1997), Wroe (1999) macknessi ? Yaraloidea, gen. and insectivore inferred from diet of Yarala burchfieldi, Muirhead and sp. indet Filan (1995) Wynyardiidae ? browser/frugivore inferred from traces of selenodont crest pattern on Namilamadeta snideri dentition (Long et al. 2002) Diprotodontidae Neohelos sp. browser/frugivore inferred from diet given for Zygomaturinae in Long et al. (2002) Neohelos tirarensis browser/frugivore inferred from diet given for Zygomaturinae in Long et al. (2002) Palorchestidae Palorchestes browser/frugivore Flannery and Archer (1985) Phalangeridae Wyulda asherjoeli browser/frugivore Crosby et al. (2001) [Illungalya aletes] browser/frugivore Crosby (2002) Pilkipildridae ? omnivore Archer et al. (1987) Acrobatidae ? omnivore based on modern acrobatid species, Woodside (1998) Insertae sedis Djaludjangi yadjana omnivore Brammall (1999) Macropodidae Gangaroo bilamina browser/frugivore Cooke (1996) Flannery et al.(1983) Wabularoo browser/frugivore Flannery et al.(1983) naughtoni Hypsiprymnodont Hypsiprymnodon omnivore Wroe et al. (1998), Long et al. (2002) and based on idae modern species (Strahan, ed. 1998) Ekaltadeta ima omnivore Wroe (1997), Wroe et al. (1998), Wroe (1996), Archer and Flannery (1985) Hipposideridae Brachipposideros insectivore S. Hand pers. comm. (2005) Megadermatidae Macroderma carnivore S. Hand pers. comm. (2005)

Table 4.3.3. Diets of species and higher-level taxa as indicated by references for Mike’s Menagerie Site

FAMILY ANIMAL DIET REFERENCE Yalkaparidont- Yalkaparidon insectivore Long et al. (2002) idae Notoryctidae ? insectivore inferred from diet of extant Notoryctes typhlops, Johnson (1998) ? Yaraloidea, gen. and insectivore inferred from diet of Yarala burchfieldi, Muirhead and sp. indet Filan (1995) Diprotodontidae ? browser/frugivore based on diets for other Riversleigh diprotodontids and for Zygomaturinae (Long et al. 2002) Pilkipildridae ? omnivore Archer et al. (1987) Phalangeridae [Illungalya aletes] browser/frugivore Crosby (2002) Petauridae ? omnivore based on diets of modern peataurids (Strahan, ed. 1998) Macropodidae Bulungamaya browser/frugivore Flannery et al.(1983) delicata Hypsiprymnodont Hypsiprymnodon omnivore Wroe et al. (1998), Long et al. (2002) and based on idae modern species (Strahan, ed. 1998) Balbaridae Nambaroo sp. 3 browser/frugivore Cooke (1996) Hipposideridae 5 species insectivore S. Hand (pers. comm. 2002)

Table 4.3.4. Diets of species and higher-level taxa as indicated by references for Quantum Leap Site

102 CHAPTER 4

FAMILY ANIMAL DIET REFERENCE Dasyuridae Ganbulanyi carnivore/bone Wroe (1998) djadjinguli cracker Thylacinidae Thylacinus sp. cf. T. carnivore Muirhead (1992, 1997), Wroe (1999) macknessi Insertae sedis Mayigriphus orbus insectivore Muirhead (1995) ? Yaraloidea, gen. and insectivore inferred from diet of Yarala burchfieldi, Muirhead and sp. indet Filan (1995) Phascolarctidae Litokoala browser/frugivore Black (1992) kanunkaensis Phascolarctos sp. browser/frugivore Black (1992) Vombatidae cf. Warenja sp. grazer? Archer et al. (1994), Wroe (1997) Thylacoleonidae Wakaleo carnivore Gillespie (1997), Muirhead (1997) vanderleurei Diprotodontidae Neohelos sp. browser/frugivore Black (1992) and inferred from diet of Zygomaturinae Long et al.(2002) Palorchestidae Palorchestes anulus browser/frugivore inferred from Black (1997) and Flannery and Archer (1985) Palorchestes sp. browser/frugivore inferred from Flannery and Archer (1985) Phalangeridae Trichosurus browser/frugivore Crosby (2002) [euphrasia] Burramyidae Burramys brutyi browser/frugivore inferred from bunodont molars described by Brammal and Archer (1997) Pseudocheiridae cf. Pseudocheirops browser/frugivore based on modern species of this genus sp. Pildra [sp. 2] browser/frugivore Carbry (2001) Macropodidae Gangaroo new sp. browser/frugivore Cooke (thesis), and inferred from diet of Bulungamayines Flannery et al. (1983) Wanbaroo hilarus browser/frugivore Cooke (1999) Wanbaroo sp. browser/frugivore based on diet of W. hilarus and Bulungamayines (Flannery et al. 1983) Sthenurinae cf. Hadronomas sp. browser/frugivore Cooke (1996) Rhyzosthenurus browser/frugivore Kirkham (2004) flanneryi Hypsiprymnodont Ekaltadeta ima omnivore Wroe (1996, 1997), Wroe et al. (1998), Archer and idae Flannery (1985) Ekaltadeta omnivore Wroe (1996, 1997), Wroe et al. (1998), Archer and jamiemulvaneyi Flannery (1985) Indet. 12 species insectivore? S. Hand pers. comm. (2005) Chiroptera Megadermatidae Macroderma carnivore S. Hand pers. comm. (2005)

Table 4.3.5. Diets of species and higher-level taxa as indicated by references for Encore Site

FAMILY ANIMAL DIET REFERENCE Dasyuridae Barinya wangala carnivore Long et al. (2002) ? Yaraloidea, gen. and sp. indet insectivore inferred from diet of Yarala burchfieldi, Muirhead and Filan (1995) Phalangeridae [Onirocuscus silvacultrix] browser/frugivore Crosby (2002) Macropodidae Gangaroo bilamina browser/frugivore Cooke (1996), Flannery et al.(1983) Hipposideridae Brachipposideros nooraleebus insectivore S. Hand pers. comm. (2005) Brachipposideros watsoni insectivore S. Hand pers. comm. (2005) Brevipalatus mccolloughi insectivore S. Hand pers. comm. (2005) Rhinonicteris tedfordi insectivore S. Hand pers. comm. (2005) Riversleigha williamsi insectivore S. Hand pers. comm. (2005) Xenorhinos halli insectivore S. Hand pers. comm. (2005) Megadermatidae Macroderma carnivore S. Hand pers. comm. (2005)

Table 4.3.6. Diets of species and higher-level taxa as indicated by references for Bitesantennary Site

103 CHAPTER 4

FAMILY ANIMAL DIET REFERENCE Notoryctidae ? insectivore inferred from diet of extant Notoryctes typhlops, Johnson (1998) Thylacinidae Maximucinus muirheadae carnivore Wroe (2001) Nimbacinus new sp. carnivore based on diet information for Nimbacinus dicksoni, Long et al. (2002) Insertae sedis [Gunawidgi tubus] omnivore extrapolated from Muirhead (1994) Diprotodontidae Nimbadon lavarackorum browser/frugivore inferred from diet of Zygomaturinae Long et al. (2002) Phalangeridae ? browser/frugivore inferred from other Riversleigh phalangerids Acrobatidae ? omnivore based on modern acrobatid species, Woodside (1998) Burramyidae Burramys brutyi browser/frugivore inferred from bunodont molars described by Brammal and Archer (1997) Cercatetus omnivore based on modern species of this genus (Strahan, ed. 1998) Petauridae ? omnivore based on diets of modern peataurids (Strahan, ed. 1998) Pseudocheiridae Pseudochirops browser/frugivore based on modern species of this genus Marlu cf. Kutjamarpensis browser/frugivore inferred from selenodont dentition Ornithorhynchidae Obdurodon dicksoni carnivore Musser and Archer (1998) Hipposideridae Brachipposideros insectivore S. Hand pers. comm. (2005) Megadermatidae Macroderma carnivore S. Hand pers. comm. (2005)

Table 4.3.7. Diets of species and higher-level taxa as indicated by references for Ringtail Site

104 CHAPTER 4

4.4. References

Archer, M., Hand, S. J. and Godthelp, H. 1994. Patterns in the history of Australia’s mammals and inferences about palaeohabitats. Pp. 80-103 in History of the Australian Vegetation: Cretaceous to Recent. Ed. R. Hill. Cambridge University Press, Cambridge.

Archer, M., Tedford, R. H., and Rich, T. H. 1987. The Pilkipildridae, a new family and four new species of ?Petauroid possums (Marsupialia: Phalangerida) from the Australian Miocene. Pp. 607-627 in Possums and Opossums: Studies in Evolution. Ed. M. Archer. Surrey Beatty and Sons and the Royal Zoological Society of New South Wales, Sydney.

Arena, D. A. 2004. The geological history and development of the terrain at the Riversleigh World Heritage Area during the middle Tertiary. PhD thesis. University of New South Wales, Sydney.

Black, K. 1992. Systematics and Palaeobiology of Fossil Phascolarctids From Riversleigh. Honours Thesis. University of New South Wales, Sydney.

Black, K. and Archer, M., 1997. Nimiokoala gen. nov. (Marsupialia, Phascolarctidae) from Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41(2): 209-228.

Brammall, J. R., 1998. A new petauroid possum from the Oligo-Miocene of Riversleigh, northwestern Queensland. Alcheringa 23: 31-50.

Brammall, J. and Archer, M., 1997. A new Oligo-Miocene species of Burramys (Marsupialia, Burramyidae) from Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41(2): 247-268.

Carbry, K. 2001. Description and Phylogenetic Analysis of New Species of Ringtail Possums of the Genus Pildra (Marsupialia: Pseudocheiridae) from the Riversleigh World Heritage Property, Queensland. Honours Thesis. University of New South Wales, Sydney.

Cooke, B. N., 1992. Primitive macropodids from Riversleigh, northwestern Queensland. Alcheringa 16: 201-217.

105 CHAPTER 4

Cooke, B. N. 1996. Researches into Fossil Kangaroos and Kangaroo Evolution. PhD Thesis. University of New South Wales, Sydney.

Crosby, K. 2002. Studies in the Diversity and Evolution of Phalangeroid Possums (Marsupialia; Phalangerida; Phalangeroidea). PhD Thesis. University of New South Wales, Sydney.

Davison, C. V. and Young, W. G. 1990. The muscles of mastication of Phascolarctos cinereus (Phascolarctidae; Marsupialia). Australian Journal of Zoology 38: 227-240.

Eisenberg, J. F. 1978. The evolution of arboreal herbivores in the class Mammalia. Pp. 135-151 in The Ecology of Arboreal Folivores. Ed. G. G. Montgomery. Smithsonian Institution Press, Washington D.C.

Flannery, T. F. and Archer, M. 1985. Palorchestes Owen, 1874. Large and small palorchestids. Pp. 234-239 in Kadimakara. Extinct Vertebrates of Australia. Eds P. V. Rich and G. F. van Tets. Pioneer Design Studio, Melbourne.

Flannery, T. F., Archer, M. and Plane, M. D. 1983. Middle Miocene kangaroos (Macropodoidea: Marsupialia) from three localities in northern Australia, with a description of two new subfamilies. Bureau of Mineral Resources Journal of Australian Geology and Geophysics 7: 287-302.

Gillespie, A. 1997. Priscaleo roskellyae sp. nov. (Thylacoleonidae, Marsupialia) from the Oligo-Miocene of Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41: 321-327.

Janis, C. M. 1995. Correlations between craniodental morphology and feeding behavior in ungulates: reciprocal illumination between living and fossil taxa. Pp. 76-98 in Functional Morphology in Vertebrate Paleontology. Ed. J. J. Thomas. Cambridge University Press, New York.

Janis, C. M. 1990. Correlation of cranial and dental variables with dietary preferences in mammals: a comparison of macropodoids and ungulates. Memoirs of the Queensland Museum 28: 349-366.

Johnson, K. A. 1998. Marsupial mole, Notoryctes typhlops (Stirling, 1889). Pp. 409-411 in The Mammals of Australia. Ed. R. Strahan. New Holland Publishers Pty Ltd, Sydney.

Kay, F. R. and Hylander, W.L. 1978. The dental structure of mammalian folivores with a special reference to primates and phalangeroids (Marsupialia). Pp. 173-191 in Ecology

106 CHAPTER 4

of Arboreal Folivores. Ed G. G. Montgomery. Smithsonian Institution Press, Washington D. C.

Lanyon, J. M. and Sanson, G. D. 1986. Koala (Phascolarctos cinereus) dentition and nutrition. I. Morphology and occlusion of cheekteeth. Journal of Zoology London 209: 155-168.

Lauder, G. V. 1995. On the inference of function from structure. Pp 1-18 in Functional Morphology in Vertebrate Paleontology. Ed J. Thomason. Cambridge University Press, Cambridge.

Long, J., Archer, M., Flannery, T. and Hand, S. 2002. Prehistoric Mammals of Australia and New Guinea, One Hundred Million Years of Evolution. University of New South Wales Press Ltd., NSW, Australia.

Mendoza, M., Janis, C. M. and Palmqvist, P. 2002. Characterizing complex craniodental patterns related to feeding behaviour in ungulates: a multivariate approach. Journal of Zoology London 258: 223-246.

Muirhead, J. 1992. A specialised thylacinid, Thylacinus macknessi, (Marsupialia: Thylacinidae) from Miocene deposits of Riversleigh, northwestern Queensland. Australian Mammalogy 15: 67-76.

Muirhead, J. 1994. Systematics, Evolution and Palaeobiology of Recent and Fossil Bandicoots (Peramelemorphia, Marsupialia). PhD Thesis. University of New South Wales, Sydney.

Muirhead, J. 1997. Two new early Miocene thylacines from Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41(2): 367-377.

Musser, A. M. and Archer, M. 1998. New information about the skull and dentary of the Miocene platypus Obdurodon dicksoni, and a discussion of ornithorhynchid relationships. Philosophical Transactions of the Royal Society of London 353: 1063- 1079.

Sanson, G. D. 1982. Evolution of feeding adaptations in fossil and recent macropodids. Pp. 490-506 in The Fossil Vertebrate Record of Australasia, eds P. V. Rich and E. M. Thompson. Monash University Offset Printing Unit, Clayton, Victoria, Australia.

Sanson, G. D. 1991. Predicting the diet of fossil mammals. Pp. 201-227 in Vertebrate Palaeontology of Australasia. Eds P. Vickers-Rich, J. M. Monaghan, R. F. Baird and T. H. Rich. Pioneer Design Studio, Lilydale.

107 CHAPTER 4

Savage, R. J. G. 1977. Evolution of carnivorous mammals. Paleontology 20: 237-271.

Seligsohn, D. and Szalay, F. S. 1978. Relationship between natural selection and dental morphology: tooth function and diet in Lepilemur and Hapalemur. Pp. 289-307 in Development, Function and Evolution of Teeth. Eds P. M. Butler and K. A. Joysey. Academic Press, London.

Van Valen, L. 1969. Evolution of dental growth and adaptation in mammalian carnivores. Evolution 23: 96-117.

Van Valkenburgh, B. 1988. Trophic diversity in past and present guilds of large predatory mammals. Paleobiology 14: 155-173.

Van Valkenburgh, B. 1989. Carnivore dental adaptations and diet: a study of trophic diversity within guilds. Pp. 410-433 in Carnivore Behaviour, Ecology and Evolution. Ed. J. Gittleman. Cornell University Press, New York.

Withers, P. C. 1992. Comparative Animal Physiology. Saunders College Publishing, United States of America.

Woodside, D. P. 1998. Feathertail glider, Acrobates pygmaeus (Shaw, 1794). Pp. 262- 264 in The Mammals of Australia. Ed. R. Strahan. New Holland Publishers Pty Ltd, Sydney.

Wroe, S., 1996. An investigation of phylogeny in the giant extinct rat kangaroo Ekaltadeta (Propleopinae, Potoroidae, Marsupialia). Journal of Paleontology 70 (4): 681-690.

Wroe, S., 1997. Mayigriphus orbus gen. et sp. nov., a Miocene dasyuromorphian from Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41 (2): 439-448.

Wroe, S., 1998. A new ‘bone – cracking’ dasyurid (Marsupialia), from the Miocene of Riversleigh, northwestern Queensland. Alcheringa 22: 277-284.

Wroe, S., 1999. The geologically oldest dasyurid (Marsupialia) from the Miocene of Riversleigh, nothwestern Queensland. Palaeontology 42: 501-527.

108

CHAPTER 5

PALAEOECOLOGY OF RIVERSLEIGH FOSSIL SITES

CHAPTER 5

PALAEOECOLOGY OF RIVERSLEIGH FOSSIL SITES

5.1. Introduction

5.1.1. Definitions

Palaeoecology is the study of the ecological organisation and the associations among species through time (Wing et al. 1992). It involves the identification of palaeocommunities, or collections of species that existed and interacted in one environment, at one time. Each species within a community (or palaeocommunity) has a position among the environmental and biotic components of the habitat relative to other species (this includes spatial distribution, activity time, trophic interactions and locomotor behaviour), referred to as its niche (Whittaker 1975, Lee and Cockburn

1985). Niche describes the partitioning of resources and habitat components between species, as discussed by Vandermeer (1972), Whittaker et al. (1973), Whittaker and

Levin (1975), Hutchinson (1978) and Lee and Cockburn (1985), and refers to a set of environmental parameters and/or functional roles.

The proportional distribution of species among available niches describes resource use by the species within a community in a way that is comparable to other communities.

Such measures, referred to as ‘ecological diversity measures’ (Fleming 1973: 558;

Andrews et al. 1979), are used to characterise or profile a community in terms of adaptive structure (Soligo 2002, Andrews et al. 1979, Reed 1998, Nesbit-Evans et al.

1980, Andrews 1989). Communities may also be described by taxonomic composition, as opposed to adaptive structure. In such cases, species diversity is the basic unit of measure. There are two components to species diversity: variety and relative abundance

109 CHAPTER 5

(Magurran 1988). Species variety, or the number of species, in a defined sampling unit is also called ‘species richness’ and relative abundance refers to the number of individuals within a species.

The use of taxa that are characterised ecologically (e.g., by body size, diet or locomotor behaviour) to describe communities, rather than the use of species diversity measures, has been called ‘taxon-free’ (Damuth 1992) or ‘phylogeny-free’ (Reed 1998) characterisation. This is a highly appropriate tool for palaeoecological analyses because it does not require determination of the number of individuals; only species numbers are necessary. As outlined in Chapter 2, methods for quantification of individuals in fossil samples have inherent assumptions and can be complicated by circumstances peculiar to individual fossil deposits. Additionally, the relative numbers of individuals preserved may not be representative of the living community due to taphonomic processes.

Irrespective of taxonomic composition and relative abundance, ecological diversity analysis reveals the extent of similarity or difference between the ecological adaptive structure of communities and allows comparisons over large geographic space and long time spans (Andrews et al. 1979, Damuth 1992, Reed 1998, Soligo 2002).

Consequently, palaeocommunities may be compared to modern communities from defined habitats or environments in efforts to establish the likely habitat types of the palaeocommunities (Reed 1998, Andrews et al. 1979, Nesbitt-Evans et al. 1981,

Andrews 1989). Fossil mammal communities are particularly good for reconstructing vegetational structure and inferring details about climate, as Janis (1984) has shown with fossil ungulates.

110 CHAPTER 5

Inference of habitat type for palaeocommunities through comparisons to modern communities is validated by the consistency in ecological diversity patterns that exists between modern communities from similar habitats (Andrews et al. 1979). Ecological adaptations of South American, African and Australian mammals from the tropics do not differ greatly, despite the differences in taxonomic composition and histories of the continents (Keast 1969). Convergences occur in a wide range of characteristics, for instance diet, social structure and escape reactions (Dubost 1968, cited in Bourlière

1973). Many authors note parallel trends in patterns of resource use for similar habitat types (for example, Glanz 1982 and Fleming 1973).

5.1.2. Aims

The aims for this component of research were: 1) to determine how representative the

Camel Sputum, Mike’s Menagerie, Quantum Leap, Encore and Ringtail fossil faunas are of the original life assemblages (in terms of the number of individuals preserved per species and the adaptive structure of the community) based on modern analogue faunas and theory as outlined below (Section 5.2.1.); 2) to describe broad habitat groups/types formed through the cluster analysis of modern faunas on the basis of ecological adaptive structure in diet and locomotion; 3) to characterise Riversleigh fossil communities using ecological diversity measures (trophic and locomotor); 4) through multiple discriminant analysis, to identify the modern broad habitat types that Riversleigh fossil faunas best fit based on community structure; 5) to explore relationships between aspects of mammalian adaptive structure within modern communities and annual rainfall in an attempt to propose a possible annual rainfall range for the Riversleigh palaeohabitats.

111 CHAPTER 5

5.1.3. Background

Material collected and processed from the fossil sites of Riversleigh over the last thirty or so years (Archer et al. 1996) has been sorted and examined morphologically, functionally, taxonomically and phylogenetically. Ecological aspects of the fossil assemblages have been investigated (Woodburne et al. 1985; Archer et al. 1989, 1995,

1997; Archer 1992) on a broad level and in terms of presence and diversity of animal

‘indicator species’. These are species considered to have particular habitat requirements, based on modern representatives, such as folivorous possums and frogs. Also, palaeoecology has been considered for isolated groups of animals at Riversleigh

(Muirhead 1994; Hand 1997 and 2001; and Crosby 2002, for bandicoots, bats and possums, respectively). The first extensive, quantitative palaeocommunity analyses of

Riversleigh material were carried out by Myers (2002). He characterised a number of

Riversleigh faunal assemblages on the basis of taxonomic representation and body mass characteristics and compared them to modern communities from a variety of Australian habitat types in an attempt to describe the likely habitats of the Riversleigh assemblages.

He also grouped discrete fossil assemblages together as palaeocommunities based on shared taxa.

Some points about the analyses of Myers (2002) that are relevant to the present study should be emphasised. These are: habitats of the modern faunas were defined a priori, only Australian faunas were used as modern analogues, and consequently the habitat descriptions for these faunas were very detailed. Myers found that some of the fossil assemblages did not match up closely to any of the modern habitats. This may be either because the fossil assemblages are from less homogenous habitats than the modern

112 CHAPTER 5

faunas used for comparison, or because the fossil communities are from habitats unlike any of the modern analogues used.

In the current study, modern analogues include communities from North, Central and

South America, Africa and New Guinea, in addition to Australia. It is anticipated that the broader habitat descriptions required for this array of sites will be more appropriate for the inclusion of fossil communities. Additionally, the modern habitat types are not described a priori on the basis of vegetation, but instead are defined by the grouping of mammalian communities by cluster and principal co-ordinates analyses. This ensures that the habitat groups are directly associated with the ecological diversity measures used to define the structure of communities as opposed to other descriptive factors.

Descriptions and definitions of habitats in literature vary widely and are not necessarily comparable. They can be based on any number of physical or biotic factors, for instance, elevation, topographic position, floristics, vegetation cover and climate.

Habitat groups formed for the modern communities in the cluster analysis are subsequently described, taking into consideration the features of vegetation structure consistent between the constituent localities.

5.2. Methods

5.2.1. Consideration of potential bias in fossil community representation

To a large extent the fossil sample and data derived from it determined the types of questions that were addressed in this palaeoecological study. Fossil assemblages need to be assessed in terms of their relationship to original species associations prior to making comparisons between extinct and living communities (Wing et al. 1992). The data in this study (specimen counts, species lists for the fossil sites and

113 CHAPTER 5

morphological/functional attributes of these species) were explored to establish whether fossil species relative abundance and fossil community profiles may potentially be biased and thus not representative of past communities. The methods used are those outlined by Damuth (1982) and Soligo (2002) for relative abundance of individuals and community structure respectively.

Damuth’s (1982) method involves plotting log body mass against log abundance of primary consumers. For an unbiased assemblage the gradient for this plot was found to be –1.05 (with a range of –0.80 to –1.30). The slope of this regression for a fossil assemblage specifies the magnitude and direction of bias that needs to be explained or corrected for (Damuth 1982). The expected slope of this regression “…represents an ecological energetic regularity of community structure that we would expect on uniformitarian grounds to be in effect in virtually all ancient communities as it is in modern ones.” (Damuth 1982: 436). This method is based on the allometric relationship of body mass to population density and turnover rate for a species (Damuth 1982 and references therein: 436). Advantages of this method of assessing bias, as identified by

Damuth, include no presumed method of estimating abundance and not all species having to be included for a valid result. The second point is important for Riversleigh sites because in many cases, the full sample has not been examined as collected material remains unprocessed.

Soligo’s (2002) method of assessing the extent to which adaptive structure, as represented by the species present, is representative of a community does not involve counting individuals. The adaptive structure, or the pattern of distribution of fossil species among ecological diversity categories, is used. Modern communities form the

114 CHAPTER 5

comparative sample, setting the limits of variability for each category of ecological diversity. The ecological diversity categories are a combination of trophic level and body mass (listed in Table 5.2.1). This method is based on principles of prey to predator ratios, species numbers in relation to body size, and patterns of distribution of small

Site Family Genus/species/specimen ID Body mass (g) Trophic/weight Specimens Encore Dasyuridae Ganbulanyi djadjinguli 3600 BS 2 Thylacinidae Thylacinus sp. cf. T. macknessi 9017 BS 1 Insertae sedis Mayigriphus orbus 17.5 AS 2 ? Yaraloidea, gen. and sp. indet ? Phascolarctidae Litokoala kanunkaensis 2213 BP Phascolarctos sp. 5008 BP Vombatidae cf. Warenja sp. 23733 CP 47 Thylacoleonidae Wakaleo vanderleurei 45 000-100 000 CS 8 Diprotodontidae Neohelos sp. ? DP? Palorchestidae Palorchestes anulus 107399 DP 1 Palorchestes sp. ? DP? Phalangeridae Trichosurus [euphrasia] 2590 BP 5 Burramyidae Burramys brutyi 21 AP 1 Pseudocheiridae cf. Pseudocheirops sp. ? AP? 1 Pildra sp. 2 385 AP 2 Macropodidae Gangaroo new sp. ? BP? Wanbaroo hilarus 4345 BP 1 Wanbaroo [wulugu] 6823 BP 1 Sthenurinae cf. Hadronomas sp. ? Rhyzosthenurus flanneryi 19481.5 CP 1 Hypsiprymnodont- Ekaltadeta ima 4805 BP idae Ekaltadeta jamiemulvaneyi 7241 BP 2 Petauroid QMF 23785 113 AP 1 Dasyurid QMF 20929 267 AS 1

Mike's Yalkaparidontidae Yalkaparidon 262 AS 1 Menagerie Notoryctidae ? 160 AS Thylacinidae Thylacinus macknessi 9017 BS 1 ? Yaraloidea, gen. and sp. indet ? AS? Wynyardiidae ? ? BP? Diprotodontidae Neohelos sp. ? DP? Neohelos tirarensis 128 883 DP 1 Palorchestidae Palorchestes ? DP? Phalangeridae Wyulda asherjoeli 865 AP 2 [Illungalya aletes] 1181 BP 1 Pilkipildridae ? ? AP? Acrobatidae ? ? AP? Insertae sedis Djaludjangi yadjana 273 AP 1 Macropodidae Gangaroo bilamina 1531 BP 1 Wabularoo naughtoni 8933 BP 2 Hypsiprymnodont- Hypsiprymnodon (AR 15134) 356 AP idae Ekaltadeta ima 4805 BP 1 Table 5.2.1. (Continued next page…) Trophic level and body mass categories for taxa from the six study sites (sensu Soligo 2002). Body mass is calculated using the equations ofMyers (2001); question marks indicate where appropriate specimens were not available for the calculation of body mass; 'specimens' refers to the number of identified specimens for each taxon, where this information is available. Abbreviations: AP = primary consumers, < 1 kg; AS = secondary consumers, < 1 kg; BP = primary consumers, 1-10 kg; BS = secondary consumers, 1-10 kg; CP = primary consumers, 10-45 kg; CS = secondary consumers, 10-45 kg; DP = primary consumers, >45 kg.

115 CHAPTER 5

Site Family Genus/species/specimen ID Body mass (g) Trophic/weight Specimens Camel Sputum Yalkaparidontidae Yalkaparidon coheni 262 AS 33 Notoryctidae ? 160 AS Thylacinidae Ngamalacinus timmulvaneyi 5743 BS 1 Wabulacinus ridei 5340 BS 2 Yaralidae Yarala burchfieldi 53 AS 9 Insertae sedis [Bulunga palara] 76 AS 11 (Peramelemorphians) [Galadi grandis] 1061 BP 1 [Gunawidgi tubus] 170.5 AP 46 [Madju ignotae] 380 AP 2 [Madju variae] 627 AP 16 Wynyardiidae Namilamadeta ? ? Phascolarctidae Nimiokoala greystanesi 2714 BP 4 Nimiokoala [whitworthi] 13436 CP 3 Priscakoala [aureco] 7770 BP 1 Diprotodontidae Neohelos sp. ? DP? Neohelos tirarensis 128 883 DP 1 Palorchestidae Propalorchestes ponticulus 94980 DP 1 Palorchestes ? DP? Phalangeridae [Ilungalya aletes] 1181 BP 5 [Ilungalya lygaeus] 1249 BP 2 [Onirocuscus inversus] 1863 BP 2 [Onirocuscus silvacultrix] 1903 BP 1 Wyulda asherjoeli 865 AP 1 Pilkipildridae ? AP? Acrobatidae ? AP? Burramyidae Burramys brutyi 21 AP 10 Pseudocheiridae Marlu cf. Kutjamarpensis 504 AP 1 Paljara maxbourkei 325 AP 1 Paljara nancyhawardae 160 AP 3 Pildra sp. 2 385 AP 1 Insertae sedis Djaludjangi yadjana 273 AP 8 Macropodidae Bulungamaya delicata 1355 BP 16 Gangaroo bilamina 1531 BP 4 Nowidgee matrix 1462 BP 7 Wabularoo naughtoni 8933 BP 19 Potoroidae ? ? ? Hypsiprymnodont- Hypsiprymnodon (AR 10704) 300 AP idae Ekaltadeta ima 4805 BP 5 Balbaridae Balbaroo gregoriensis 6039 BP 1 Ganawamaya acris 5852 BP 1 Nambaroo [camilleriae] 6018.5 BP 1 Wururoo [gadiyuli] 9527 BP 2

Table 5.2.1. (Continued…) Trophic level and body mass categories for taxa from the six study sites (sensu Soligo 2002). Abbreviations: AP = primary consumers, < 1 kg; AS = secondary consumers, < 1 kg; BP = primary consumers, 1-10 kg; BS = secondary consumers, 1-10 kg; CP = primary consumers, 10- 45 kg; CS = secondary consumers, 10-45 kg; DP = primary consumers, >45 kg.

116 CHAPTER 5

Site Family Genus/species/specimen ID Body mass (g) Trophic/weight Specimens Quantum Yalkaparidontidae Yalkaparidon 262 AS 4 Leap Notoryctidae QMF 50799 160 AS 1 Dasyurid QMF 30763 141 AS 1 ? Yaraloidea, QMF 50793 103 AS 1 Yaraloidea, QMF 50803 810 AP 1 Yaraloidea, QMF 50883 555 AP 1 Yaraloidea, QMF 50800 88 AS 1 Yaraloidea, QMF 50792 130 AS 1 Diprotodontidae ? ? DP? 2 Phalangeridae [Illungalya aletes] 1181 BP 1 Pilkipildridae ? ? AP? 1 Petauridae QMF 41090 67 AP 3 Macropodidae Bulungamaya delicata 1355 BP 1 Hypsiprymnodontidae Hypsiprymnodon (QMF 41071) 654 AP 1 Balbaridae Nambaroo sp. 3 7820 BP 1

Ringtail Notoryctidae ? 55 AS Thylacinidae Maximucinus muirheadae 18400 CS Nimbacinus new sp. 5023 BS 1 Insertae sedis [Gunawidgi tubus] 200 AP 1 Diprotodontidae Nimbadon lavarackorum 52000 DP Phalangeridae ? ? ? Acrobatidae ? ? AP Burramyidae Burramys brutyi 21 AP 7 Cercatetus ? AP Petauridae ? ? AP Pseudocheiridae Pseudochirops ? AP Marlu cf. kutjamarpensis 504 AP 3 Ornithorhynchidae Obdurodon dicksoni ? ? 19 Macropodoid AR 18501 8666 BP 1 Macropodoid QMF 41247 505 AP 1

species over functional groups; these principles indicate that “…there must be limits to the variability with which species are distributed over the functional groups present at a given locality” (Soligo 2002: 25). Soligo does not describe the trophic modes that he includes within the ‘primary’ and ‘secondary’ categories. As a result, these categories may not be directly equivalent to those presented here. In this study, primary consumers include browsers, frugivores, root and fungi eaters, pollen eaters, exudivores, nectarivores, grazers and omnivores. Omnivores are included because it is often difficult to determine the relative proportions of plant and animal matter in the diet,

117 CHAPTER 5

especially for fossil omnivores. Secondary consumers (insectivores and carnivores) over

45kg in body mass are not found in modern or Oligo-Miocene faunas of Australia, and are therefore excluded here. The comparative data set for this analysis consisted of twenty-nine modern Australian faunas from localities listed in Table 5.2.2 and illustrated in Figure 5.2.1.

5.2.2. Modern communities: localities and habitats

The fauna lists for the modern mammalian communities used in the comparative data set (84 in total) for the palaeoecological analyses were obtained from a number of sources (Table 5.2.2; species lists in Appendices C1 – C5). Mammal lists for North,

Central and South American sites were obtained from Fleming (1973), Ojeda and Mares

(1989), and Medellin (1994). Reed (1998) provides counts of mammal species in ecological diversity categories for African sites, though actual species lists for the sites are not provided. The habitats of these mammalian communities are described by the respective authors, but in many cases it is difficult to obtain a measure of habitat heterogeneity from the descriptions.

118 CHAPTER 5

Site Family Genus/species/specimen ID Body mass (g) Trophic/weight Specimens Site Family Genus/species/specimen ID Body mass (g) Trophic/weight Specimens Dasyuridae Ganbulanyi djadjinguli 3600 BS 2 Yalkaparidontidae Yalkaparidon coheni 262 AS 33 Thylacinidae Thylacinus sp. cf. T. macknessi 9017 BS 1 Notoryctidae ? 160 AS Insertae sedis Mayigriphus orbus 17.5 AS 2 Ngamalacinus timmulvaneyi 5743 BS 1 Thylacinidae ? Yaraloidea, gen. and sp. indet ? Wabulacinus ridei 5340 BS 2 Litokoala kanunkaensis 2213 BP Yaralidae Yarala burchfieldi 53 AS 9 Phascolarctidae Phascolarctos sp. measure at uni BP Insertae sedis [Bulunga palara] 76 AS 11 Vombatidae cf. Warenja sp. 23733 CP 47 [Galadi grandis] 1061 BP 1 Thylacoleonidae Wakaleo vanderleurei 45 000-100 000 CS 8 [Gunawidgi tubus] 170.5 AP 46 (Peramelemorphians) Diprotodontidae Neohelos sp. ? DP? [Madju ignotae] 380 AP 2 Palorchestes anulus 107399 DP 1 [Madju variae] 627 AP 16 Palorchestidae Palorchestes sp. ? DP? Wynyardiidae Namilamadeta ?? Phalangeridae Trichosurus [euphrasia] 2590 BP 5 Nimiokoala greystanesi 2714 BP 4 Burramyidae Burramys brutyi 21 AP 1 Phascolarctidae Nimiokoala [whitworthi] 13436 CP 3 Encore cf. Pseudocheirops sp. ? AP? 1 Priscakoala [aureco] 7770 BP 1 Pseudocheiridae Pildra sp. 2 385 AP 2 Neohelos sp. ? DP? Diprotodontidae Gangaroo new sp. ? BP? Neohelos tirarensis 128 883 DP 1 Macropodidae Wanbaroo hilarus 4345 BP 1 Propalorchestes ponticulus 94980 DP 1 Palorchestidae Wanbaroo [wulugu] 6823 BP 1 Palorchestes ? DP? cf. Hadronomas sp. ? [Ilungalya aletes] 1181 BP 5 Sthenurinae Rhyzosthenurus flanneryi 19481.5 CP 1 [Ilungalya lygaeus] 1249 BP 2 Ekaltadeta ima 4805 BP Phalangeridae [Onirocuscus inversus] 1863 BP 2 Hypsiprymnodontidae Ekaltadeta jamiemulvaneyi 7241 BP 2 [Onirocuscus silvacultrix] 1903 BP 1 Petauroid QMF 23785 113 AP 1 Wyulda asherjoeli 865 AP 1

Dasyurid QMF 20929 267 AS 1Camel Sputum Pilkipildridae ? AP? Acrobatidae ? AP? Yalkaparidontidae Yalkaparidon 262 AS 1 Burramyidae Burramys brutyi 21 AP 10 Notoryctidae ? 160 AS Marlu cf. Kutjamarpensis 504 AP 1 Thylacinidae Thylacinus macknessi 9017 BS 1 Paljara maxbourkei 325 AP 1 Pseudocheiridae ? Yaraloidea, gen. and sp. indet ? AS? Paljara nancyhawardae 160 AP 3 Wynyardiidae ? ? BP? Pildra sp. 2 385 AP 1 Neohelos sp. ? DP? Insertae sedis Djaludjangi yadjana 273 AP 8 Diprotodontidae Neohelos tirarensis 128 883 DP 1 Bulungamaya delicata 1355 BP 16 Palorchestidae Palorchestes ? DP? Gangaroo bilamina 1531 BP 4 Macropodidae Wyulda asherjoeli 865 AP 2 Nowidgee matrix 1462 BP 7 Phalangeridae [Illungalya aletes] 1181 BP 1 Wabularoo naughtoni 8933 BP 19 Pilkipildridae ? ? AP? Potoroidae ? ??

Mike's Menagerie Acrobatidae ? ? AP? Hypsiprymnodon (AR 10704) 300 AP Hypsiprymnodontidae Insertae sedis Djaludjangi yadjana 273 AP 1 Ekaltadeta ima 4805 BP 5 Gangaroo bilamina 1531 BP 1 Balbaroo gregoriensis 6039 BP 1 Macropodidae Wabularoo naughtoni 8933 BP 2 Ganawamaya acris 5852 BP 1 Balbaridae Hypsiprymnodon (AR 15134) 356 AP Nambaroo [camilleriae] 6018.5 BP 1 Hypsiprymnodontidae Ekaltadeta ima 4805 BP 1 Wururoo [gadiyuli] 9527 BP 2

Yalkaparidontidae Yalkaparidon 262 AS 4 Notoryctidae ? 55 AS Notoryctidae QMF 50799 160 AS 1 Maximucinus muirheadae 18400 CS Thylacinidae Dasyurid QMF 30763 141 AS 1 Nimbacinus new sp. 5023 BS 1 Yaraloidea, QMF 50793 103 AS 1 Insertae sedis [Gunawidgi tubus] 200 AP 1 Yaraloidea, QMF 50803 810 AP 1 Diprotodontidae Nimbadon lavarackorum 52000 DP ? Yaraloidea, QMF 50883 555 AP 1 Phalangeridae ? ? ? Yaraloidea, QMF 50800 88 AS 1 Acrobatidae ? ? AP Yaraloidea, QMF 50792 130 AS 1 Burramys brutyi 21 AP 7 Burramyidae

Diprotodontidae ? ? DP? 2 Ringtail Cercatetus ?AP Phalangeridae [Illungalya aletes] 1181 BP 1 Petauridae ? ? AP Quantum Leap Pilkipildridae ? ? AP? 1 Pseudochirops ?AP Pseudocheiridae Petauridae QMF 41090 67 AP 3 Marlu cf. kutjamarpensis 504 AP 3 Macropodidae Bulungamaya delicata 1355 BP 1 Ornithorhynchidae Obdurodon dicksoni ?? 19 Hypsiprymnodontidae Hypsiprymnodon (QMF 41071) 654 AP 1 Macropodoid AR 18501 8666 BP 1 Balbaridae Nambaroo sp. 3 7820 BP 1 Macropodoid QMF 41247 505 AP 1

Table 5.2.1. Trophic level and body mass categories for taxa from the six study sites (sensu Soligo 2002). Body mass is calculated using the equations of Myers (2001); question marks indicate where appropriate specimens were not available for the calculation of body mass; 'specimens' refers to the number of identified specimens for each taxon, where this information is available. Abbreviations: AP = primary consumers, < 1 kg; AS = secondary consumers, < 1 kg; BP = primary consumers, 1-10 kg; BS = secondary consumers, 1-10 kg; CP = primary consumers, 10-45 kg; CS = secondary consumers, 10-45 kg; DP = primary consumers, >45 kg.

119 CHAPTER 5

15O

NORTHERN TERRITORY

WESTERN QUEENSLAND AUSTRALIA

SOUTH AUSTRALIA

NSW

35O VICTORIA

1000 km

TASMANIA O 125O 140

Figure 5.2.1. Map of locations of modern Australian sites used in palaeoecological analyses. The Grampians and Gippsland regions of Victoria include four and ten habitats respectively (as described in Table 5.2.2).

Lawn Hill National Park Grampians region, Victoria Central Australia Gippsland region, Victoria Ku-Ring-Gai Chase National Park Jervis Bay National Park Lamington National Park South Coast TasWS TasH TasS TasR Cape York Atherton Uplands Mossman Lowlands Ingham Lowlands Lee Uplands

120 CHAPTER 5

For Australian localities (Fig. 5.2.1), an attempt was made to incorporate information about the extent of habitat heterogeneity. The purpose of this was to test the sensitivity of the analyses in distinguishing mosaic habitats from uniform habitats. Mosaic, or heterogeneous, habitats consist of more than one vegetation type. For example, the present day environment at Riversleigh, as represented at Lawn Hill (Figure 5.2.2), is strikingly heterogeneous with a tall, shaded gallery forest along the river edge that abruptly changes to woodland and grassland within metres of the river (O’Keefe 1996).

Distinction between mosaic and uniform habitats for the modern sites may be useful in determining the extent of habitat heterogeneity for fossil sites.

Figure 5.2.2. Lawn Hill Gorge and surrounding landscape.

National parks with mosaic habitat types were chosen to represent heterogenous habitats in Australia. These include Jervis Bay National Park, Ku-Ring-Gai Chase National

Park, Lamington National Park and Lawn Hill National Park. Additional sites with

121 CHAPTER 5

habitat heterogeneity were included: South Coast (a strip, approximately 150 km, of coastal vegetation from Batemen’s Bay in the north to Eden in the south), Cape York, and central Australia. Localities with uniform habitat types, in Tasmania and northern

Queensland (Atherton uplands, Mossman lowlands, Ingham lowlands and Lee uplands) were also identified. Mammal lists for Australian sites, except those in Victoria, are taken from references as indicated in Table 5.2.2.

Mammal lists for Australian sites in Victoria and for New Guinea were prepared through the use of range maps of mammals and vegetation. The purpose of this was to obtain community data for relatively homogenous habitat types in terms of vegetation.

For Victoria, two bioregions were selected, East Gippsland and the Grampians, with ten and four habitat types identified respectively. The habitat types were determined from a combination of vegetation type descriptions (Conn 1993, Menkhorst ed. 1995 and

Anon. 1997a); listed in Table 5.2.3 are the vegetation types and the resulting ‘adjusted habitat type’ used in this study. Three references were used for habitat type definition because the mammal distribution descriptions, with respect to vegetation, are not standardised – some use vegetation categories, others list plant species that particular mammals are normally associated with. Distribution maps of the mammals and descriptions of their preferred habitat (Menkhorst ed. 1995) were used in combination with a map of broad vegetation types for Victorian bioregions (Anon. 1997a) to produce the lists of mammals for each habitat type identified in these bioregions. East

Gippsland (12 820 km2 in area; Conn 1993) was chosen because it is the bioregion least disturbed by human activity (Anon. 1997b). The Grampians covers an area of 2 356 km2

(Conn 1993), surrounded by agricultural land, thus representing a relatively isolated

122 CHAPTER 5

natural region (though sites within the region are adjacent, allowing exchange of fauna between sites).

Alliance equivalent Broad vegetation Conn’s (1993) Adjusted Habitat type Bioregion of Menkhorst, ed. type equivalent category habitat types abbreviation (1995) of Anon. (1997) primary dune coast scrubs (wattle coastal scrubs and coastal scrubs, scrub or banksia) grassland grasslands and VGipSGH saltmarsh saltmarsh herbland complex herblands coastal coastal wet heaths ? VGipCH heathland heathland Banksia heathy wet heathy heathy woodland VGipHW woodland woodland woodland rainshadow rainshadow rainshadow dry woodland VGipRW woodland woodland woodland lowland lowland and dry sclerophyll dry forest foothill forest dry lowland forest complex VGipDLF forests dry sclerophyll dry forest (Box- dry foothill forest forest Ironbark forests?) complex montane montane dry montane sclerophyll dry forest woodland woodland and VGipMW woodland and complex forest montane forest

East Gippsland Gippsland East coastal riparian forest sclerophyll wet (gully?) forest riparian (in river VGipRF forest gullies) riparian forest wet forest lowland and moist wet sclerophyll wet lowland wet (gully?) forest foothill forest VGipWLF forest forests complex warm warm temperate moist foothill lowland rainforest temperate/low VGipWTR rainforest forest complex land rainforest montane moist cool cool temperate forest complex upland rainforest temperate/upl VGipCTR rainforest and moist foothill and rainforest forest complex swamplands and wet heaths heathlands herb-rich herb-rich heath woodland/ woodland VGrHRW heathy wet woodland sclerophyll complexes woodlands woodland plains grassy grassy grassy dry woodland woodland VGrGW woodland woodland complexes low open- woodlands/ shrublands

Grampians shrublands inland slopes woodland/ rocky outcrop woodland VGrWDF ? dry-forest woodland complexes forest (open, dry forests dry) wet sclerophyll dry foothill forest wet forests open-forest VGrOF forest complexes

Table 5.2.3. Vegetation types of the East Gippsland and Grampians Regions of Victoria as described by Conn (1993), Anonymous (1997a) and Menkhorst (ed. 1995), and ‘adjusted habitats types’ derived from these descriptions.

123 CHAPTER 5

The mammal list for New Guinea was obtained from mammal distribution maps in

Flannery (1995) and is restricted to a specific habitat type – medium-crowned lowland hill forest vegetation (Paijmans 1975). This vegetation occurs on hills and mountains below 1 400 m elevation and is floristically diverse with crown sizes in the canopy from

8 to 15 m in diameter, crown height between 25 and 30 m and 60-80% crown closure

(Paijmans 1975). Aspects of geology were also taken into consideration in the collation of this list. A geological map (D’Addario et al. 1975) was used in conjunction with the vegetation map (Paijmans 1975) and mammal distribution maps (Flannery 1995).

Distributions were restricted to limestone terrain.

5.2.3.Ecological diversity analyses

5.2.3.1. Variables used

Taxa from all modern sites and from Camel Sputum, Mike’s Menagerie, Quantum

Leap, Encore and Ringtail Sites were placed into categories of trophic and locomotor behaviour (Appendices C1 – C5 for mammal species lists from modern localities;

Appendices C6. 1 – C6. 4 for trophic and locomotor adaptations of modern species;

Table 3.3.6 in Chapter 3 for locomotor adaptations of fossil taxa; Tables 4.3.2 – 4.3.7 in

Chapter 4 for diet adaptations of fossil taxa). The categories used reflect the extent of information obtainable from fossil morphology, as discussed in Chapters 3 and 4. They represent functional groups and are the variables used in further analyses of the communities. Taxonomic uniformitarianism and functional and ecological morphology

(ecomorphology) methods can be used in obtaining ecological diversity information for fossil animals (Reed 1998). The taxonomic uniformitarian approach, as described by

Reed (1998), compares a fossil taxon with its closest living relative to predict ecological behaviour. Ecomorphology is comparative functional morphology; it involves

124 CHAPTER 5

morphological analyses of extant taxa as a basis for comparison to fossil animals to which ecological behaviour is ascribed (Karr and James 1975; Damuth 1992; Reed

1998). The use of functional morphology in predicting ecological adaptations of fossil taxa is preferable to the use of taxonomic analogy (Reed 1998). Ecomorpholoy has been applied in the current study for the determination of locomotor behaviour (Chapter 3), and a combination of ecomorphology and taxonomic analogy has been used for the determination of diets of fossil taxa (Chapter 4).

Each mammal community (modern and fossil) is characterised by the percentage of species in each of the categories for diet and locomotion (Tables 5.2.4 and 5.2.5).

Vegetation is the source of food and shelter for animals (Andrews and O’Brien 2000), thus trophic and locomotor adaptations were considered appropriate in this study for the formation of habitat groupings that reflect general vegetation structure.

5.2.3.2. Mammal taxa

Exotic species and humans were excluded from species lists for modern communities.

Chiropteran species were not included for the analysis of Camel Sputum, Mike’s

Menageris, Quantum Leap, Encore and Ringtail Sites. Despite the fact that bats enhance habitat distinctions in palaeoecological studies (Andrews and O’Brien 2000), it can not be assumed that bat specimens from these sites are representative of the bat communities. In addition to this, the bats from these sites have been identified to genus level at best. The most reliable Riversleigh fossil fauna for bat community analyses is the Bitesantennary Site local fauna (see Chapter 2). The absence of bats from communities used in palaeoecological analyses is unlikely to alter the interpretation of

125 CHAPTER 5

Site Arboreal Scansorial SmallMix Terrestrial Hopper Other Terr+hop TotalLoc V.GipSGH 7.69 0 38.46 38.46 15.38 0 53.84 13 V.GipCH 5.26 036.84 26.31 21.05 10.53 47.36 19 V.GipHW 9.09 4.54 36.36 22.73 18.18 9.09 40.91 22 V.GipRW 22.22 11.11 22.22 22.22 22.22 0 44.44 18 V.GipDLF 14.81 11.11 33.33 18.52 14.81 7.41 33.33 27 V.GipMW 16.67 16.67 27.78 22.22 16.67 0 38.89 18 V.GipRF 13.64 13.64 31.82 22.73 13.64 4.54 36.37 22 V.GipWLF 14.81 11.11 29.63 22.22 14.81 7.41 37.03 27 V.GipWTF 14.29 14.29 33.33 28.57 4.76 4.76 33.33 21 V.GipCTR 19.05 9.52 28.57 23.81 14.29 4.76 38.1 21 V.GrHRW 11.76 5.88 47.06 11.76 23.53 0 35.29 17 V.GrGW 0 020 20 60 0 80 5 V.GrWDF 8.69 4.35 47.83 8.69 26.09 4.35 34.78 23 V.GrOF 10 5 45 5 25 10 30 20 Jervis 23.53 5.88 35.29 17.65 17.65 0 35.3 17 Kuringai 14.28 9.52 28.57 23.81 14.28 9.52 38.09 21 SouthCoast 16.67 8.33 36.11 13.89 19.44 5.56 33.33 36 CapeYork 11.63 11.63 37.21 13.95 20.93 4.65 34.88 43 Lamington 10.53 7.89 42.1 10.53 26.32 2.63 36.85 38 Mossman 10.71 14.28 42.86 10.71 14.28 7.14 24.99 28 Atherton 19.15 10.64 46.81 6.38 14.89 2.13 21.27 47 Ingham 7.41 11.11 51.85 11.11 14.81 3.7 25.92 27 LeeUp 11.11 13.89 36.11 11.11 25 2.78 36.11 36 LawnHill 0 11.76 35.29 11.76 35.29 5.88 47.05 17 TasWS 9.09 18.18 27.27 18.18 18.18 9.09 36.36 11 TasH 7.69 23.08 15.38 23.08 23.08 7.69 46.16 13 TasS 5.88 23.53 29.41 17.65 17.65 5.88 35.3 17 TasR 6.25 18.75 37.5 12.5 12.5 12.5 25 16 CentAust 0 0 75 12.5 12.5 0 25 8 NewGuinea 21.43 11.9 50 14.29 0 2.34 14.29 39 Tundra 0 050 43.75 0 6.25 43.75 16 Wsp 0 21.43 35.71 35.71 0 7.14 35.71 14 Vhf 0 27.59 37.93 31.03 0 3.45 31.03 29 Dbm 0 21.43 35.71 39.29 0 3.57 39.29 28 Dtf 17.5 2035 22.5 0 2.5 22.5 38 MTf 19.51 19.51 26.83 21.95 0 9.76 21.95 37 Puna 0 5.56 50 44.44 0 0 44.44 19 Prepuna 0 16.67 50 33.33 0 0 33.33 6 Monte 7.69 15.38 46.15 30.77 0 0 30.77 14 UppMontane 0 44.44 33.33 22.22 0 0 22.22 10 LowMontane 8 20 40 24 0 8 24 27 Transitional 11.11 15.56 31.11 37.78 0 4.44 37.78 43 MontChaco 5.26 15.79 31.58 47.37 0 0 47.37 19 Chaco 2.78 11.11 27.78 55.56 0 2.78 55.56 36 Chamela 0 21.88 43.75 34.37 0 3.12 34.37 32 MtAzules 11.11 22.22 26.67 35.56 0 4.44 35.56 45 LaSelva 21.57 17.65 27.45 29.41 0 3.92 29.41 51 Barro 25.64 17.95 23.08 25.64 0 7.69 25.64 39 Guatapo 13.16 15.79 36.84 31.58 0 2.63 31.58 38 Masaguaral 10.71 21.43 25 39.29 0 3.57 39.29 28 Cuzco 23.61 15.23 29.17 25 0 6.94 25 72 Manu 25.4 19.05 28.57 22.22 0 4.76 22.22 63 Belem 17.74 20.97 24.19 29.03 0 8.06 29.03 62 Camel Sputum 12 4 40 16 28 0 44 25 Mike's Menagerie 0 0 37.5 25 37.5 0 62.5 8 Quantum Leap 12.5 0 50 25 12.5 0 37.5 8 Encore 0 6.25 25 12.5 56.25 0 68.75 16 Table 5.2.4. Percentage of taxa in ecological diversity categories for locomotion for modern and fossil localities. ‘Terr+Hop’ is the sum of terrestrial and hopper categories; ‘Total Loc’ is not a percentage; it is the total number of taxa included for the localities.

126 CHAPTER 5

Site Root/fungi Granivore Grazer Brows/fruit Omnivore Insectivore Carnivore Total diet V.GipSGH 0 015.38 7.69 30.77 38.46 7.69 13 V.GipCH 0 5.26 15.79 10.53 21.05 31.58 15.79 19 V.GipHW 0 4.55 13.64 13.64 22.73 27.27 18.18 22 V.GipRW 0 022.22 27.78 22.22 11.11 16.67 18 V.GipDLF 3.7 011.11 22.22 25.93 18.52 18.52 27 V.GipMW 0 022.22 22.22 27.78 16.67 11.11 18 V.GipRF 0 013.64 22.73 31.82 18.18 13.64 22 V.GipWLF 3.7 014.81 22.22 22.22 22.22 14.81 27 V.GipWTF 4.76 0 4.76 28.57 19.05 28.59 14.29 21 V.GipCTR 4.76 0 14.29 23.81 23.81 19.05 14.29 21 V.GrHRW 0 11.76 17.65 23.53 23.53 23.53 0 17 V.GrGW 0 060 0 20 20 0 5 V.GrWDF 0 8.7 17.39 17.39 26.09 21.74 8.7 23 V.GrOF 0 1020 20 20 15 15 20 Jervis 0 011.76 23.53 29.41 29.41 5.88 17 Kuringai 0 014.29 19.05 28.57 19.05 19.01 21 SouthCoast 5.56 5.56 16.67 16.67 19.44 22.22 13.89 36 CapeYork 0 2.33 18.6 23.26 20.93 18.6 16.28 43 Lamington 2.56 2.56 17.95 20.51 23.08 17.95 15.38 38 Mossman 0 03.57 28.57 32.14 14.28 21.43 28 Atherton 2.13 4.25 6.38 29.79 27.66 14.89 14.89 47 Ingham 3.7 3.7 3.7 22.22 33.33 22.22 11.11 27 LeeUp 2.78 011.11 27.78 30.56 13.89 13.89 36 LawnHill 0 5.88 41.18 23.53 0 11.76 17.65 17 TasWS 0 027.27 18.18 27.27 9.09 18.18 11 TasH 0 023.08 15.38 15.38 15.38 30.77 13 TasS 5.88 017.65 11.76 17.65 17.65 29.41 17 TasR 0 018.75 12.5 25 18.75 25 16 CentAust 0 12.5 12.5 0 25 37.5 12.5 8 NewGuinea 0 5.13 7.69 35.9 23.08 23.08 5.13 39 Tundra 0 043.75 0 18.75 12.5 25 16 Wsp 0 021.43 7.14 28.57 7.14 35.71 14 Vhf 0 013.79 6.9 41.38 13.79 31.03 29 Dbm 0 017.86 7.14 32.14 10.71 32.14 28 Dtf 2.63 2.63 5.26 21.05 42.11 10.53 15.79 38 MTf 2.7 5.41 2.7 24.32 37.84 8.11 18.92 37 Puna 0 5.56 50 5.56 5.56 22.22 11.11 18 Prepuna 0 33.33 33.33 0 0 0 33.33 6 Monte 0 14.28 14.28 7.14 7.14 28.57 28.57 14 UppMontane 0 10 10 0 10 20 50 10 LowMontane 0 3.7 11.11 18.52 22.22 14.81 29.63 27 Transitional 0 2.32 11.63 13.95 18.6 32.56 27.91 43 MontChaco 0 5.26 21.05 0 10.52 31.58 31.58 19 Chaco 0 2.78 27.78 5.56 11.11 30.56 22.22 36 Chamela 0 18.18 15.15 6.06 18.18 12.12 30.3 33 MtAzules 0 19.56 8.69 21.74 19.56 15.22 17.39 46 LaSelva 1.96 11.76 7.84 25.49 21.57 13.72 17.65 51 Barro 2.56 2.56 5.13 30.77 30.77 15.38 12.82 39 Guatapo 2.63 2.63 5.26 23.68 26.32 18.42 21.05 38 Masaguaral 0 7.14 7.14 21.43 25 14.28 25 28 Cuzco 3.95 5.26 7.89 21.05 26.31 18.42 17.1 76 Manu 4.48 7.46 8.96 23.88 26.86 13.43 14.93 67 Belem 1.59 6.35 7.94 20.63 15.87 20.63 26.98 63

Table 5.2.5. Percentage of taxa in ecological diversity categories for diet for modern and fossil localities. ‘Total’ refers to the number (not percentage) of taxa included for each locality

127 CHAPTER 5

Site Root/fungi Granivore Grazer Brows/frug Omnivore Insectivore Carnivore Total diet ENiger 3 024.3 33.3 18.2 12.1 9.1 CongoBasin 3.8 0 9.4 41.5 13.2 5.7 26.4 Knysna 11.8 05.9 29.4 23.6 0 29.4 Kilimanjaro 5.3 0 15.8 47.4 21 0 10.5 WNiger 6.3 018.8 34.4 28.1 6.3 6.3 ECross 5 015 35 25 5 15 Makakou 4.9 017.1 39 26.8 0 12.2 RwenzoriNP 2 025.5 21.6 15.6 3.9 31.4 GuineaWood 1.7 0 28.9 23.7 8.5 5.1 32.2 NatalWood 1.9 0 35.8 20.8 9.5 3.8 35 LakeMweru 0 0 40 25.7 5.8 0 28.6 WLungaNP 2.3 0 34.1 20.5 11.4 0 31.8 SerengetiNP 1.3 0 34.7 18.7 9.4 6.7 29.3 SerengetiBush 1.6 0 34.4 18.8 7.8 6.3 31.3 RukwaValley 1.9 0 36.5 19.3 9.7 3.9 28.9 KafueNP 1.8 032.7 14.6 10.9 5.5 34.5 KrugerNP 1.6 033.3 17.4 8 4.8 34.9 LinyantiSwamp 0 0 51.3 14.3 5.8 0 28.6 SSWoodland 1.2 0 39.8 16.8 6 6 30.1 Sudan 2 029.3 15.7 9.8 3.9 39.2 SWarid 1.7 039.9 13.4 3.4 8.3 33.3 KalahariTV 6.7 0 33.3 6.7 0 20 33.3 Sahel 3.2 032.3 6.5 9.7 6.5 41.9 Chobe 1.8 041.1 12.5 7.2 8.9 28.6 AmboseliNP 2.2 0 36.9 17.4 8.7 6.5 23.9 TarangireNP 2.1 0 33.4 18.7 8.3 10.4 27.1 Okavango 1.9 037.1 11.2 7.5 9.3 33.3 KafueFlats 0 045.9 10.8 5.4 0 37.8 SerengetiPlains 0 0 47.4 0 10.5 10.5 31.6 SSGrassland 2.4 0 41.4 7.3 0 9.8 36.6 Namibdesert 5.6 0 22.2 11.1 0 5.6 50 Camel Sputum 0 2.5 0 57.5 25 10 5 40 Mike's 0 5.88 0 47.06 23.53 17.65 5.88 17 Menagerie Quantum Leap 0 7.14 0 28.57 14.29 50 0 14 Encore 0 04.55 63.63 9.09 9.09 13.64 22 Ringtail 0 00 46.67 26.67 6.67 20 15

Table 5.2.5. continued…

128 CHAPTER 5

habitat (Andrews and O’Brien 2000) so in the cases of Camel Sputum, Mike’s

Menagerie, Quantum Leap, Encore and Ringtail Sites, the more conservative approach of excluding bats is taken. For the Bitesantennary Site local fauna, only bat community structure should be analysed due to the major under representation of other orders of mammals.

5.2.3.3. Statistical analyses – determining habitat types and annual rainfall for fossil communities

The procedure for grouping modern mammalian communities, as described by their ecological diversity variables, into ‘broad habitat types’ is outlined as follows: cluster analysis was applied to the ecological diversity variables for modern communities; principal co-ordinates analysis was used to confirm the groups formed by the cluster analysis; these groups/clusters of mammalian communities are referred to as ‘broad habitat types’. To predict which broad habitat types the fossil communities most likely fit into, based on ecological diversity variables, multiple discriminant analyses

(canonical variates analysis) were carried out. Following this, a simple linear regression analysis was performed on principal co-ordinates scores and average annual rainfall for sites of the modern communities in order to predict a range of annual rainfall for

Riversleigh study sites.

The cluster analyses were carried out on two data sets; one for the diet categories only and including all modern sites (referred to as ‘Diet Only’) and one for the diet and locomotion categories excluding modern sites from Africa (referred to as ‘Diet and

Locomotion’). African sites are excluded from the latter data set because the locomotor categories for the African sites are not equivalent to locomotor categories used in the

129 CHAPTER 5

current study (the ‘small mixed’ category is not included for the African sites). This could not be determined for African sites because Reed (1998) does not provide species lists for each site separately, but only percentages of species in particular ecological diversity categories.

Cluster analysis groups objects on the basis of similarity (Hair et al. 1998). In this study, the similarity measure used was the correlation coefficient. This takes the patterns across the variables, or their structural characteristics, into account rather than their magnitudes (Hair et al. 1998). A heirarchical cluster procedure was used so that the number of clusters did not have to be predetermined. Average linkage, or paired group method was used to create clusters with approximately the same variance (Hair et al. 1998). The analysis of the Diet and Locomotion data set will presumably be more sensitive than that on the Diet Only data set because the adaptive structure of the community is represented in more detail, however, the Diet and Locomotion data set includes considerably fewer modern sites. Both analyses were performed and an attempt was made to combine/merge the results.

Cluster solutions were validated by splitting the sample of each data set and running each sub sample through cluster analysis again, then comparing resulting clusters to those formed for the entire data set in each case. Ordination techniques were used in conjunction with cluster analysis as further support for cluster groups as advised by

Sneath and Sokal (1973) and Field et al. (1982). The ordination technique used was principal co-ordinates analysis because distance/similarity measures can be specified and, in this case, needed to be correlation measures as for the cluster analyses. Principal

130 CHAPTER 5

co-ordinates analysis supports cluster groupings visually by displaying the inter-point relationships and the areas of greatest variation.

Multiple discriminant, or canonical variates, analysis was used to group faunas from

Riversleigh fossil sites, using their ecological diversity profiles, into the broad habitat types formed by the cluster analysis of modern localities. This type of analysis is described and used in Chapter 3. The fauna from Ringtail Site could only be analysed through the use of the Diet Only data set because locomotor behaviour information was not available (the calcaneum sample size from this site is very small). Fossil faunas from Camel Sputum, Mike’s Menagerie, Quantum Leap and Encore Site were grouped into broad habitat types using both the Diet Only and the Diet and Locomotion data sets.

Rainfall is related to vegetation structure and productivity (Damuth et al. 2002) and has been found to correlate with vegetation complexity in the lowland tropics (Gentry 1988;

Kay and Madden 1997). Because vegetation determines animal community structure, mammalian community structure may be used to predict rainfall for fossil sites.

Principal co-ordinates scores for modern mammalian communities, obtained from ecological diversity variables, were regressed on average annual rainfall at the localities of these communities, separately for each data set. Even though these regressions use the same ecological diversity data as the cluster analyses that create habitat groups, the regressions and cluster analyses are independent and provide distinct information. The cluster analyses group localities based on the similarity between their ecological diversity characteristics, whereas the regression analyses identify correlations between the ecological diversity characteristics and annual rainfall at the sites. It is therefore,

131 CHAPTER 5

possible to compare and combine the results obtained in each of these analyses in the interpretation of palaeoenvironment for the fossil sites.

Cluster and principal coordinates analyses and simple linear regressions were performed with PAST statistical software (Hammer et al. 2004); multiple discriminant analyses were performed with SYSTAT version 7.0 (SPSS 1997); graphs and figures of results were produced with PAST software.

5.3. Results

5.3.1. Consideration of potential bias in fossil community representation

Taphonomic analyses of the Riversleigh fossil assemblages (Chapter 2) revealed no bias in taxonomic representation due to predation, hydraulic transport or spatial mixing of elements at the sites, which led to the interpretation of the fossil assemblages as single communities in space. It is considered that time averaging for Riversleigh faunas may not have had a significant effect on the ecological diversity characteristics/community characterisation, provided environmental conditions were stable. These analyses however, did not investigate the extent to which remains at the sites (even if they are from a single community) are representative of the original community. Further examination of the data has been undertaken here.

Logarithms of abundance plotted against body mass for primary consumers from

Riversleigh fossil sites shows that slopes for all sites are outside the range (-0.80 to -

1.30) specified by Damuth (1982) for an unbiased assemblage (Fig. 5.3.1). Relative abundance of species therefore, is not representative of a community for any of the

132 CHAPTER 5

fossil sites included here. Camel Sputum, Quantum Leap and Ringtail Sites have a considerable under-representation of individuals of small species (low body mass).

Mike’s Menagerie and Encore Sites have especially low numbers of individuals within species of low body mass. This result appears to be in contradiction to the results from the taphonomic analysis (Chapter 2) that reveal much greater specimen numbers for small animals than for medium sized or large animals at most sites. A possible explanation is that not all specimens from the sites have been identified to species-level and counts used here are based predominantly on literature where specimens have been assigned to species with certainty. I make no attempt to personally assign unidentified specimens to species-level taxa. Consequently, many of the specimens included in the taphonomic analysis have not been included in the current analysis of species relative abundance.

133 CHAPTER 5

0.35 1.8 CS MM

0.3 1.6 1.4 0.25 1.2 0.2 1

0.8 0.15 log abundance log abundance 0.6 0.1

0.4 y = -0.005x + 0.1034 0.05 0.2 y = -0.2159x + 1.1433 0 0 0123456 0123456 log body mass log body mass

0.6 QL 1.8 EC

0.5 1.6 1.4 0.4

1.2 0.3 1 0.2 0.8 log abundance log abundance 0.1 0.6 y = 0.1355x - 0.1037

0 0.4 00.511.522.533.544.5 -0.1 0.2 y = -0.2412x + 0.8065 0 -0.2 0123456 log body mass log body mass

1 RG

0.8

0.6

0.4

log abundance 0.2

0 012345 y = -0.2447x + 1.0172 -0.2 log body mass

Figure 5.3.1. Logarithms of species abundance against body mass for primary consumers from Riversleigh fossil sites. Red lines represent expected gradient (-1.05) for no taphonomic bias (sensu Damuth 1982). Abbreviations are as follows: CS, Camel Sputum Site; MM, Mike’s Menagerie Site; QL, Quantum Leap Site; EC, Encore Site; RG, Ringtail Site.

134 CHAPTER 5

Abundance of individuals within species is not representative for the fossil sites, for those individuals that have been identified to species-level, however, this does not mean that species presence itself is not representative of the original communities in terms of adaptive structure. Figure 5.3.2. shows the range of the proportion of species within diet/body mass groups for twenty nine modern Australian faunas with the proportion of species in each group for the fossil sites superimposed. The ranges for the Australian modern faunas are similar to those for sites around the world, presented by Soligo

(2002); however, secondary consumers over 45 kg and primary and secondary consumers over 180 kg are not present in Australian faunas.

The Ringtail Site fossil fauna has an over-representation of primary consumer species under 1 kg body mass; the Quantum Leap Site fossil fauna has a slight over- representation of secondary consumer species under 1 kg body mass; and the Camel

Sputum and Encore Site fossil faunas have an over-representation of primary consumer species of 1-10 kg body mass. All other diet/body mass group proportions for the fossil sites are within the range for modern faunas. Thus, despite numbers of individuals within small species being under-represented for the fossil sites, in most cases small species numbers are over-represented. The Quantum Leap fossil fauna has no representatives in the 1-10 kg secondary consumer category, or in the 10-45 kg primary and secondary consumer categories. This may indicate a bias in the size of animals preserved there, despite the fact that it does not fall outside the range for modern faunas.

Note that Quantum Leap Site has produced a representative in the >45 kg primary consumer category, however, this is based on isolated molar teeth which may have fallen into the deposit even if there was a physical restriction on the size of animals

135 CHAPTER 5

0.9

0.8

0.7

0.6

0.5

0.4

0.3 Relative abundance

0.2

0.1

0 1 2 3 4 5 6 7

Diet/body mass group

Figure 5.3.2. Boxplots showing range of relative abundance of species covered by 25 modern Australian faunas for the diet/body mass groups. Superimposed are values for Riversleigh fossil faunas (refer to Table 5.2.1). Symbols are as follows: 1 – AP (primary consumer, < 1 kg) Ringtail Site 2 – AS (secondary consumer, <1 kg) Quantum Leap Site 3 – BP (primary consumer, 1-10 kg) Camel Sputum Site 4 – BS (secondary consumer, 1-10 kg) Mike’s Menagerie Site 5 – CP (primary consumer, 10-45 kg) Encore Site 6 – CS (secondary consumer, 10-45 kg) 7 – DP (primary consumer, >45 kg)

136 CHAPTER 5

preserved. All of the other fossil faunas contain species in the 0-10 kg body mass groups. Soligo (2002) explains that more significance should be placed on species that occur in greater numbers rather than the less numerous species. Species numbers are higher in the under 10 kg body mass range for modern and fossil faunas, so it is presumed that the absence of species in the 10-45 kg body mass range for Camel

Sputum, Mike’s Menagerie, Quantum Leap and Ringtail Site fossil faunas will have minimal impact.

To test if the over-representation of small mammals at fossil sites affects their palaeohabitat interpretations, the ecological diversity variables for Camel Sputum,

Quantum Leap, Ringtail and Encore Sites were adjusted to fit within the range for modern Australian faunas. Table 5.3.1 shows these adjusted values, and the resulting classification into habitat type for the Diet Only data set is presented in Table 5.3.8, with the classification results for the multiple discriminant analysis of the actual values.

No difference is seen in the grouping of the fossil sites into habitat types with the altered values; the only effect is that their Mahalanobis distances from the habitat group centroids are reduced. This indicates that the over-representation of small mammals at

Camel Sputum, Quantum Leap, Ringtail and Encore Sites does not affect palaeohabitat interpretations for the sites. The adaptive structure of these fossil faunas is therefore considered to be reliable for palaeoecological analyses. A bias against small mammals

(which is not the case here) is presumed to lead to loss of palaeoecological information

(Soligo 2002).

137 CHAPTER 5

Root/ Browser/ Site Granivore Grazer Omnivore Insectivore Carnivore Total fungi frugivore CS 0 (0) 2.5 (2.5) 0 (0) 50.5 (57.5) 25 (25) 12 (10) 10 (5) 40 7.14 14.29 QL 0 (0) 0 (0) 33.57 (28.57) 40 (50) 5 (0) 14 (7.14) (14.29) 4.55 14.09 12.09 13.64 EC 0 (0) 0 (0) 55.63 (63.63) 22 (4.55) (9.09) (9.09) (13.64) 29.67 RG 0 (0) 0 (0) 0 (0) 40.67 (46.67) 9.67 (6.67) 20 (20) 15 (26.67)

Table 5.3.1. Altered ecological diversity variables for fossil sites (CS, Camel Sputum; QL, Quantum Leap; EC, Encore; RG, Ringtail). Numbers in red indicate actual values. Adjustments were made by increasing/decreasing ecological diversity variables that fall outside the range expected for modern Australian faunas in order to place them within the expected range.

5.3.2. Ecological diversity analyses

5.3.2.1. Determining broad habitat types from modern communities

Results of the cluster analysis for all modern communities, for the Diet Only data set are shown in Fig. 5.3.3. The clusters are supported in the principal co-ordinates analysis of this data (Fig. 5.3.4); the affinity of Tas WS to other localities is not well defined. The cluster analysis is validated by splitting the sample (results shown Appendix C7.1). The clusters formed in the split-sample analysis mostly do not differ from those for the entire sample. Only two modern communities from Group A (Monte and Upper

Montane from Argentina) have clustered differently to the un-split sample, and three communities from Group B (Chamela, Masaguaral and Belem) that clustered together in the un-split sample have been separated into different clusters. Broad habitat types assigned to the clusters of modern communities are indicated in Figure 5.3.3.

138 CHAPTER 5

Figure 5.3.3. Cluster analysis results on diet categories for all modern sites. Broad habitat types are indicated above clusters

OPEN 1 FOREST 2 FOREST 3 FOREST 1 OPEN 2

1 Law nHill Tundra Serengeti RwenzoriN GuineaWoo NatalWood KrugerNP KafueNP Sudan WLungaNP LakeMweru Serengeti Serengeti Rukw aVall SSWoodlan Tarangire AmboseliN LinyantiS SWarid Okavango Chobe KafueFlat SSGrassla KalahariT Sahel Namibdese V.GrGW Puna Prepuna V.GrOF TasWS Mossman Guatapo Cuz c o Atherton Barro Manu LeeUp Ingham Dtf MTf V.GipDLF Kuringai V.GipRF V.GipWLF V.GipCTR V.GipMW CapeYor k Lamington V.GipWTF New Guinea V.GrHRW V.GrWDF Jervis MtAzules LaSelva CongoBasi Knysna Kilimanja ECr os s Makakou WNiger V.GipRW ENiger TasH TasS TasR Ws p Dbm Vhf Low Montan Masaguara Belem UppMontan Chamela Cent Aus t Transitio V.GipCH V.GipHW SouthCoas V.GipSGH Monte MontChaco Chac o

0.9

0.8

0.7

0.6

0.5

Similarity

0.4

0.3

0.2

0.1

10 20 30 40 50 60 70 80 139 CHAPTER 5

0.3

FOREST 3 Kilimanjaro Makakou 0.2 ECross CongoBasin WNiger Knysna ENiger OPEN 1 LakeMweru FOREST 2 0.1 WLungaNPGuineaWoodRwenzoriNP V.GipRW NatalWoodRukwaValley AmboseliNP LinyantiSwampKrugerNPSerengetiBushLawnHillSerengetiNP LeeUpAthertonBarro SSWoodlandTarangireNP TasWS LaSelvaMTfManu KafueFlatsKafueNPSudan MossmanNewGuinea SWaridChobe V.GipMW V.GipCTR 0 V.GrOFCapeYork Dtf OkavangoNamibdesert LamingtonMasaguaralV.GipDLFV.GipRF Sahel Guatapo SSGrassland TasH LowMontane V.GipWLFMtAzules Cuzco Wsp Kuringai V.GrHRWV.GipWTF PrepunaSerengetiPlains Ingham Tundra V.GrGW Dbm Belem Puna TasS Vhf Jervis -0.1 2 Coordinate KalahariTV TasR V.GrWDF FOREST 1 SouthCoast Chamela

UppMontane V.GipHW -0.2 OPEN 2 Transitional Chaco V.GipCH V.GipSGH MontChacoMonte

-0.3 CentAust

-0.1 0 0.1 0.2 Coordinate 1

Figure 5.3.4. Plot of principal co-ordinates 1 and 2 for diet categories for all modern sites. Colours correspond to broad habitat types from cluster analysis.

Results of the cluster analysis for the Diet and Locomotion data set, excluding African sites are shown in Fig. 5.3.5. The clusters are supported in the principal co-ordinates analysis of this data set (Fig. 5.3.6); again, the affinity of TasWS to other localities is not entirely clear. The cluster analysis is validated by splitting the sample (results shown in Appendix C7. 2). The clusters formed in the split-sample analysis are very similar to those of the entire sample, with only one community being re-clustered in each case.

Broad habitat types assigned to the clusters of modern communities are indicated in

Figure 5.3.5.

140 CHAPTER 5

GRASS/DESERT DENSE FOREST MIXED FOREST TEMPERATE MIX

CentAust Prepuna Monte Chamela UppMontan V.GrGW Law nHill Tundra Puna TasH MontChaco Chaco Mossman Guatapo Ingham Atherton New Guinea Dtf MTf LaSelva Barro Cuzco Manu V.GipDLF V.GipRF LeeUp V.GipWLF V.GipCTR Kuringai Jervis V.GipWTF V.GrHRW V.GrWDF V.GrOF SouthCoas CapeYork Lamington V.GipCH V.GipHW V.GipSGH V.GipRW V.GipMW TasWS TasS Wsp Dbm Vhf TasR Low Montan Transitio Belem MtAzules Masaguara

0.9

0.8

Similarity

0.7

0.6

0.5 10 20 30 40 50

Figure 5.3.5. Cluster analysis results on diet and locomotion categories for all modern sites excluding African sites. Broad habitat types are indicated for clusters.

141 CHAPTER 5

0.3

MIXED FOREST V.GrGW 0.2 V.GrHRW V.GipSGH Jervis Puna V.GipCH V.GrWDFSouthCoast V.GipCTRV.GipWLF LawnHill 0.1 V.GrOFV.GipHW V.GipRWLamington Chaco V.GipMWCapeYork Tundra NewGuinea V.GipWTF Kuringai CentAust V.GipRF GRASS/DESERT LeeUpV.GipDLF 0 Ingham Atherton TasWS Barro Cuzco Transitional MontChaco LaSelva MIX Manu Guatapo Monte -0.1 MtAzules Mossman TasH TasR Prepuna Dtf

Coordinate 2 Coordinate Masaguaral TasS DENSE FOREST Belem Dbm -0.2 MTf Chamela LowMontane Wsp Vhf TEMPERATE -0.3

-0.4 UppMontane

-0.2 -0.1 0 0.1 0.2 0.3 Coordinate 1

Figure 5.3.6. Plot of principal co-ordinates 1 and 2 for diet and locomotion categories for all modern sites excluding Africa. Colours correspond to broad habitat types from cluster analysis.

Descriptions of the broad habitat types derived from the cluster analyses are presented in the discussion of this chapter. Table 5.4.1, in the discussion section, indicates which habitat types from the two data sets are presumed to be equivalent.

5.3.2.2. Grouping fossil communities into broad habitat types

Cluster and principal co-ordinates analyses do not assume normality of variables, however, multiple discriminant analysis does. Assumptions of multiple discriminant analysis are described in Chapter 3. Normality of each variable was statistically tested using Zskewness values as described in Chapter 3. For the Diet Only data set, the grazer, browser, omnivore, insectivore, and carnivore variables are normally distributed (see

Appendix C8.1 for Zskewness values); the root/fungus variable is normally distributed when log transformed; and the granivore variable is not included in multiple

142 CHAPTER 5

discriminant analyses for this data set because it is not normally distributed even if log transformed. All variables are included for the Diet and Locomotion data set.

Root/fungi, granivore, grazer, small mixed, and ‘terrestrial + hoppers’ are log transformed in order to obtain normal distribution (see Appendix C8.2 for Zskewness values).

The broad habitat types determined by cluster analyses have approximately the same variance because the average-linkage or paired group method was used to create the clusters (Hair et al. 1998). Thus the assumption of equivalence of variances across groups for multiple discriminant analysis is satisfied. Tolerance values for the Diet Only data set are over 0.3 for all variables used, and for the Diet and Locomotion data set are over 0.1 for all variables used. This indicates little multicollinearity of variables. Tables

5.3.2 and 5.3.3 show summary statistics for the two data sets used in multiple discriminant analyses.

Group Statistic GRAZER BROWS OMNI INSECT CARN LOG ROOT X 37.77 13.54 7.70 7.32 30.36 0.80 OPEN 1 37.80 25.70 20.00 22.22 50.00 2.04 (N=29) OR SD 7.99 7.63 5.08 5.86 9.15 0.60 X 16.52 8.35 18.48 31.14 19.81 0.21 OPEN 2 16.15 16.67 23.63 16.24 23.89 1.88 (N=9) OR SD 5.03 5.94 7.63 4.97 8.22 0.63 X 14.90 11.59 22.85 15.02 31.45 0.26 FOREST 1 15.94 21.43 31.38 13.49 25.00 1.93 (N=11) OR SD 5.38 6.84 8.89 4.10 6.89 0.62 X 10.87 24.05 26.94 18.08 14.19 0.82 FOREST 2 19.52 18.51 23.06 21.30 21.43 1.75 (N=26) OR SD 5.73 4.07 5.60 4.94 4.93 0.74 X 16.07 35.97 22.27 5.03 15.70 1.61 FOREST 3 18.40 19.62 14.90 12.10 23.10 2.55 (N=8) OR SD 6.13 6.45 4.84 4.85 8.24 0.74

Table 5.3.2. Summary statistics for broad habitat types from ‘Diet Only’ data set. X, group means; OR, observed ranges; SD standard deviations; N, number of modern community localities within each group; GRAZER, grazing category; BROWS, browsers; OMNI, omnivores; INSECT, insectivores; CARN, carnivores; LOG ROOT, log (root/fungus).

143 CHAPTER 5

Group Statistic BR OM INS CA AR SC OTH LSM LTH LRF LGN LGZ 5.27 11.80 20.18 24.42 1.95 13.81 2.14 3.60 3.69 0 1.77 3.23 GRASS/ X DESERT OR 23.53 25.00 37.50 50.00 7.69 44.44 7.69 1.58 1.28 0 3.54 1.79 (N=12) SD 7.43 7.91 10.61 12.95 3.13 12.63 2.94 0.44 0.35 0 1.21 0.59 10.66 27.83 13.81 30.49 3.36 22.12 6.76 3.58 3.44 0.32 0.26 2.80 TEMPE- X RATE OR 11.62 23.73 11.61 10.71 8.00 8.84 9.05 0.31 0.49 1.93 1.55 0.66 (N=6) SD 4.59 8.32 4.34 3.53 3.74 3.12 3.36 0.11 0.20 0.79 0.63 0.24 X 19.44 19.76 20.67 24.32 12.67 20.05 5.13 3.28 3.56 0.24 2.80 2.16 MIX 7.79 9.13 18.28 10.52 7.03 6.66 4.49 0.25 0.30 0.95 1.82 0.49 (N=4) OR SD 3.69 3.83 8.40 4.78 3.39 3.03 2.00 0.11 0.13 0.48 0.75 0.21 20.57 24.81 21.12 13.56 13.24 9.13 4.85 3.54 3.62 0.55 0.76 2.72 MIXED X FOREST OR 20.88 12.77 29.37 19.01 18.27 18.18 10.53 0.77 0.58 1.88 2.55 1.75 (N=20) SD 5.45 4.03 7.17 4.78 4.75 5.01 3.68 0.20 0.13 0.78 1.00 0.36 26.07 29.82 15.68 15.23 18.65 15.74 4.87 3.55 3.16 1.11 1.57 1.70 DENSE X FOREST OR 14.85 20.54 14.97 16.30 18.23 9.36 7.63 0.81 0.79 1.70 2.55 1.20 (N=11) SD 4.67 6.21 4.56 4.69 5.98 3.41 2.61 0.28 0.21 0.58 0.65 0.39

Table 5.3.3. Summary statistics for broad habitat types from ‘Diet and Locomotion’ data set. X, group means; OR, observed ranges; SD standard deviations; N, number of modern community localities within each group; BR, browsers; OM, omnivores; INS, insectivores; CA, carnivores; AR, arboreal; SC, scansorial; OTH, other locomor behaviour; LSM, log (small mixed); LTH, log (terrestrial + hoppers); LRF, log (root/fungi); LGN, log (granivores); LGZ, log grazers.

For the Diet Only data set, the mean of the grazer variable is greatest for the OPEN 1 broad habitat type; the browser variable means are considerably higher for FOREST 2 and FOREST 3 habitat types; the omnivore variable has a considerably lower mean value for OPEN 1 habitat type; the mean of the insectivore variable is highest for OPEN

2 habitat type; and the carnivore variable means are lower for FOREST 2 and FOREST

3 habitat types. For the Diet and Locomotion data set, the log grazer variable means are highest for the GRASS/DESERT and lowest for the DENSE FOREST habitat types; the log (root/fungus) variable mean is highest for the DENSE FOREST habitat type; the log granivore variable mean is highest for the MIX habitat type; the browser and omnivore variable means are lowest for the GRASS/DESERT habitat type; carnivore variable means are lower for MIXED FOREST and DENSE FOREST habitat types than for the others; and TEMPERATE and GRASS/DESERT habitat types have low arboreal variable means.

144 CHAPTER 5

Figure 5.3.7 (A and B) shows the pattern of separation on the canonical variates plot for the modern communities using the Diet Only data set. Score 1 distinguishes OPEN 1 from all other habitat types, and FOREST 2 from FOREST 3; score 2 distinguishes

FOREST 1 and OPEN 2 from FOREST 2 and FOREST 3. FOREST 1 and OPEN 2 are distinguished on score 3. Figure 5.3.8 (A and B) shows the pattern of separation on the canonical variates plot for the modern communities using the Diet and Locomotion data set. Score 1 distinguishes GRASS/DESERT from all other habitat types; TEMPERATE from MIXED FOREST and DENSE FOREST; and MIX from DENSE FOREST. Score

2 distinguishes DENSE FOREST and MIX habitat types from TEMPERATE. A combination of scores 1 and 3 separates MIXED FOREST from DENSE FOREST.

Eigenvalues (Tables 5.3.4 and 5.3.5) indicate the variation explained by the canonical axes of plots. For the Diet Only data set, the first canonical variate accounts for 76.5% of the variation in proportional distribution of communities among broad habitat types;

19.8% of the variation is explained by the second canonical variate; and 3.7% variation in total is explained by the third and fourth variates. For the Diet and Locomotion data set, 74.5% of variation is explained by the first canonical variate; 12.3% by the second; and 13.2% in total by the third and fourth variates.

145 CHAPTER 5

A. 5

3 OPEN 2 FOREST 4 2

0 OPEN 1

2 Score -1

-2 FOREST 2 -3 FOREST 3

-4

-5

-4 -3 -2 -1 0 1 2 3 4 5 6 Score 1

B. 3

OPEN 2 OPEN 1 1 FOREST 3

0 FOREST 2

3 Score -1

-2 FOREST 4

-3

-4 -3 -2 -1 0 1 2 3 4 5 6 Score 1

Figure 5.3.7. Canonical variates plots for Diet Only data set. A: score 1 against score 2; B: score 1against score 3.

146 CHAPTER 5

4 A.

TEMPERATE 3 MIXED FOREST 2

0 Score 2 GRASS/DESERT

-1

MIX -2

DENSE FOREST -3

-5 -4 -3 -2 -1 0 1 2 3 4 5 Score 1

B. 4

3 DENSE FOREST TEMPERATE

GRASS/DESERT 1

Score 3 MIX 0

-1 MIXED FOREST -2

-5 -4 -3 -2 -1 0 1 2 3 4 5 Score 1

Figure 5.3.8. Canonical variates plots for Diet and Locomotion data set. A: score 1 against score 2; B: score 1against score 3.

147 CHAPTER 5

CANONICAL VARIATE VARIABLE 1 2 3 4 CONSTANT -4.100 -2.942 0.482 -5.939 GRAZER 0.171 (1.120) 0.014 (0.090) 0.030 (0.198) 0.005 (0.030) BROWSER 0.022 (0.138) -0.066 (-0.417) 0.015 (0.093) 0.115 (0.724) OMNIVORE -0.040 (-0.243) 0.011 (0.068) -0.079 (-0.487) -0.025 (-0.156) INSECTIVORE -0.079 (-0.410) 0.125 (0.647) 0.090 (0.465) 0.105 (0.545) CARNIVORE 0.060 (0.457) 0.098 (0.736) -0.073 (-0.547) 0.114 (0.857) LOGROOT/ 0.553 (0.368) -0.459 (-0.305) 0.586 (0.390) 0.183 (0.122) FUNGUS Eigenvalue 10.127 2.630 0.467 0.023 Variance (%) 76.5 19.8 3.5 0.2

Table 5.3.4. Canonical discriminant functions (values in brackets are standardised coefficients for canonical variates) for Diet Only data set.

CANONICAL VARIATE VARIABLE 1 2 3 4 CONSTANT -16.036 -20.886 43.026 12.981 BROWSER 0.117 (0.669) 0.139 (0.792) -0.100 (-0.572) -0.152 (-0.869) OMNIVORE 0.118 (0.722) 0.188 (1.151) -0.027 (-0.165) -0.115 (-0.702) INSECTIVORE -0.013 (-0.094) 0.075 (0.553) -0.081 (-0.603) -0.098 (-0.723) CARNIVORE 0.023 (0.173) 0.114 (0.852) -0.050 (-0.372) -0.130 (-0.965) ARBOREAL 0.165 (0.754) -0.057 (-0.262) -0.129 (-0.587) 0.062 (0.284) SCANSORIAL -0.060 (-0.427) 0.031 (0.218) -0.032 (-0.226) -0.025 (-0.174) OTHER LOC. - - - - LOGSMMIX 1.312 (0.367) 1.664 (0.466) -3.502 (-0.980) 1.731 (0.485) LOGTERRH 2.234 (0.505) -0.973 (-0.220) -5.880 (-1.328) -1.962 (-0.443) LOGROOT/F 0.333 (0.209) 0.771 (0.484) 0.031 (0.019) 0.239 (0.150) LOGGRAN 0.508 (0.482) -0.225 (-0.213) -0.220 (-0.209) -0.831 (-0.788) LOGGRAZ -1.190 (-0.486) 3.188 (1.301) -1.176 (-0.480) -0.815 (-0.333) Eigenvalue 9.054 1.495 1.216 0.385 Variance (%) 74.5 12.3 10 3.2

Table 5.3.5. Canonical discriminant functions (values in brackets are standardised coefficients for canonical variates) for Diet and Locomotion data set. The variable ‘Other locomotion’ is not included because this variable is below the tolerance level of 0.1.

A global test of differences between multivariate means showed a significant difference between habitat types for both data sets. For the Diet Only data set: Pillai’s trace =

1.975, d.f. = 24, 304, P<0.001; for the Diet and Locomotion data set: Pillai’s trace =

2.326, d.f. = 44, 160, P<0.001. Tables 5.3.6 and 5.3.7 show the classification results.

148 CHAPTER 5

Allocated habitat type OPEN 1 OPEN 2 FOREST 1 FOREST 2 FOREST 3 OPEN 1 (29) 100/100 0/0 0/0 0/0 0/0 OPEN 2 (9) 0/0 100/100 0/0 0/0 0/0 FOREST 1 0/0 0/0 82/82 18/18 0/0 (11) FOREST 2 0/0 0/0 0/0 100/100 0/0 (26) Actual habitat type type Actual habitat FOREST 3 (8) 0/0 0/0 0/0 0/12 100/88

Table 5.3.6. Classification results from canonical variates analysis of Diet Only data set (values indicate the percentage of allocations of individuals to each group; values in italics indicate jackknife classification results; values in parentheses are the numbers of modern communities in each group)

Allocated habitat type MIXED DENSE GRASS/DESERT TEMPERATE MIX FOREST FOREST GRASS/DESERT (12) 100/67 0/25 0/0 0/8 0/0 TEMPERATE 0/0 100/83 0/0 0/17 0/0 (6) MIX (4) 0/0 0/0 100/75 0/25 0/0 MIXED 0/0 0/0 5/11 95/ 84 0/5 FOREST (19) Actual habitat type type habitat Actual DENSE 0/0 0/0 0/0 0/0 100/100 FOREST (11)

Table 5.3.7. Classification results from canonical variates analysis of Diet and Locomotion data set (values indicate the percentage of allocations of individuals to each group; values in italics indicate jackknife classification results; values in parentheses are the numbers of species for each group)

The communities from the Riversleigh sites used in this analysis were allocated to broad habitat types as indicated in Table 5.3.8. This table also includes miss-classified modern communities and the habitat types that they most likely belong to.

149 CHAPTER 5

‘DIET ONLY’ DATA SET LOCALITY OPEN 1 OPEN 2 FOREST 1 FOREST 2 FOREST 3 (113.1), 0% (75.4), 0% (68.3), 0% (35.9), 67% (37.4), 33% CS [98.8, 0%] [55.9, 0%] [49.8, 0%] [24.1, 98%] [31.8, 2%] MM (105.4), 0% (48.4), 0% (50.3), 0% (22), 100% (36.9), 0% (172.7), 0% (44.7), 100% (86.1), 0% (58.9), 0% (114.9), 0% QL [137.6, 0%] [34, 88%] [62.9, 0%] [38, 12%] [81, 0%] (101.4), 0% (90.3), 0% (81.1), 0% (54.9), 0% (43.3), 100% EC [88.2, 0%] [64.8, 0%] [58.3, 0%] [35.1, 24%] [32.8, 76%] (82.4), 0% (51.7), 0% (36.8) 0% (21.1), 93% (26.4), 7% RG [81.1, 0%] [40, 0%] [28.1, 0%] [14.7, 100%] [27.4, 0%] South Coast (37.7), 0% (15.3), 3% (18.1), 1% (8.5), 95% (19), 1% Tas WS (19.4), 1% (29.2), 0% (10.5), 71% (14.1), 12% (13.5), 16% Masaguaral (53.9), 0% (14.2), 1% (6.1), 41% (5.3), 59% (26.2), 0% Belem (44.2), 0% (6.8), 27% (7), 25% (5.4), 47% (24.9), 0% ‘DIET AND LOCOMOTION’ DATA SET LOCALITY GRASS/ MIXED DENSE TEMPERATE MIX DESERT FOREST FOREST CS (283.4), 0% (286.5), 0% (175.3), 0% (183.5), 0% (148.6), 0% MM (255.3), 0% (271.9), 0% (164.3), 0% (183.5), 0% (157.8), 0% QL (205.8), 0% (253.2), 0% (151.8), 0% (169.9), 0% (149.2), 0% EC (261.4), 0% (270.7), 0% (176), 0% (195.9), 0% (185.6), 0% V.GipCH (25.7), 0% (32.4), 0% (12.2), 66% (13.5), 34% (35), 0% Tas WS (21.6), 0% (6.4), 100% (26.6), 0% (17.4), 0% (39.7), 0%

Table 5.3.8. Mahalanobis distance-square from group means (in brackets) and percent probabilities for broad habitat type allocations for fossil communities and miss-classified modern communities for the two data sets. Values in red are the Mahalanobis distances, followed by percent probabilities, for broad habitat type allocations for fossil faunas, based on the altered ecological diversity variables for those sites (see section 5.3.1, p. 137-138 for explanation). Abbreviations: CS, Camel Sputum; MM, Mike’s Menagerie; QL, Quantum Leap; EC, Encore; RG, Ringtail. Fossil sites were not grouped within habitat type in the ‘Diet and Locomotion’ data set analysis because their Mahalanobis distances are very large. Shading indicates smallest Mahalanobis distance.

5.3.2.2. Determining annual rainfall for fossil communities

Simple linear regressions on the first principal co-ordinates (PCO) score against average annual rainfall for modern faunas for both data sets (Figs 5.3.9 and 5.3.10) reveal that there are significant correlations. The assumption of normality of variables is satisfied when the square root of rainfall values is taken and, for the Diet Only data set, the PCO values are cubed. The PCO values for the Diet and Locomotion data set are normally distributed. The Zskewness values for each variable are as follows: all modern sites (square root of rainfall) = 2.21; Diet Only (PCO)3 = 0.83; Diet and Locomotion PCO = 0.26;

150 CHAPTER 5

-4 0.006 gradient = 1.74x10 y-intercept = -5.96x10-3 0.005 Pearson’s r = 0.73 p(uncorr) = 3.23x10-15 0.004

95% on gradient: 0.003 1.43x10-4; 2.08x10-4 0.002

95% on y-intercept: 0.001 -7.12x10-3; -4.92x10-3

0 PCOSc1Cube -0.001 1) score (PCO -0.002

-0.003

-0.004

10 20 30 40 50 60 70 Square root of average annual rainfall (mm)

Figure 5.3.9. Simple linear regression of square root of average annual rainfall against the (first principal co-ordinates score)3 for the ‘Diet Only’ data set, for modern communities including those from Africa. Colours represent ranges of rainfall: 100 – 1 000 mm (red cross); 1 000 – 2 000 mm (blue squares); 2 000 – 4 000 mm (green diamonds).

0.4 gradient = 0.012 y-intercept = -0.422 Pearson’s r = 0.77 0.3 p(uncorr) = 1.82x10-11

0.2 95% on gradient: 9.96x10-3; 0.014

0.1 95% on y-intercept: -0.509; -0.352 0

PCO score 1 score PCO -0.1

-0.2

-0.3

10 20 30 40 50 60 70 Square root of average annual rainfall (mm)

Figure 5.3.10. Simple regression of square root of average annual rainfall against the first principal co- ordinates score for the ‘Diet and Locomotion’ data set, for modern communities excluding those from Africa. Colours represent ranges of rainfall: 100 – 1 000 mm (red cross); 1 000 – 2 000 mm (blue squares); 2 000 – 4 000 mm (green diamonds).

151 CHAPTER 5

modern sites excluding Africa (square root of rainfall) = 0.89 (Zskewness values under

+/- 2.58 indicate normal distribution at the 0.01 probability level).

Table 5.3.9 shows the predicted annual rainfall for the five Riversleigh study sites based on these regressions. Additionally, an attempt was made to determine the seasonality of rainfall at the five sites. Figure 5.3.11 illustrates the percentage of modern localities with relatively even rainfall throughout the year within given ranges of rainfall. It is apparent that the higher the overall rainfall for a locality, the less likely that the locality is subject to seasonal variations in rainfall. Given the predicted annual rainfall ranges for the five Riversleigh fossil sites, it seems plausible that the Camel Sputum palaeocommunity did not experience large seasonal fluctuations in rainfall. The Mike’s

Menagerie and Quantum Leap Site palaeocommunities fall within the 2 000 – 3 000 mm rainfall range for the Diet Only data set analysis, and in the 1 000 – 2 000 mm rainfall range for the Diet and Locomotion data set analysis; the Ringtail Site palaeocommunity is within the 1 000 – 2 000 mm rainfall range for the Diet Only data set analysis. It is difficult to speculate on the seasonality experienced by these palaeocommunities, but they are towards the upper limit of the 1 000 – 2 000 mm range and therefore, perhaps experienced mildly seasonal climates. The Encore Site palaeocommunity, with the lowest predicted rainfall (1 470 – 1 690 mm per year), is likely to have had a more pronounced seasonal pattern.

152 CHAPTER 5

Diet Only Diet and Locomotion SITE Predicted Range (95% confidence) Predicted Range (95% confidence) CS 2515 1359 – 4705 2649 1435 – 4894 MM 3095 1718 – 5721 1853 954 – 3593 QL 2108 1111 – 4075 1786 914 – 3482 EC 1468 728 – 2982 1687 855 – 3317 RG 1891 980 – 3708 - -

Table 5.3.9. Estimated annual rainfall (mm) for five Riversleigh fossil communities based on simple linear regressions on the first principal co-ordinates score on the data sets ‘Diet Only’ and ‘Diet and Locomotion’. Abbreviations: CS = Camel Sputum; MM = Mike’s Menagerie; QL = Quantum Leap; EC = Encore; RG = Ringtail.

0.9 0.8

0.7 0.6

Seasonality 0.5 0.4 13%

Seasonality 0.3 19% 0.2 69% 0.1 0

1000 2000 3000 Rainfall

Figure 5.3.11. Percentage of modern habitats with even seasonality within specified ranges of annual rainfall. Red indicates 100 – 1 000 mm annual rainfall; blue indicates 1 000 – 2 000 mm annual rainfall; green indicates 2 000 – 4 000 mm annual rainfall. Habitats with seasonal rain are plotted along the value ‘1’ and habitats with relatively even rain throughout the year are plotted along the value ‘0’.

153 CHAPTER 5

5.4. Discussion

5.4.1. Consideration of potential bias in fossil community representation

Fossil sampling and collecting techniques were examined by Wolff (1975) in terms of how representative the resulting sample is of original community structure. Surface collecting and small bulk samples were found not to be satisfactory in representing quantitative aspects of original community structure (involving relative numbers of individuals), whereas very large bulk samples (5000 – 10 000 kg of bulk sediment) are considered potentially representative. This weight range for bulk samples obviously varies with sediment type and bone density and can not be applied as an absolute figure to the Riversleigh material. Bone density varies between and within Riversleigh fossil sites, but in any case, Camel Sputum, Encore and Ringtail Sites have been sampled extensively, with bulk weights of sediment removed from these sites being in the order of tons (A. Gillespie pers. comm. 2005). Samples from Quantum Leap and Mike’s

Menagerie Sites are estimated to be under one ton.

According to Wolff’s (1975) study, 72% of the mammalian taxa contained within a site may be obtained from surface collecting, but this may represent only 36% of the total number of taxa expected to have lived there. Bulk sediment processing may increase the yield to about 55% (Wolff 1975). These figures may broadly apply to the Riversleigh study sites and indicate that the Quantum Leap and Mike’s Menagerie Site local faunas should be interpreted with caution. Further to this, specimen and species numbers recovered and described from Quantum Leap, Mike’s Menagerie and Ringtail Sites also indicate that interpretations of these fossil faunas should be made with caution (just over 1 000 specimens recovered from Quantum Leap and under

154 CHAPTER 5

1 000 specimens recovered from Mike’s Menagerie and Ringtail Sites; 9, 16 and 14 species identified from Quantum Leap, Mike’s Menagerie and Ringtail Sites respectively).

Mares and Willig (1994) examine the extent to which incomplete sampling of several non-volant mammal faunas gives rise to uncertainty concerning the habitat description of each fauna. They found that, except for tropical grassland habitats, over 75% of the fauna needed to be sampled to correctly associate a site with its actual biome; for tropical grasslands, only 33% of the fauna needed to be sampled. It must be noted that the study of Mares and Willig (1994) refers only to species lists and not to adaptive structure deduced from these lists. Given that the present study deals only with proportional distribution of species among categories of ecological diversity and not with species presence as such, and that the proportion of the fauna represented by the sample from each Riversleigh site is unknown, these figures are not particularly informative.

Results obtained for the five sites indicate that relative abundance of individuals within species is not representative (very likely due to many specimens not having been identified to species level); however, all five sites may be potentially representative in terms of adaptive structure as represented by species presence. Animals of lower body mass and, therefore, smaller size are more reliable as indicators of habitat compared to larger animals, because they are more restricted spatially (Mares and Willg 1994, Soligo

2002). Small species (0 – 10 kg body mass range) are not under-represented for any of the fossil sites studied (Fig. 5.3.2) and it is therefore presumed that local palaeoenvironmental conditions are reliably represented at these sites.

155 CHAPTER 5

Relative abundance of individuals could not be incorporated into the palaeoecological analyses undertaken in this study. The implication of this is less refinement of the palaeoecological questions addressed. Characterisation of communities through adaptive structure alone may indirectly reflect some aspects of ecosystems, thus independent information about the physical environment should also be considered

(Damuth 1992). Such independent information about climate and vegetation in

Australia through the Tertiary is presented in Chapter 6.

5.4.2. Habitat descriptions based on modern localities

The habitat types formed by cluster analysis of modern localities may be similar or equivalent between the two data sets used. To facilitate interpretation of the results an attempt is made to group similar habitat types from the two data sets into ‘combined habitat type’ groups (Table 5.4.1). Looking at distribution of modern localities among broad habitat types for the Diet and Locomotion data set (Fig. 5.3.5), it is apparent that the GRASS/DESERT category incorporates most of the localities, except those in

Africa, from OPEN 1 and OPEN 2 categories in the Diet Only data set (Fig. 5.3.3).

Three FOREST 1 localities from the Diet Only data set (Chamela, Upper Montane and

Tas H) are also included within the GRASS/DESERT category, but the remaining six

FOREST 1 localities are incorporated into the TEMPERATE category (of the Diet and

Locomotion data set). GRASS/DESERT, OPEN 1 and OPEN 2 habitat types are grouped together into G1 (a combined habitat type from both data sets). TEMPERATE and FOREST 1 habitat types are grouped together into G2.

The MIX category from the Diet and Locomotion data set includes three FOREST 2 localities (Belem and Masaguarel are included because the multiple discriminant

156 CHAPTER 5

analysis reclassifies them as FOREST 2 localities) and one OPEN 2 locality

(Transitional). Because this is a small cluster for the Diet and Locomotion data set and is dominated by FOREST 2 localities, it is considered equivalent to FOREST 2 habitat type, along with DENSE FOREST and MIXED FOREST categories which are also dominated by FOREST 2 localities. Therefore, FOREST 2, MIX, MIXED FOREST and

DENSE FOREST categories are grouped together into the combined habitat type G3.

The FOREST 3 category from the Diet Only data set includes only forest localities from

Africa (V.GipRW is reclassified by the multiple discriminant analysis as a FOREST 2 locality). This category has no equivalent in the Diet and Locomotion data set and will continue to be referred to as FOREST 3.

Combined habitat Annual rainfall Data set Broad habitat type type range (mm) D+L GRASS/DESERT D OPEN 1 G1 100-1200 D OPEN 2 D+L TEMPERATE G2 300-1250 D FOREST 1 D+L MIX D+L MIXED FOREST G3 800-4000 D+L DENSE FOREST D FOREST 2 D FOREST 3 FOREST 3 1000-1800

Table 5.4.1. Combined habitat types for the analyses of Diet and Locomotion data set (D+L) and Diet Only data set (D). Localities reclassified by the multiple discriminant analyses are included in the habitat type indicated by the results of these analyses.

Modern localities that do not cluster consistently between the two data set analyses and therefore are difficult to assign to a combined habitat type group include Chamela,

Upper Montane and Tas H (G1 or G2), Transitional (G1 or G3), and V.GipCH,

V.GipHW and V.GipSGH (G1 or G3).

157 CHAPTER 5

Chamela, Upper Montane and Tas H cluster within the FOREST 1 category in the analysis of the Diet Only data set and within the GRASS/DESERT category in the analysis of the Diet and Locomotion data set. It is interesting to note that these three localities have the highest annual rainfall within the GRASS/DESERT category. For the

Diet and Locomotion data set analysis, the GRASS/DESERT category is spatially close to the TEMPERATE category, with some overlap between the two (Fig. 5.3.6). The multiple discriminant analysis distinguishes these two categories on the first and second variates (Fig. 5.3.8. A). The variables that contribute most to this separation are

Browser, Omnivore and Arboreal. Chamela and Upper Montane have no arboreal species and a low proportion of browsers, and Tas H and Upper Montane have a low proportion of omnivores. These factors contribute to the grouping of these sites in the

GRASS/DESERT broad habitat type. For the Diet Only data set analysis, variable means are very similar between the FOREST 1 and OPEN 2 broad habitat types, except for the Insectivore and Carnivore variables. The OPEN 2 category has a higher proportion of insectivorous and a lower proportion of carnivorous mammals compared to FOREST 1. Tas H, Upper Montane and Chamela all have two or more times the number of carnivorous species relative to insectivorous species. This factor contributes to the position of the localities within the FOREST 1 cluster.

The Transitional locality clusters within the OPEN 2 category in the analysis of the Diet

Only data set and within the MIX category in the analysis of the Diet and Locomotion data set. Ojeda and Mares (1989) describe this locality as subtropical moist forest with

20-30 m tall trees, so its position within the OPEN 2 category is an unexpected result.

158 CHAPTER 5

Further information about the locality is required to explain this inconsistency, such as extent of habitat disturbance and proximity to areas with different vegetation structure.

V.GipCH, V.GipHW and V.GipSGH cluster within the OPEN 2 category in the analysis of the Diet Only data set and within the MIXED FOREST category in the analysis of the Diet and Locomotion data set (V.GipCH is reclassified by the multiple discriminant analysis into the MIX category). The vegetation at these localities is described by Conn

(1993) as mostly treeless heathland for V.GipCH, grass and herbland with low shrubs for V.Gip.SGH, and heathland/woodland in salt-influenced conditions around estuaries with a diverse shrub-layer understorey for V.GipHW. These three vegetation types cover a small area, are adjacent to one another and are also in close proximity (within

20 km) to the dry lowland vegetation type, V.GipDLF. This vegetation type is described by Conn (1993) as forest dominated by species of Eucalyptus with a canopy height of

20-37 m, a shrub layer consisting of various species, and a sparse ground layer. The discrepancy in the clustering of these three localities may be due to their restricted area and their proximity to a more densely forested region, which may allow for movement of animals between the vegetation types.

G1 and G2 combined habitat types have a similar range of rainfall, roughly

100 – 1 200 mm annual average. The broad habitat types of G1 consist predominantly of grassland, bushland/shrubland, and open woodland with ground cover. The

Australian localities, Lawn Hill and V.GrGW are grassy woodlands. The Lawn Hill vegetation includes riverine forest (tall, shaded gallery forest on the river’s edge), but consists mostly of grassy and spinifex open woodland and open grassland (O’Keefe

1996). V.GrGW is in the Grampians where the climate is mild and temperate. The

159 CHAPTER 5

vegetation of V.GrGW includes species of Eucalyptus and Allocasuarina, a shrub layer of various species, and a mainly herbaceous ground layer (Conn 1993). African localities within G1 include six vegetation types described by White (1983) as: woodland (light, open canopy and ground layer of grasses); closed woodland (closed canopy and poorly developed grass layer; may be part of forest/woodland ecotone); bushland (40% or more cover of bushes between 3-7 m tall with grasses usually present); scrubland (dominated by shrubs 10 cm – 2 m in height); grassland (grasses dominate, herbs are present and often associated in mosaic with lightly wooded communities); and desert (arid regions with sparse plant cover). White (1983) points out that transition towards arid regions is usually gradual and ‘desert’ cannot be defined precisely. The Namib desert of Africa is floristically diverse and not well differentiated from its surrounding semi-desert vegetation, which includes grassland and shrubland

(White 1983). This is not unlike the central Australian locality which includes diverse vegetation of open woodland with grass or shrub understory or shrubland or grassland

(Yeatman 1992). The North American locality included within G1 is the structurally simple tundra in Alaska. This consists of boreal or coniferous forests that become more open, with shrubs, sedges and grasses along the northern margins of the tundra

(Packham et al. 1992). Five Argentinian localities are also included within the G1 combined habitat type. These are Puna, Prepuna, Chaco, Montane Chaco and Monte.

The Prepuna vegetation consists of cacti, shrubs and bromeliads on hillsides; the Puna vegetation consists of shrub steppe with one metre tall bushes, and at higher elevations of a low grassland with low shrubs (Ojeda and Mares 1989). Chaco, Montane Chaco and Monte vegetation consists of spiny trees, shrubs and cacti (Ojeda and Mares 1989).

160 CHAPTER 5

The broad habitat types of G2 consist of three North American, one South American and two Australian localities. The North American Wsp, Dbm and Vhf localities consist of white spruce-paper birch, virgin hemlock-hardwood, and deciduous beech-maple forests respectively (Fleming 1973). The Wsp locality is within evergreen boreal coniferous forest (taiga) and open coniferous woodland vegetation, and the Dbm and

Vhf localities are within cold-deciduous broadleaved forests (Packham et al. 1992).

There are prominent herb and shrub layers during spring in areas where deciduous trees predominate within these temperate deciduous forests (Packham et al. 1992). The two

Australian localities are TasS and TasR, both temperate with mild winters and cool summers (Williams 1974). TasR consists of variable vegetation, dominated by

Nothofagus, Atherosperma, Eucryphia, Phyllocladus and Anodopetalum, with an understorey that may or may not be densely vegetated (Green 1974). TasS ranges from dense wet sclerophyll to dry, semi-open forest, dominated by species of Eucalyptus, and understorey presence dependent on moisture and fertility (Green 1974). The temperate rainforests of Tasmania have lower tree species diversity than tropical or subtropical rainforests (Packham et al. 1992). The Argentinian locality, Lower Montane, is a partially deciduous, dense forest with a closed canopy (Ojeda and Mares 1989). This locality occurs at considerably lower latitude than the other localities within the G2 combined habitat type, but Lower Montane is in northwestern Argentina, which is described by Ojeda and Mares (1989) as the interface between tropical and temperate climates. The Lower Montane locality may have a reduction of some tropical elements as a result of impoverished vegetation and increased seasonality, and possibly, the effects of habitat modification (Ojeda and Mares 1989).

161 CHAPTER 5

The G3 and FOREST 3 combined habitat types overlap in annual rainfall range, with

G3 having a very broad range (800-4000 mm), and FOREST 3 having a more restricted range (1000-1800mm). The African localities in the FOREST 3 habitat type are all referred to as ‘forest’ habitats by Reed (1998). White (1983) describes African forests as continuous stands of trees, with a canopy of 10-50 m or more, with several layers and overlapping crowns. He lists several types of vegetation within this broad category: rainforest, dry forest, semi-evergreen forest, deciduous forest and undifferentiated forest, but warns that these vegetation descriptions are often applied imprecisely or in conflicting senses.

A point worth mentioning is that localities from Africa have a tendency to cluster together within broad habitat types. This is particularly striking for the FOREST 3 habitat type, which consists only of African forest localities. One explanation for this is that Australia and South America have more marked affinities (Keast 1972a). African faunas differ from both South American and Australian faunas in having a greater diversity of medium-sized to large terrestrial herbivores, and consequently, a greater diversity of large carnivores (Keast 1972b). Both South America and Australia are depauperate in their original (i.e. pre 20 000 years ago) megafauna. Community species composition, diversity and ecological adaptive structure are largely determined by geographical, evolutionary and historical factors (Ricklefs and Schluter 1993), in addition to the physical environment. Isolating the physical environment from evolutionary history as a cause of differences between faunas from different continents is possible to some extent. Westoby (1993: 171) suggests that “if a difference between continents is due to the physical environment, a similar difference in the physical environment within a continent ought to have the same effect on diversity” and

162 CHAPTER 5

presumably on adaptive structure. For example, the difference between Australian

FOREST 2 sites and African OPEN 1 sites is comparable to the difference between

Australian FOREST 2 sites and Australian OPEN 1 sites. This is seen only for considerably different habitats, or physical environments, in the current study. The resemblance of faunas from similar vegetation types within Africa is greater than that of faunas from similar vegetation types between Africa and other continents. The implication of this is that a cluster consisting of localities from different continents indicates that these localities do share features of the physical environment to a large extent. However, a cluster consisting of localities from one continent only, although indicating that the localities do have features of the physical environment in common, could potentially be based more on geographical, evolutionary and historical factors.

The similarities of the physical environment between the localities may therefore be very general and this imposes limits to the detail with which these features can be described (as seen above for African FOREST 3 sites).

The broad habitat types constituting the G3 combined habitat type are variable in vegetation, which reflects the wide range of annual rainfall. DENSE FOREST includes heavily vegetated tropical localities less than 20o from the equator. The South American localities contained within this habitat type are all described as tropical rainforest

(Medellin 1994), except for Guatapo which is a broken canopy riverine Ilanos; the

Australian, Central American and New Guinea localities are tropical or subtropical wet forests.

MIXED FOREST includes localities of variable vegetation cover. Some of these localities have heterogeneous habitats (Ku-Ring-Gai, Jervis and Lamington National

163 CHAPTER 5

Parks, and South Coast and Cape York), that include heath, grassland, woodland and rainforest vegetation types within them. Others, such as V.GipWLF, may be ecotonal or transitional between other forest types, or may be adjacent to different vegetation types, allowing for significant faunal interchange. Heterogeneous or vegetationally mixed habitats contain diverse faunas with a great variety of adaptations (Williams et al.

1996). This may explain the clustering of communities from vegetationally heterogeneous habitats with the very diverse faunas of rainforest habitats in the

FOREST 2 broad habitat type of the Diet Only analysis. Rosenzweig and Abramsky

(1993: 57) describe the relationship between diversity and productivity, where diversity increases with productivity up to a point (as a result of a rise in resource heterogeneity), then declines (because “productivity tends to be spread more evenly within and between years, reducing the variety of viable temporal specialisations”), resulting in a hump- shaped pattern. Such a relationship would explain comparable diversity patterns between rainforest, or densely vegetated habitats and more structurally mixed habitats.

The analysis based on the Diet and Locomotion data set is more sensitive in distinguishing the dense forests from the mixed forests than the analysis based on the

Diet Only data set. Often heterogeneous habitats may be dominated by one type of vegetation structure, for example, the landscape of Lawn Hill National Park is dominated by grassland. The dominant vegetation structure is reflected in the clustering of these heterogeneous localities using ecological diversity analyses of mammalian faunas. In other cases, the mammalian faunas of homogenous habitats are affected by adjacent habitats, resulting in a seemingly mixed/heterogeneous profile (as for many of the Victorian sites in Gippsland). Cale and Hobbs (1994) point out that landscape heterogeneity occurs over many scales (e.g. patch, landscape and regional).

Heterogeneity, as it applies here, refers not only to variations in vegetation structure

164 CHAPTER 5

within a locality, but also to interactions with adjacent habitats that differ in vegetation structure.

The MIX broad habitat type includes Masaguaral, a riverine Ilanos (Medellin 1994), which is described as subtropical vegetational mosaic with savannah by Keast (1972b);

Belem, a tropical rainforest which is now urban and suburban (Kay and Madden 1997 and Pine 1973, cited within); and Mt Azules, a lowland tropical rainforest (Medellin

1994) that clusters within FOREST 2 in the Diet Only data set analysis. G3 is dominated by localities that have tropical rainforest elements or are in close proximity to such vegetation types.

5.4.3. Determining habitat types and annual rainfall for fossil communities

The ecological adaptive structure of modern mammalian communities has been related to the general vegetation structure of their habitats. On this basis, fossil mammalian communities from the five Riversleigh study sites are grouped into habitat types as follows: Camel Sputum, Mike’s Menagerie and Ringtail Sites represent G3 habitats;

Encore Site represents a FOREST 3 habitat; and Quantum Leap Site represents a G1 habitat. This is based on the Diet Only data set. For the Diet and Locomotion data set, fossil sites cannot be grouped into broad habitat types because their Mahalanobis distances from the group centroids are too large. This indicates either, palaeocommunity structure very different to modern community structure (which is not likely because such differences are not apparent in dietary adaptive structure alone), or the effect of certain biases in preservation on the apparent locomotor adaptive structure of the fossil communities (foot and hand bones occur in generally lower numbers than expected for the study sites – see Chapter 2).

165 CHAPTER 5

Camel Sputum and Mike’s Menagerie Sites are interpreted here to represent heavily forested habitats with tropical rainforest elements. This interpretation is supported by the fact that their lowest Mahalanobis distances are from the DENSE FOREST broad habitat type in the Diet and Locomotion data set analysis. Also, estimates of annual rainfall at these sites support this interpretation. Using estimates from both the Diet

Only and the Diet and Locomotion data sets, the range for Camel Sputum Site is approximately 2 575 – 2 650 mm of annual rainfall, and the range for Mike’s Menagerie

Site is approximately 1 780 – 3 000 mm of annual rainfall. These values are well within the annual rainfall range for G3 habitats and, in fact, are within the range for the

DENSE FOREST habitat type (>2 000 mm).

Ringtail Site may also be interpreted as a densely forested palaeohabitat, however, the only rainfall estimate available is from the Diet Only data set (approximately

1 890 mm annual rainfall). This value is close to the lower limit for the DENSE

FOREST broad habitat type. Further support for this interpretation comes from the vegetation depicted by Arena (in prep) for the Riversleigh middle Miocene. Variations in landform and topography that appear in karst terrain have an effect on the distribution of vegetation. Arena (2004) presents the karst environment mosaic model of vegetation and soil distribution. He describes a humid, wet environment with thick soil and continuous vegetation cover in the ‘lower’ parts of the terrain, and discontinuous soil cover with drier surface conditions, exposed rock surfaces and drier vegetation in the

‘upper’ parts. His suggestion is that these ‘lower’ parts of the terrain expanded through denudation during the early Miocene and that the dense vegetation associated with them became dominant in the landscape by the middle Miocene.

166 CHAPTER 5

Encore Site, as a FOREST 3 habitat, would have potentially resembled the forested habitats of modern African localities. It is difficult to describe the possible structure of the vegetation because the African localities that define this habitat category are quite varied, as explained above. However, according to White’s (1983) general description of African forests, the Encore Site palaeovegetation structure may be described as continuous stands of trees with overlapping crowns and a canopy more than ten metres high. Estimated annual rainfall for Encore Site is approximately 1 470 – 1 690 mm. This falls within the range of annual rainfall for the FOREST 3 habitat type. The FOREST 3 habitat type includes dry forest and deciduous forest vegetation and this is reflected in the generally lower annual rainfall bracket compared with the G3 habitat type. It is therefore postulated that the Encore Site palaeohabitat was generally drier and possibly more open relative to the Camel Sputum, Mike’s Menagerie and Ringtail Site palaeohabitats. Woodland, mixed with closed forest components, seems a likely palaeohabitat for Encore Site based on parallels drawn from Van Couvering (1999) for

East Africa. A shift from forest domination towards a drier, seasonal woodland community is apparent in East Africa in the middle Miocene from the ecological diversity structure of the fauna, which is dominated by browsing herbivores. Browsers dominate the Encore Site local fauna, representing over 63% of the species. This dominance of browsers is an interesting phenomenon worth further investigation, especially in light of evidence presented by Janis et al. (2004). They outline potential reasons for high numbers of browsing ungulates in North American mid-Miocene communities and suggest that this was a global phenomenon due to high primary productivity. The timing of this phenomenon in Australia may be examined through

167 CHAPTER 5

palaeoecological studies of late Miocene and Pliocene sites, such as Alcoota (Northern

Territory), Bluff Downs (eastern Queensland) and Curramulka (South Australia).

The habitat grouping of Quantum Leap Site within the OPEN 2 category suggests a drastic difference in vegetation at Riversleigh between the late Oligocene and the early

Miocene. An ‘icehouse’ phase, with cool temperatures and relatively dry conditions was prevailing in the late Oligocene (McGowran and Li 1994). In the Diet and Locomotion analysis, the shortest Mahalanobis distance for Quantum Leap Site is from the MIX broad habitat type. Taking the estimated annual rainfall range for Quantum Leap Site

(approximately 1 780 – 2 100 mm) into account, it seems plausible that this site represents a mozaic palaeohabitat, in terms of vegetation, rather than an open woodland or grassland. The estimated annual rainfall is well above the 100 – 1 200 mm annual rainfall range for the G1 habitat type. Myers’ (2002) interpretation of the Riversleigh

White Hunter Site local fauna (equivalent in age to the Quantum Leap Site local fauna) as closed forest with patches of significantly drier and more open vegetation is possibly applicable to the palaeohabitat at Quantum Leap Site in the late Oligocene.

Results and interpretations presented here are broadly consistent with previous palaeoecological conclusions for Riversleigh. Archer et al. (1995) interpret the

Riversleigh Miocene palaeoenvironment as cool, wet rainforest. Tyler et al. (1990,

1994) suggest that seasonal aridity did not occur at Riversleigh throughout the Oligo-

Miocene, based on the presence of small, non-burrowing frog species. The current study indicates that the early Miocene Camel Sputum Site palaeoenvironment probably did not have a seasonal climate with respect to rainfall. Sampling at Quantum Leap and

Mike’s Menagerie Sites has been lower than for other study sites and consequently

168 CHAPTER 5

results for these local faunas need to be interpreted with caution. Based on estimated rainfall values, Quantum Leap, Mike’s Menagerie and Ringtail Sites potentially had a mildly seasonal climate. If Mike’s Menagerie Site however, is found to be part of the same deposit as Camel Sputum Site (as discussed in Chapter 2), then the pattern of seasonality should be the same as for the Camel Sputum local fauna. For the Encore Site local fauna, considered to be early late Miocene in age (Myers et al. 2000), a stronger pattern in seasonality is likely to have occurred.

5.4.4. The influence of abiotic factors on vegetation structure and animal communities

Abiotic factors that have an influence on vegetation structure and composition, and therefore, indirectly impact on animal communities, include rainfall, evaporation, soil type, light, temperature, geology, topography, landmass area and latitudinal position

(Keast 1972b, Pielou 1975, Turton and Sexton 1996, Westoby 1993, Brown et al. 1997,

Reed 1998, Andrews and O’Brien 2000). Of these, only rainfall was considered in the present study, and only to a limited extent. Rainfall was represented by a figure for average annual rainfall, thereby not taking into account the effect of variation in rainfall between years. This may be important in some cases, for instance, extensive and prolonged drought in Australia (Adam 1994), where there may be effects on vegetation and animal communities. However, even if such detail is included for modern localities, it would probably not be possible to find a correlation between this and the indirect information available from mammalian community adaptive structure, which would make it useless for application to the fossil sites under study. In any case, average annual rainfall figures are sufficient for a broad picture as they indicate a zonal pattern, which is reflected in major vegetation types and mammal species distribution (Keast

1972b).

169 CHAPTER 5

Although latitudinal gradients in species diversity are apparent today and existed in the past (Pielou 1975, Rosenzweig 1995), these trends were not examined in the present study because the confounding effects of topography, continental positions and associated oceanic currents could not be factored in. These factors, through their direct influence on climate, affect faunal communities. For example, variations in elevation and topographic relief cause changes in temperature and rainfall patterns (Andrews and

O’Brien 2000). Rainfall and seasonality, along with habitat heterogeneity at different scales, are invariably related to faunal diversity in the tropics, more so than is latitude

(Williams et al. 1996). Seasonal climatic variability results in unstable, unpredictable environments, which in turn affects availability of food resources and thus faunal richness and diversity are affected (Root 1967, Pielou 1975, Andrews and O’Brien

2000). Seasonality and unpredictability act to decrease diversity in species and in their ecological adaptations. This is because species must be highly generalised and flexible in their responses, in other words have wide niches, in order to survive in habitats that fluctuate widely and a result of greater niche width is fewer niches and therefore fewer species (Root 1967, Pielou 1975). High environmental stability, predictability and productivity, on the other hand, lead to high community stability, which permits high diversity (Pielou 1975).

Soils and geology also affect the vegetational landscape through their effects on moisture availability to plants. The availability of moisture to vegetation is dependent on annual rainfall to a large extent but other factors are also of great importance, such as intensity of rain, evaporation, surface runoff and seepage (Brown et al. 1997). These factors interact with the moisture-holding capacity of the particular soil type to

170 CHAPTER 5

determine the amount of water available to vegetation (Brown et al. 1997). The karst terrain in which Riversleigh Tertiary fossils were deposited would have had discontinuous drainage (Taylor 1994), which would have had some influence on vegetation, for instance a variable occurrence of soil moisture that would result in a mosaic of vegetation types.

Certain abiotic factors are considered, as they relate to climate, in Chapter 6. This, and other independent evidence, is used to assess the reliability of the results obtained here.

5.5. Conclusions

This study aimed to provide broad descriptions of palaeohabitats for five Riversleigh fossil sites. Trophic and locomotor ecological adaptations of the fossil mammals from these sites were related to broad aspects of vegetation structure on the basis of analogy with 84 modern mammalian communities and their physical environments.

The results of the analyses undertaken here indicate that Riversleigh Miocene palaeohabitats were forested, not open. The Quantum Leap Site local fauna (late

Oligocene in age) possibly inhabited a heterogeneous environment with structurally mixed vegetation including forest elements. Annual rainfall for the Quantum Leap palaeocommunity is estimated to have been up to, approximately 2 000 mm. Rainfall estimates are higher for the Camel Sputum and Mike’s Menagerie Site local faunas

(early Miocene in age), which most likely inhabited densely forested environments. The

Ringtail Site local fauna (middle Miocene in age) also inhabited densely forested environments. The Encore Site local fauna (early late Miocene in age) inhabited slightly drier, forested environments with a correspondingly lower annual rainfall, well under 2

171 CHAPTER 5

000 mm. The trend of slowly decreasing rainfall through the Miocene will be discussed with respect to other evidence of climatic change throughout this period in Chapter 6.

The diet and locomotor adaptive structure of mammalian communities was found to provide better resolution than the dietary adaptive structure alone, of different types of forested habitats, with the distinction of densely forested (or tropical rainforest) localities from localities with structurally mixed vegetation. Riversleigh fossil faunas were found to be closely associated with the broad habitat groups formed by the dietary adaptive structure of modern communities, but not with those formed by the diet and locomotor adaptive structure of modern communities. This may be due to: 1) a preservation bias in calcanea, with subsequent effects on the apparent locomotor adaptive structure of the palaeocommunities, or 2) the possibility that ecological characteristics may actually have been different in the past, as has been suggested for palaeocommunities from Africa (Van Couvering 1999) and North America (Janis et al.

2002). Also, Janis (1993) considers the possibility that modern communities may not represent exact equivalents for past communities, despite general similarities existing between them.

Only the mammalian component of the fossil communities has been examined here, but perhaps much more insight can be gained by the inclusion of birds, fish, reptiles, amphibians and molluscs in such palaeoecological analyses.

172 CHAPTER 5

5.6. References

Adam, P. 1994. Australian Rainforests. Oxford University Press, Oxford.

Andrews, P. 1989. Paleoecology of Laetoli. Journal of Human Evolution 18: 173-181.

Andrews, P., Lord, J. M. and Nesbit Evans, E. M., 1979. Patterns of ecological diversity in fossil and modern mammalian faunas. Biological Journal of the Linnean Society 11: 177-205.

Andrews, P. and O’Brien, E. M. 2000. Climate, vegetation, and predictable gradients in mammal species richness in southern Africa. Journal of Zoology, London 251: 205-231.

Anonymous, 1980. Atlas of NSW Wildlife (On-line), NSW Department of Environment and Conservation. Accessed May 28, 2003 at http://www.wildlifeatlas.nationalparks.nsw.gov.au/wildlifeatlas/watlasSpecies.jsp

Anonymous, 1997a. Victoria – broad vegetation types (coverage in 1987) and Victorian bioregions, 1:500 000. Map in Victoria’s Biodiversity – Directions in Management. Department of Natural Resources and Environment, Melbourne.

Anonymous, 1997b. Victoria’s Biodiversity – Directions in Management. Department of Natural Resources and Environment, Melbourne, Victoria.

Anonymous, 1998. Annual Rainfall of Australia 1997, 1:9 000 000. Map, Bureau of Meteorology, National Climate Centre, Commonwealth of Australia.

Anonymous, 2000. Lamington National Park (On-line), Green Mountains Natural History Association, The University of Queensland. Accessed Feb 14, 2003 at http://lamington.nrsm.uq.edu.au/MainMenu.html

Anonymous, 2005. Annual Cycle of Seasonal Total Rainfall for Pacific Islands, (On- line), NOAA/National Weather Service Climate Prediction Centre. Accesses May 21, 2005 at http://www.cpc.ncep.noaa.gov/pacdir/html-figdir/html4dir/mada4.html

Archer, M. 1992. Ringtail possums (Pseudocheiridae, Marsupialia) from the Tertiary deposits of Riversleigh. The Beagle, Records of the Northern Territory Museum of Arts and Sciences 9: 257.

Archer, M., Hand, S. J. and Godthelp, H. 1996. Riversleigh. Reed Books: Chatswood, NSW.

173 CHAPTER 5

Archer, M., Hand, S. J., Godthelp, H. and Creaser, P. 1997. Correlation of the Cainozoic sediments of the Riversleigh World Heritage Fossil Property, Queensland, Australia. Pp. 131-52 in Actes du Congres BiochroM’97. Ed. J. P. Aguilar, S. Legendre, J. Michaux. Mem. Trav. E.P.H.E., Inst. Montpellier.

Arena, D. A. 2004. The Geological History and Development of the Terrain at the Riversleigh World Heritage Area During the Middle Tertiary. PhD Thesis. University of New South Wales, Sydney.

Bourlière, F. 1973. The comparative ecology of rain forest mammals in Africa and tropical America: some introductory remarks. Pp. 279-292 in Tropical Forest Ecosystems in Africa and South America: A Comparative Review. Eds R. C. Meggers, E. E. Ayensu and W. D. Duckworth. Smithsonian Institution Press, Washington D. C.

Brown, A. G., Turnbull, J. W. and Booth, T. H. 1997. The Australian environment. Pp. 1-18 in Australian Trees and Shrubs: Species for Land Rehabilitation and Farm Planting in the Tropics. Eds J. C. Doran and J. W. Turnbull. Australian Centre for International Agriculture Research, Canberra.

Cale, P. G. and Hobbs, R. J. 1994. Landscape heterogeneity indices: problems of scale and applicability, with particular reference to animal habitat description. Pacific Conservation Biology 1: 183-193.

Conn, B. J. (1993). Natural regions and vegetation of Victoria. Pp. 79-158 in D. B. Foreman and N. G. Walsh (eds), Flora of Victoria, Volume 1 Introduction. Inkata Press, Melbourne.

Crosby, K. 2002. Studies in the Diversity and Evolution of Phalangeroid Possums (Marsupialia; Phalangerida; Phalangeroidea). PhD Thesis. University of New South Wales, Sydney.

D’Addario, G. W., Dow, D. B. and Swoboda, R. 1975. Geology of Papua New Guinea, 1:2500 000. Canberra, Bureau of Mineral Resources, Australia.

Damuth, J. 1982. Analysis of the preservation of community structure in assemblages of fossil mammals. Paleobiology 8: 434-446.

Damuth, J. D. 1992. Taxon-free characterization of animal communities. Pp. 183-203 in Terrestrial ecosystems through time. Eds A. K. Behrensmeyer, J. D. Damuth, W. A. Di Michele, R. Polts, H. D. Sues and S. L. Wing. The University of Chicago Press, Chicago.

De Bruine, L. 2002. Saguinus fuscicollis (On-line), Animal Diversity Web. Accessed May 23, 2005 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Saguinus fuscicollis.html

174 CHAPTER 5

Eisenberg, J. F., O’Connell, M. A. and August, P. V. 1979. Density, productivity, and distribution of mammals in two Venezuelan habitats. Pp. 187-207 in Vertebrate Ecology in the Northern Neotropics. Ed J. F. Eisenberg. Smithsonian Institution Press, Washington, D.C.

Eisenberg, J. F. and Thorington, R. W. Jr. 1973. A preliminary analysis of a neotropical mammal fauna. Biotropica 5 (3): 150-161.

Field, J. G., Clarke, K. R. and Warwick, R. M. 1982. A practical strategy for analysing multispecies distribution patterns. Marine Ecology Progress Series 8: 37-52.

Flannery, T. 1995. Mammals of New Guinea. Reed Books, Australia.

Fleming, T. H. 1973. Numbers of mammal species in North and Central American forest communities. Ecology 54: 555-563.

Gardner, J. L. (project editor) 1995. Reader’s Digest Atlas of the World. Toucan Books Limited, London.

Gentry, A. H. 1988. Tree species richness of upper Amazonian forests Proceedings of the Natural Academy of Science 85: 156-159.

Glanz, W. E. 1982. Adaptive zones of neotropical mammals: a comparison of some temperate and tropical patterns. Pp. 95-110 in Mammalian Biology in South America. Eds M. A. Mares and H. H. Genoways. Pymatuning Laboratory of Ecology, Special Publication 6, University of Pittsburgh.

Green, R. H. 1974. Mammals. Pp. 367-396 in Biogeography and Ecology in Tasmania. Ed. W. D. Williams. Dr. W. Junk b.v., Publishers, The Hague.

Hair, J. F. Jr, Anderson, R. E., Tatham, R. L. and Black, W. C. 1998. Multivariate Data Analysis. Prentice-Hall International, Inc., United States of America.

Hammer, Ø., Harper, D. A. T. and Ryan, P. D. 2004. PAST – PAlaeontological STatistics, Version 1.27. Computer program available online at http://folk.uio.no/ohammer/past

Hand, S. 1997. New Miocene leaf-nosed bats (Microchiroptera: Hipposideridae) from Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41 (2): 335-349.

Hand, S. J., Archer, M. and Godthelp, H. 2001. New Miocene Icarops material (Microchiroptera: Mystacinidae) from Australia, with a revised diagnosis of the genus. Memoirs of the Association of Australasian Palaeontologists 25: 139-146.

Hoffman, J. A. J. (director of preparation) 1975. Climatic Atlas of South America – Maps of Mean Temperature and Precipitation, 1:5 000 000. WMO, Unesco, Cartographia, Hungary.

175 CHAPTER 5

Hutchins, M., Kleiman, D. G., Geist, V. and McDade, M. C. (eds) 2003. Grzimek's Animal Life Encyclopedia, 2nd Edition, Volumes 12-16. Mammals I-V. Gale Group, Farmington Hills, Michigan.

Hutchinson, G. E. 1978. An Introduction to Population Ecology. Yale University Press, New Haven, Connecticut.

Jablonski, D. and Sepkoski, J. J. Jr. 1996. Paleobiology, community ecology, and scales of ecological pattern. Ecology 77: 1367-1378.

Janis, C. M. 1984. The use of fossil ungulate communities as indicators of climate and environment. Pp. 85-104 in Fossils and Climate. Ed. P. J. Brenchley. John Wiley and Sons Ltd, Chichester.

Janis, C. M. 1993. Tertiary mammal evolution in the context of changing climates, vegetation, and tectonic events. Annual Review of Ecology and Systematics 24: 467-500.

Janis, C. M., Damuth, J. and Theodor, J. M. 2002. The origins and evolution of the North American grassland biome: the story from the hoofed mammals. Palaeogeography, Palaeoclimatology, Palaeoecology 177: 183-198.

Janis, C. M., Damuth, J. and Theodor, J. M. 2004. The species richness of Miocene browsers, and implications for habitat type and primary productivity in the North American grassland biome. Palaeogeography, Palaeoclimatology, Palaeoecology 207: 371-398.

Karr, J. R. and James, F. C. 1975. Ecomorphological configurations and convergent evolution in species and communities. Pp. 258-291 in Ecology and Evolution of Communities. Eds M. L. Cody and J. M. Diamond. Belknap Press of Harvard University Press, Cambridge.

Kay, R. F. and Madden, R. H. 1997. Mammals and rainfall: paleoecology of the middle Miocene at La Venta (Colombia, South America). Journal of Human Evolution 32: 161- 199.

Keast, A. 1969. The evolution of mammals on southern continents, VII: comparisons of the contemporary mammalian faunas of the southern continents. Quarterly Review of Biology 44: 121-167.

Keast, A. 1972a. Continental drift and the biota of the mammals on southern continents. Pp. 23-87 in Evolution, Mammals, and Southern Continents. Eds A. Keast, F. C. Erk and B. Glass. State University of New York Press, Albany.

Keast, A. 1972b. Comparisons of contemporary mammal faunas of southern continents. Pp. 433-501 in Evolution, Mammals, and Southern Continents. Eds A. Keast, F. C. Erk and B. Glass. State University of New York Press, Albany.

Koch, C. F. 1978. Bias in the published fossil record. Paleobiology 4: 367-372.

176 CHAPTER 5

Koch, C. F. 1987. Prediction of sample size effects on the measured temporal and geographic distribution patterns of species. Paleobiology 13: 100-107.

Lee, K. A. and Cockburn, A. 1985. Evolutionary Ecology of Marsupials. Cambridge University Press, Cambridge.

Macdonald, D. (ed), 2001. The New Encyclopedia of Mammals. Oxford University Press, Oxford.

McGowran, B. and Li, Q. 1994. The Miocene oscillation in southern Australia. Records of the South Australian Museum 27 (2): 197-212.

Magurran, A. E. 1988. Ecological Diversity and its Measurement. Princeton University Press, Princeton, New Jersey.

Mares, M. A. and Willig, M R. 1994. Inferring biome associations of recent mammals from samples of temperate and tropical faunas: paleoecological considerations. Historical Biology 8: 31-48.

Medellin, R. A. 1994. Mammal diversity and conservation in the Selva Lacandona, Chiapas, Mexico. Conservation Biology. 8: 780-799.

Menkhorst, P. W. (ed.), 1995. Mammals of Victoria – Distribution, Ecology and Conservation. Oxford University Press, Melbourne.

Muirhead, J. 1994. Systematics, Evolution and Palaeobiology of Recent and Fossil Bandicoots (Peramelemorphia, Marsupialia). PhD Thesis. University of New South Wales, Sydney.

Myers, T., Crosby, K., Archer, M. and Tyler, M. 2000. The Encore Local Fauna, a late Miocene assemblage from Riversleigh, northwestern Queensland. Memoirs of the Association of Australasian Palaeontologists 25: 147-154.

Myers, T. J. 2001. Prediction of marsupial body mass. Australian Journal of Zoology 49: 99-118.

Myers, T. J. 2002. Palaeoecology of Oligo-Miocene Local Faunas from Riversleigh. PhD Thesis. University of New South Wales, Sydney.

Nesbit-Evans, E. M., Van Couvering, J. A. H., Andrews, P. 1980. Paleoecology of Miocene sites in western Kenya. Journal of Human Evolution 10: 99-116.

Nowak, R. M. and Paradiso, J. L. 1983. Walker’s Mammals of the World, 4th Edition. Johns Hopkins University Press, Baltimore.

Ojeda, A. R. and Mares, M. A. 1989. A Biogeographic Analysis of the Mammals of Salta Province, Argentina – Patterns of Species Assemblage in the Neotropics. Special Publications, The Museum Texas Tech University, No. 27, Texas.

177 CHAPTER 5

O’Keefe, C. 1996. Lawn Hill National Park. Robert Brown and Associates Pty Ltd, Queensland.

Packham, J. R., Harding, D. J. L., Hilton, G. M. and Stuttard, R. A.1992. Functional Ecology of Woodlands and Forests. Chapman and Hall, London.

Paijmans, K. 1975. Vegetation of Papua New Guinea, 1 : 1 000 000. Map, Division of Land Use Research, Commonwealth Scientific and Industrial Research Organisation.

Pielou, E. C. 1975. Ecological Diversity. John Wiley and Sons Inc., New York.

Ray, A., Fitzherbert, S. and Jehne, R. 1996. Kuring-gai Chase National Park. National Parks and Wildlife Service, Wentworth Press, Australia.

Redford, K. H. and Eisenberg, J. F. 1992. Mammals of the Neotropics - the Southern Cone. The University of Chicago Press, Chicago.

Reed, K. E. 1998. Using large mammal communities to examine ecological and taxonomic structure and predict vegetation in extant and extinct assemblages. Paleobiology 24: 384-408.

Ricklefs, R. E. and Schluter, D. 1993. Species diversity: regional and historical influences. Pp. 350-363 in Species Diversity in Ecological Communities – Historical and Geographical Perspectives. Eds R. E. Ricklefs and D. Schluter. The University of Chicago Press, Chicago.

Root, B. R. 1967. The niche exploitation pattern of the blue-gray gnatcatcher. Ecological Monographs 37: 317-350.

Rosenzweig, M. L. 1995. Species Diversity in Space and Time. Cambridge University Press, Cambridge.

Rosenzweig, M. L. and Abramsky, Z. 1993. How are diversity and productivity related? Pp. 52-65 in Species Diversity in Ecological Communities – Historical and Geographical Perspectives. Eds R. E. Ricklefs and D. Schluter. The University of Chicago Press, Chicago.

Sheppe, W. and Osborne, T. 1971. Patterns of use of a flood plain by Zambian mammals. Ecological Monographs 41 (3): 179-205.

Sneath, P. H. A. and Sokal, R. R. 1973. Numerical Taxonomy – the Principles and Practice of Numerical Classification. W. H. Freeman and Company, San Francisco.

Soligo, C. 2002. Primatology, paleoecology, and a new method for assessing taphonomic bias in fossil assemblages. Evolutionary Anthropology Suppl 1: 24-27.

Steinhauser, F. (technical supervisor) 1979. Climatic Atlas of North and Central America – Maps of Mean Temperature and Precipitation, 1:5 000 000. WMO, Unesco, Cartgraphia, Hungary.

178 CHAPTER 5

Strahan, R. (ed) 1998. The Mammals of Australia. New Holland Publishers Pty Ltd, Sydney.

SYSTAT – Version 7.0 for Windows. 1997. SPSS Inc., Chicago (Computer program).

Taylor, G. 1994. Landscapes of Australia: their nature and evolution. Pp. 60-79 in History of the Australian Vegetation: Cretaceous to Recent. Ed. R. S. Hill. Cambridge University Press, Cambridge.

Teeter, K. 2000. Pappogeomys castanops (On-line), Animal Diversity Web. Accessed May 21, 2005 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Pappogeomys_castanops.html

Tracey, J. G. and Webb, L. J. 1975. The Vegetation of the Humid Tropical Region of North Queensland (Maps and Key). CSIRO, Indooroopilly, Queensland.

Turton, S. M. and Sexton, G. J. 1996. Environmental gradients across four rainforest- open forest boundaries in northeastern Queensland. Australian Journal of Ecology 21: 245-254.

Tyler, M., Godthelp, H. and Archer, M. 1994. An overview of the Riversleigh frog fauna. Riversleigh Symposium 1994: Program and Abstracts, University of New South Wales, Sydney.

Tyler, M., Hand, S. and Ward, V. 1990. Analysis of the frequency of Lechriodus intergerivus Tyler (Anura: Leptodactylidae) in Oligo-Miocene local faunas of Riversleigh Station, Queensland. Proceedings of the Linnean Society of New South Wales 112:105-109.

Van Couvering, J. A. H. 1999. Community evolution in east Africa during the late Cenozoic. Pp. 272-298 in African Biogeography, Climate Change, and Human Evolution. Eds T. G. Bromage and F. Schrenk. Oxford University Press, New York.

Vandermeer, J. H. 1972. Niche theory. Annual Review of Ecology and Systematics 3: 107-132.

Vieira, E. M., Pizo, M. A., Izar, P. (2003). Fruit and seed exploitation by small rodents of the Brazilian Atlantic forest. Mammalia 67(4): pp?

Watts, C. H. S. 1977. The foods eaten by some Australian Rodents (Muridae). Australian Widlife Research 4: 151-158.

Watts, C. H. S. and Aslin, H. J. 1981.The Rodents of Australia. Angus and Robertson Publishers, London.

Westoby, M. 1993. Biodiversity in Australia compared with other continents. Pp. 170- 177 in Species Diversity in Ecological Communities – Historical and Geographical Perspectives. Eds R. E. Ricklefs and D. Schluter. The University of Chicago Press, Chicago.

179 CHAPTER 5

White, F. 1983. The Vegetation of Africa – a Descriptive Memoir to Accompany the Unesco/AETFAT/UNSO Vegetation Map of Africa. United Nations Educational, Scientific and Cultural Organization, Paris.

Whittaker, R. H. 1975. Communities and Ecosystems. Macmillan Publishing Co., Inc., New York.

Whittaker, R. H. and Levin, S. A. (Eds). 1975. Niche: Theory and Application. Benchmark Papers in Ecology 3, Halted Press, Pennsylvania.

Whittaker, R. H., Levin, S. A. and Root, R. B. 1973. Niche, habitat and ecotope. American Naturalist 107: 321-338.

Williams, W. D. 1974. Introduction. Pp.3-16 in Biogeography and Ecology in Tasmania. Ed. W. D. Williams. Dr. W. Junk b.v., Publishers, The Hague.

Williams, S. E., Pearson, R. G., and Walsh, P. J. 1996. Distribution and biodiversity of the terrestrial vertebrates of Australia’s wet tropics: a review of current knowledge. Pacific Conservation Biology 2:327-362.

Wing, S. L., Sues, H. S., Potts, R., DiMichele, W. A. and Behrensmeyer, A. K. 1992. Evolutionary paleoecology. Pp. 1-13 in Terrestrial Ecosystems Through Time – Evolutionary Paleoecology of Terrestrial Plants and Animals. Eds A. K. Behrensmeyer, J. D. Damuth, W. A. DiMichele, R. Potts, H. D. Sues, S. L. Wing. The University of Chicago Press, Chicago.

Winter, J. W. and Allison, F. R. 1980. The native mammals of Cape York Peninsula – changes in status since the 1948 Archbold expedition. Pp. 31-44 in Contemporary Cape York Peninsula, eds N. C. Stevens and A. Bailey. The Royal Society of Queensland, Brisbane.

Witt, E. 2000. Sciurus deppei (On-line), Animal Diversity Web. Accessed May 21, 2005 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sciurus_deppei.html

Wolff, R. G. 1975. Sampling and sample size in ecological analyses of fossil mammals. Paleobiology 1: 195-204.

Woodburne, M. O., Tedford, R.H., Archer, M., Turnbull, W. D., Plane, M. D. andLundelius, E. L. 1985. Biochronology of the continental mammal record of Australia and New Guinea. Special Publications of the South Australian Deptartment of Mines and Energy 5: 347-63.

Yeatman, E. M. 1992. Fauna survey of the Lake Amadeus area, Central Australia. South Australian Nature, 66: 64-81.

180 APPENDIX C1. Species lists for Australian sites (see Chapter 5 for habitat descriptions). Dark fill indicates presence; light fill indicates uncertain but probable presence. Species lists for Victorian sites were obtained through the use of mammal distribution and vegetation maps (as described in Chapter 5); species lists for the South Coast, Jervis Bay and Ku-ring-gai Chase National Parks were obtained from Anon. (1980) and for Lamington National Park, from Anon. (2000); species list for Cape York is from Winter and Allison (1979); species list for Lawn Hill is from O'Keefe (1996); species list for Central Australia is from Yeatman (1992); species lists for Tasmania are from Green (1974); species lists for Mossman, Atherton Uplands, Lee Uplands and Ingham Lowlands are from Williams et al.(1996).

SPECIES VicGipSGHVicGipCH VicGipHW VicGipRW VicGipDLF VicGipMWFVicGipRF VicGipWLFVicGipWTRVicGipCTRVicGrHRW VicGrGW VicGrWDF VicGrOF Jervis Bay Kuringai South Coast Cape York Lamington Mossman Atherton Ingham Lee Up Lawn Hill Tas.WS Tas.H Tas.S Tas.R Cent. Aust Acrobates pygmaeus Aepyprymnus rufescens Antechinus agilis Antechinus flavipes Antechinus godmani Antechinus minimus minimus Antechinus stuartii Antechinus swainsonii Bettongia gaimardi cuniculus Canis lupus dingo Cercatetus caudatus Cercatetus lepidus Cercatetus nanus Chaerephon jobensis Chalinolobus dwyeri Chalinolobus gouldii Chalinolobus morio Chalinolobus nigrogriseus Dactylopsila trivirgata Dasyurus hallucatus Dasyurus maculatus Dasyurus viverrinus Dendrolagus bennettianus Dendrolagus lumholtzi Dobsonia molluccensis Echymipera rufescens Falsistrellus tasmaniensis Hemibelideus lemuroides Hipposideros ater Hipposideros cervinus Hipposideros diadema Hipposideros semoni Hydromys chrysogaster Hypsiptrymnodon moschatus Isoodon macrourus Isoodon obesulus Kerivoula papuensis Lagorchestes conspicillatus Leggadina forresti Macroderma gigas Macroglossus minimus Macropus agilis Macropus antilopinus Macropus dorsalis Macropus fuliginosus Macropus giganteus Macropus parryi Macropus robustus Macropus rufogriseus Macropus rufus Mastacomys fuscus Melomys burtoni Melomys cervinipes Melomys littoralis Mesembriomys gouldii Miniopterus australis Miniopterus schreibersii Mormopterus beccarii Mormopterus loriae Mormopterus norfolkensis Mormopterus planiceps Mormopterus spp Myotis adversus Ningaui ridei Notomys alexis Nyctimene robinsoni Nyctinomus australis Nyctophilus bifax Nyctophilus geoffroyi Nyctophilus gouldi Nyctophilus walkeri Onychogalea unguifera Ornithorhynchus anatinus Perameles gunnii Perameles nasuta Petauroides volans Petaurus australis Petaurus breviceps Petaurus gracilis Petaurus norfolcensis Petrogale assimilis Petrogale godmani Petrogale lateralis purpureicollis Petrogale mareeba Petrogale penicillata Petrogale sharmani Petropseudes dahli Phalanger intercastellanus Phascogale tapoatafa Phascolarctos cinereus Pipistrellus adamsi Planigale ingrami Planigale maculata Pogonomys mollipilosus Potorous longipes Potorous tridactylus Pseudocheirops archeri Pseudochirulus herbertensis Pseudocheirus peregrinus Pseudomys delicatulus Pseudomys desertor Pseudomys fumeus Pseudomys gracilicaudatus Pseudomys hermannsburgensis Pseudomys higginsi Pseudomys nanus Pseudomys novaehollandiae Pseudomys oralis Pseudomys shortridgei Pteropus alecto Pteropus conspicillatus Pteropus poliocephalus Pteropus scapulatus Rattus fuscipes Rattus leucopus Rattus lutreolus Rattus sordidus Rattus tunneyi Rattus villosissimus Rhinolophus megaphyllus Rhinolophus philippinensis Rhinonicteris aurantius Saccolaimus flaviventris Saccolaimus saccolaimus Sarcophilus harrisii Scoteanax rueppellii Scotorepens balstoni Scotorepens greyii Scotorepens orion Sminthopsis leucopus Sminthopsis macroura Sminthopsis murina Sminthopsis ooldea Sminthopsis virginiae Sminthopsis youngsonii Spilocuscus maculatus Syconycteris australis Tachyglossus aculeatus Taphozous australis Taphozous georgianus Taphozous hilli Thylacinus cynocephalus Thylogale billardierii Thylogale stigmatica Thylogale thetis Trichosurus caninus Trichosurus vulpecula Uromys caudimaculatus Uromys hadrourus Vespadelus baverstocki Vespadelus darlingtoni Vespadelus finlaysoni Vespadelus pumilus Vespadelus regulus Vespadelus troughtoni Vespadelus vulturnus Vombatus ursinus Wallabia bicolor Zyzomys argurus Total species richness 14 20 27 18 37 24 32 39 32 31 26 7 34 30 31 28 55 64 63 37 56 38 38 33 16 18 22 21 18 Species richness minus bats 13 19 22 18 27 18 22 27 21 21 17 5 23 20 17 21 36 43 38 28 47 27 36 17 11 13 17 16 8

181 CHAPTER 5 CHAPTER 5

APPENDIX C2. Species list for New Guinea locality (see Chapter 5 for site and habitat description - data was obtained from mammal distribution maps, vegetation maps and geological maps, as described in Chapter 5).

Anisomys imitator Antechinus melanurus Chiruromys vates Crossomys moncktoni Total species richness = 48 Dactylopsila trivirgata Species richness minus bats = 42 Dasyurus albopunctatus Dendrolagus dorianus Dendrolagus spadix Distoechurus pennatus Echymipera kalubu Emballonura furax Hipposideros maggietaylorae Leptomys elegans Lorentzimys nouhuysi Mallomys aroaensis Melomys levipes Melomys lutillus Melomys rubex Microhydromys richardsoni Microperoryctes longicaudata Murexia longicaudata Myoictis melas Nyctimene albiventer Nyctimene cyclotis Nyctimene draconilla Parahydromys asper Peroryctes raffrayana Petaurus breviceps Phalanger carmelitae Phalanger orientalis Phalanger sericeus Phalanger vestitus Pogonomys loriae Pogonomys macrourus Pseudocheirops corinnae Pseudocheirus canescens Pseudocheirus forbesi Rattus leucops Rattus niobe Rattus steini Rattus verecundus Rhinolophus arcuatus Spilocuscus maculatus Strigocuscus gymnotis Tachyglossus aculeatus Uromys caudimaculatus Xenuromys barbatus Zaglossus bruijn

182 CHAPTER 5

APPENDIX C3. Species lists for Central American sites (from Medellin, 1994 and references within). Genus Species Chamela Mts Azules La Barro Guatopo Masaguaral Cuzco Manu Belem Selva Primates Alouatta belzebul 1 Alouatta palliata 1 1 Alouatta pigra 1 Alouatta seniculus 1 1 1 1 Aotus lemurinus 1 Aotus nigriceps Aotus trivirgatus 1 1 Ateles belzebuth 1 1 1 Ateles geoffroyi 1 1 1 Ateles paniscus 1 1 Callicebus moloch 1 1 Callimico goeldii 1 1 Cebuella pygmaea 1 1 Cebus albifrons 1 1 Cebus apella 1 1 1 Cebus capucinus 1 1 Cebus nigrivittatus 1 1 Chiropotes satanas 1 lagorthix lagorthricha 1 1 Pithecia monachus 1 Pithecia sp. 1 Saguinus fuscicollis 1 1 Saguinus imperator 1 1 Saguinus midas 1 Saguinus oedipus 1 Saimiri boliviensis 1 Saimiri sciureus 1 1 Edentates Bradypus tridactylus 1 Bradypus variegatus 1 1 1 1 1 Cabassous centralis 1 1 1 Cabassous unicinctus 1 1 1 Choloepus didactylus 1 1 Choloepus hoffmanni 1 1 1 Cyclopes didactylus 1 1 1 1 1 1 Dasypus kappleri 1 1 Dasypus novemcinctus 1 1 1 1 1 1 1 1 1 Dasypus septemcinctus 1 Euphractus sexcinctus 1 Myrmecophaga tridactyla 1 1 1 1 Priodontes maximus 1 1 1 Tamandua mexicana 1 Tamandua tetradactyla 1 1 1 1 1 1 1 Artiodactyls Mazama americana 1 1 1 1 1 1 1 Mazama gouazoubira 1 1 1 Odocoileus virginianus 1 1 1 1 1 1 Perissodactyls Tapirus bairdii 1 1 1 Tapirus terrestris 1 1 1 1 Tayassu pecari 1 1 1 1 1 Tayassu tajacu 1 1 1 1 1 1 1 1 1 Carnivores Atelocynus microtis 1 1 Bassaricyon gabbii 1 1 Bassariscus astutus 1 1 1 1 Canis latrans 1 Cerdocyon thous 1 1 Conepatus mesoleucos 1 Conepatus semistriatus 1 1 1 1 Eira barbara 1 1 1 1 1 1 1 1 Felis yagouaroundi 1 1 1 1 1 1 1 1 1 Galictis vittata 1 1 1 1 1 Leopardis pardalis 1 1 1 1 1 1 1 1 1 Leopardis tigrinus 1 Leopardis wiedii 1 1 1 1 1 Lontra longicaudus 1 1 1 1 1 1 1 Mephitis macroura 1 Mustela africana 1

183 CHAPTER 5

APPENDIX C3. Continued…

Carnivores Genus Species Chamela Mts Azules La Barro Guatopo Masaguaral Cuzco Manu Belem Selva Mustela frenata 1 1 Mustela sp. Nasua nasua 1 1 1 1 1 1 1 Panthera onca 1 1 1 1 1 1 1 Potos flavus 1 1 1 1 1 1 Procyon cancrivorus 1 1 1 1 Procyon lotor 1 1 1 Pteronura brasiliensis 1 1 1 Puma concolor 1 1 1 1 1 1 1 1 Speothos venaticus 1 1 Spilogale pygmaea 1 Urocyon cinereoargenteus 1 1 Rodents Agouti paca 1 1 1 1 1 1 1 1 Akodon urichi 1 Baiomys musculus 1 Coendou bicolor 1 1 Coendou koopmani 1 Coendou prehensilis 1 1 Dactylomys dactylinus 1 1 Dasyprocta agouti 1 1 Dasyprocta prymnolopha 1 Dasyprocta punctata 1 1 1 Dasyprocta variegata 1 1 Dinomys branickii 1 1 Diplomys labilis 1 Echimys "armatus" 1 Echimys "chrysurus" 1 Echimys semivillosus 1 1 Echimys sp. 1 Heteromys anomalus 1 1 Heteromys desmarestianus 1 1 1 Holochilus brasilliensis 1 Hoplomys gymnurus 1 Hydrochaeris hydrochaeris 1 1 1 1 1 Isothrix bistriata 1 Liomys pictus 1 Mesomys hispidus 1 1 Microsciurus alfari 1 Myoprocta pratti 1 1 Neacomys spinosus 1 Neacomys tenuipes 1 1 Nectomys squamipes 1 1 1 Neotoma alleni 1 Nyctomys sumichrasti 1 1 1 Oecomys bicolor 1 1 1 1 1 1 Oecomys concolor 1 1 1 Oecomys tapajinus 1 Oecomys superans 1 1 Oligoryzomys microtis 1 1 Orthogeomys cherriei 1 Orthogeomys hispidus 1 Oryzomys alfari 1 Oryzomys albigularis 1 Oryzomys alfaroi 1 Oryzomys bombicinus 1 Oryzomys caliginosus 1 Oryzomys capito 1 1 1 1 1 Oryzomys couesi 1 1 Oryzomys delicatus 1 Oryzomys fulvescens 1 1 Oryzomys melanotis 1 1 Oryzomys macconelli 1 1 Oryzomys nitidus 1 1 Oryzomys yunganus 1 Osgoodomys banderanus 1

184 CHAPTER 5

APPENDIX C3. Continued…

Rodenta Genus Species Chamela Mts La Barro Guatopo Masaguaral Cuzco Manu Belem Azules Selva Ototylomys phyllotis 1 Oxymycterus sp 1 Pappogeomys bulleri 1 Peromyscus mexicanus 1 Peromyscus perfulvus 1 Proechimys brevicauda 1 1 Proechimys "guyanensis" 1 Proechimys semispinosus 1 1 1 Proechimys simonsi 1 1 Proechimys steerei 1 1 Reithrodontomys fulvescens 1 Rhipidomys couesi 1 1 Rhipidomys mastacalis 1 Rhipidomys venezuelae 1 Rhipidomys sp. 1 Rhipidomys sp2. 1 Sciurus aureogaster 1 Sciurus colliaei 1 Sciurus deppei 1 Sciurus gilvigularis 1 Sciurus granatensis 1 1 1 1 Sciurus ignitus 1 1 Sciurus sanborni 1 Sciurus spadiceus 1 1 Sciurus variegatoides 1 Sciurus yucatanensis 1 Sigmodon alstoni 1 Sigmodon hispidus 1 Sigmodon mascotensis 1 Sphiggurus mexicanus 1 1 Sphiggurus rothschildi 1 Tylomys nudicaudatus 1 Tylomys watsoni 1 Zygodontomys brevicauda 1 Zygodontomys "lasiurus" 1 Xenomys nelsoni 1 Lagomorphs Sylvilagus brasilliensis 1 1 1 1 1 1 1 1 Sylvilagus cunicularius 1 Marsupials Caluromys derbianus 1 1 1 Caluromys lanatus 1 1 Caluromys philander 1 1 Caluromysiops irrupta 1 Chironectes minimus 1 1 1 1 1 1 Didelphis marsupialis 1 1 1 1 1 1 1 1 Didelphis virginiana 1 1 Glironia sp. 1 Marmosa canescens 1 Marmosa fuscata 1 Marmosa mexicana 1 1 Marmosa murina 1 1 1 Marmosa robinsoni 1 1 Marmosops noctivagus 1 1 Marmosops parvidens 1 1 Metachirus nudicaudatus 1 1 1 1 1 Micoureus cinerea 1 1 1 Micoureus regina 1 Monodelphis adusta 1 Monodelphis americana 1 Monodelphis brevicaudata 1 Philander andersoni 1 Philander opossum 1 1 1 1 1

Chiroptera Balantiopteryx io Balantiopteryx plicata Centronycteris maximiliani

185 CHAPTER 5

APPENDIX C3. Continued…

Chiroptera Genus Species Chamela Mts La Barro Guatopo Masaguaral Cuzco Manu Belem Azules Selva Cormura brevirostris Cyttarops alecto Diclidurus albus Peropteryx kappleri Peropteryx leucoptera Peropteryx macrotis Rhynchonycteris naso Saccopteryx bilineata Saccopteryx canescens Saccopteryx leptura Noctilio albiventris Noctilio leporinus Mormoops megalophylla Pteronotus davyi Pteronotus gymnonotus Pteronotus parnellii Pteronotus personatus Chrotopterus auritus Lonchorhina aurita Macrophyllum macrophyllum Macrotus waterhousii Micronycteris brachyotis Micronycteris daviesi Micronycteris hirsuta Micronycteris megalotis Micronycteris minuta Micronycteris nicefori Micronycteris schimdtorum Micronycteris sylvestris Mimon crenulatum Mimon cozumelae Phyllostomus discolor Phyllostomus elongatus Phyllostomus hastatus Phyllostomus stenops Tonatia bidens Tonatia brasiliense Tonatia carrikeri Tonatia evotis Tonatia silvicola Trachops cirrhosus Vampyrum spectrum Lionycteris spurrelli Lonchophylla mordax Lonchophylla robusta Lonchophylla thomasi Anoura caudifer Choeroniscus godmani Choeroniscus minor Glossophaga commissarisi Glossophaga longirostris Glossophaga soricina Hylonycteris underwoodi Leptonycteris curasoae Lichonycteris obscura Musonycteris harrisoni Carollia brevicauda Carollia castanea Carollia perspicillata Carollia subrufa Rhinophylla pumilio Rhinophylla sp Ametrida centurio Artibeus jamaicensis Artibeus lituratus

186 CHAPTER 5

APPENDIX C3. Continued…

Chiroptera Genus Species Chamela Mts La Barro Guatopo Masaguaral Cuzco Manu Belem Azules Selva Artibeus obscurus Artibeus planirostris Centurio senex Chiroderma salvini Chiroderma trinitatum Chiroderma villosum Dermanura anderseni Dermanura azteca Dermanura cinerea Dermanura glauca Dermanura gnoma Dermanura hartii Dermanura phaeotis Dermanura tolteca Dermanura watsoni Dermanura sp Ectophylla alba Mesophylla macconelli Platyrrhinus brachycephalus Platyrrhinus dorsalis Platyrrhinus helleri Platyrrhinus infuscus Sphaeronycteris toxophyllum Sturnira lilium Sturnira ludovici Sturnira luisi Sturnira magna Sturnira tildae Uroderma bilobatum Uroderma magnirostrum Vampyressa bidens Vampyressa macconelli Vampyressa nymphaea Vampyressa pusilla Vampyrodes major Desmodus rotundus Diaemus youngi Diphylla ecaudata Natalus stramineus Furipterus horrens Thyroptera discifera Thyroptera tricolor Eptesicus brasiliensis Eptesicus furinalis Lasiurus borealis Lasiurus ega Lasiurus intermedius Myotis albescens Myotis elegans Myotis fortidens Myotis keaysi Myotis nigricans Myotis riparius Myotis simus Pipistrellus subflavus Rhogeessa parvula Rhogeessa tumida Bauerus dubiaquercus Eumops auripendulus Eumops bonariensis Eumops dabbenei Eumops glaucinus Eumops hansae? Eumops trumbulli Molossops greenhalli

187 CHAPTER 5

APPENDIX C3. Continued…

Chiroptera Genus Species Chamela Mts La Barro Guatopo Masaguaral Cuzco Manu Belem Azules Selva Molossops neglectus Molossops temminckii Molossus ater Molossus bondae Molossus coibensis Molossus molossus Molossus pretiosus Molossus sinaloae Nyctinomops aurispinosus Nyctinomops laticaudatus Promops centralis Total species 67 112 117 94 69 71 133 114 112 richness Species richness 34 48 51 39 40 29 8070 66 minus bats

188 CHAPTER 5

APPENDIX C4. Species lists for South American sites (from Ojeda and Mares 1989). See Chapter 5 for site and habitat descriptions. Taxon Genus Species Puna Prepuna Monte Upper Lower Transi- Montane Chac montane montane tional chaco o Marsupials Monodelphis dimidiata Marmosa constantiae Marmosa elegans Marmosa pusilla Lutreolina crassicaudata Didelphis albiventris Chiroptera Noctilio leporinus Tonatia silvicola Glossophaga soricina Chrotopterus auritus Anoura caudifer Sturnira lilium Artibeus jamaicensis Artibeus lituratus Pygoderma bilabiatum Desmodus rotundus Desmodus youngi Myotis levis Myotis nigricans Myotis albescens Eptesicus diminutus Eptesicus furinalis Histiotus macrotus Histiotus montanus Lasiurus borealis Lasiurus cinereus Molossops temminckii Molossops planirostris Tadarida brasiliensis Nyctinomops macrotis Nyctinomops laudicaudatus Eumops bonariensis Eumops glaucinus Eumops perotis Promops nasutus Molossus atter Molossus molossus Primates Alouatta caraya Cebus apella Edentates Myrmecophaga tridactyla Tamandua tetradactyla Chaetophractus vellerosus Euphractus sexcinctus Priodontes maximus Tolypeutes matacus Dasypus novemcinctus Dasypus septemcinctus Chlamyphorus retusus Lagomorpha Sylvilagus brasiliensis Rodentia Sciurus ignitus Oryzomys legatus Oryzomys longicaudatus Rhipidomys leucodactylus Akodon andinus Akodon boliviensis Akodon varius Akodon albiventer Oxymycterus paramensis Calomys laucha Calomys callosus Eligmodontia moreni Eligmodontia puerulus Phyllotis darwini Auliscomys sublimis Graomys domorum

189 CHAPTER 5

APPENDIX C4. Continued…

Taxon Genus Species Puna Prepuna Monte Upper Lower Transi- Montane Chaco montane montane tional chaco Rodentia Graomys griseoflavus Andinomys edax Holochilus brasiliensis Coendou prehensilis Microcavia australis Microcavia shiptoni Galea musteloides Pediolagus salinicola Hydrochaeris hydrochaeris Dasyprocta punctata Lagostomus maximus Lagidium viscacia Chinchilla brevicaudata Myocastor coypus Octodontomys gliroides Ctenomys frater Ctenomys mendocinus Ctenomys opimus Ctenomys saltarius Abrocoma cinerea Carnivora Dusicyon culpaeus Dusicyon griseus Dusicyon gymnocercus Cerdocyon thous Procyon cancrivorus Nasua nasua Lyncodon patagonicus Galictis cuja Eira barbara Conepatus chinga Lutra longicaudis Felis colocolo Felis tigrina Felis geoffroyi Felis weidii Felis yagouaroundi Felis concolor Panthera onca Perisso- Tapirus terrestris dactyla Artiodact- Tayassu tajacu yla Catagonus wagneri Mazama americana Mazama gouazoubira Blastoceros dichotomus Ozotoceros bezoarcticus Lama guanicoe Vicugna vicugna

Total species richness 19 7 21 11 39 70 27 46 Species richness minus 19614102646 1936 bats

190 CHAPTER 5

APPENDIX C5. Species lists for North and Central American sites (from Fleming 1973). See Chapter 5 for site and habitat descriptions. Taxon Species Tundra Wsp Vhf Dbm DTf MTf Marsupials Caluromys derbianus Chironectes minimus Didelphis marsupialis Didelphis virginiana Marmosa robinsoni Metachirus nudicaudatus Philander opossum Chiroptera Artibeus jamaicensis Artibeus lituratus Artibeus phaeotis Artibeus watsoni Carollia castanea Carollia perspicillata Chiroderma trinitatum Chiroderma villosum Chrotopterus auritus Cormura brevirostris Desmodus rotundus Eptesicus fuscus Glossophaga soricina Lasionycteris noctivagans Lasiurus borealis Lasiurus cinereus Lonchorhina aurita Macrophyllum macrophyllum Micronycteris brachyotis Micronycteris megalotis Micronycteris minuta Micronycteris sylvestris Molossus bondae Molossus coibensis Myotis keeni Myotis lucifugus Myotis nigricans Noctilio leporinus Nyctecius humeralis Phyllostomus discolor Phyllostomus hastatus Pteronotus parnellii Rhogeesa tumida Saccopteryx bilineata Tonatia sylvicola Trachops cirrhosus Uroderma bilobatum Uroderma magnirostrum Vampyressa nymphaea Vampyressa pusilla Vampyrodes carriccioli Vampyrops helleri Vampyrum spectrum Insectivores Blarina brevicauda Scalopus aquaticus Sorex arcticus Sorex cinereus Primates Allouata villosa Aotus trivirgatus Cebus capucinus Saguinus geoffroyi Edentates Bradypus infuseatus Choelepus hoffmanni Cyclopes didactylus Dasypus novemcinctus Tamandua tetradactyla Artiodactyla Alces alces Cervus canadensis Mazama americana Odocoileus virginiana Tayassu tajacu

191 CHAPTER 5

APPENDIX C5. Continued… Taxon Species Tundra Wsp Vhf Dbm DTf MTf Rodentia Agouti paca Citellus undulatus Clethrionomys gapperi Clethrionomys rutilus Coendu rothschildi Dasyprocta punctata Dicrostonyx groenlandicus Diplomys darlingi Erethizon dorsatum Glaucomys sabrinus Glaucomys volans Heteromys desmarestianus Hoplomys gymnurus Lemmus trimucronatus Liomys adspersus Marmota monax Microtus miurus Microtus oeconomus Microtus pinetorum Napeozapus insignis Nectomys alfari Nyctomys sumichrasti Ondatra zibethica Oryzomys bicolor Oryzomys capito Oryzomys concolor Peromyscus leucopus Peromyscus maniculatus Proechimys semispinosus Scuirus carolinensis Scuirus granatensis Scuirus niger Scuirus variegatoides Sigmodon hispidus Tamias striatus Tamiastriatus hudsonicus Tylomys panamensis Zygodontomys microtinus Lagomorphs Lepus othus Lepus americanus Sylvilagus floridanus Sylvilagus brasiliensis Carnivora Bassaricyon gabbi Canis lupus Eira barbara Felis concolor Felis onca Felis pardalis Galictis allamandi Gulo luscus Lutra annectens Lutra canadensis Lynx canadensis Lynx rufus Martes americana Martes pennanti Mephitis mephitis Mustela erminea Mustela frenata Mustela rixosa Mustela vison Nasua narica Potos flavus Procyon lotor Urocyon cinereoargenteus Ursus americana Ursus horribilis Vulpes fulva Total species richness 16 15 25 35 7070 Species richness minus bats 16 14 19 28 4041

192 CHAPTER 5

APPENDIX C6. 1. Trophic and locomotor adaptations for modern marsupial and monotreme species, listed by area. * denotes body mass < 500 g

AUSTRALIA - all references are from Strahan (ed.), 1998 Species Diet Locomotion Reference Acrobates pygmaeus* omnivore arboreal Woodside 1998 Aepyprymnus rufescens root/fungus hopper Dennis and Johnson 1998 Antechinus agilis* insectivore scansorial Dickman 1998 Antechinus flavipes* carnivore scansorial Van Dyck 1998 Antechinus godmani* carnivore scansorial Van Dyck 1998 Antechinus minimus insectivore terrestrial Wainer and Wilson 1998 minimus* Antechinus stuartii* insectivore scansorial Braithwaite 1998 Antechinus swainsonii* insectivore terrestrial Dickman 1998 Bettongia gaimardi root/fungus hopper Rose and Johnson 1998 cuniculus Cercatetus caudatus* omnivore arboreal Atherton and Haffenden 1998 Cercatetus lepidus* insectivore arboreal Green 1998 Cercatetus nanus* omnivore arboreal Turner and Ward 1998 Dactylopsila trivirgata* insectivore arboreal Van Dyck 1998 Dasyurus hallucatus carnivore scansorial Braithwaite and Begg 1998 Dasyurus maculatus carnivore scansorial Edgar and Belcher 1998 Dasyurus viverrinus carnivore scansorial Godsell 1998 Dendrolagus bennettianus browser/frug arboreal Martin and Johnson 1998 Dendrolagus lumholtzi browser/frug arboreal Johnson 1998 Echymipera rufescens insectivore terrestrial Gordon 1998 Hemibelideus lemuroides browser/frug arboreal Winter and Goudberg 1998 Hypsiptrymnodon omnivore scansorial Dennis and Johnson 1998 moschatus Isoodon macrourus insectivore terrestrial Gordon 1998 Isoodon obesulus insectivore terrestrial Braithwaite 1998 Lagorchestes browser/frug hopper Burbridge and Johnson 1998 conspicillatus Macropus agilis grazer hopper Merchant 1998 Macropus antilopinus grazer hopper Calaby 1998 Macropus dorsalis grazer hopper Kirkpatrick 1998 Macropus fuliginosus grazer hopper Poole 1998 Macropus giganteus grazer hopper Poole 1998 Macropus parryi grazer hopper Johnson 1998 Macropus robustus grazer hopper Poole 1998 Macropus rufogriseus grazer hopper Calaby 1998 Macropus rufus grazer hopper Newsome 1998 Ningaui ridei* insectivore scansorial McKenzie and Dickman 1998 Onychogalea unguifera browser/frug hopper Ingleby and Gordon 1998 Ornithorhynchus anatinus carnivore aquatic Carrick 1998 Perameles gunnii omnivore terrestrial Seebeck 1998 Perameles nasuta insectivore terrestrial Stodart 1998 Petauroides volans browser/frug arboreal McKay 1998 Petaurus australis omnivore arboreal Russell 1998 Petaurus breviceps* omnivore arboreal Suckling 1998 Petaurus gracilis* omnivore arboreal Van Dyck 1998 Petaurus norfolcensis* omnivore arboreal Suckling 1998 Petrogale assimilis browser/frug hopper Eldridge and Close 1998 Petrogale godmani browser/frug hopper Eldridge and Close 1998 Petrogale lateralis grazer hopper Eldridge and Close 1998 purpureicollis Petrogale mareeba browser/frug hopper Eldridge and Close 1998 Petrogale penicillata grazer hopper Eldridge and Close 1998 Petrogale sharmani browser/frug hopper Eldridge and Close 1998 Petropseudes dahli browser/frug scansorial Kerle and Winter 1998 Phalanger intercastellanus browser/frug arboreal Winter and Leung 1998 Phascogale tapoatafa* carnivore arboreal Soderquist 1998

193 CHAPTER 5

APPENDIX C6.1. continued…

AUSTRALIA - all references are from Strahan (ed.), 1998 Species Diet Locomotion Reference Phascolarctos cinereus browser/frug arboreal Martin and Handasyde 1998 Planigale ingrami* carnivore terrestrial Van Dyck 1998 Planigale maculata* insectivore terrestrial Redhead 1998 Potorous longipes root/fungus hopper Seebeck 1998 Potorous tridactylus omnivore hopper Johnson 1998 Pseudocheirops archeri browser/frug arboreal Winter and Gouldberg 1998 Pseudochirulus browser/frug arboreal Winter and herbertensis Gouldberg 1998 Pseudocheirus peregrinus browser/frug arboreal McKay and Ong 1998 Sarcophilus harrisii carnivore scansorial Jones 1998 Sminthopsis leucopus* insectivore terrestrial Lunney 1998 Sminthopsis macroura* insectivore terrestrial Morton 1998 Sminthopsis murina* insectivore terrestrial Fox 1998 Sminthopsis ooldea* insectivore terrestrial Aslin 1998 Sminthopsis virginiae* insectivore terrestrial Woolley 1998 Sminthopsis youngsonii* insectivore terrestrial McKenzie and Cole 1998 Spilocuscus maculatus browser/frug arboreal Winter and Leung 1998 Tachyglossus aculeatus insectivore terrestrial Augee 1998 Thylacinus cynocephalus carnivore terrestrial Rounsevell and Mooney 1998 Thylogale billardierii grazer hopper Johnson and Rose 1998 Thylogale stigmatica browser/frug hopper Johnson and Vernes 1998 Thylogale thetis grazer hopper Johnson 1998 Trichosurus caninus browser/frug scansorial How 1998 Trichosurus vulpecula browser/frug scansorial How and Kerle 1998 Vombatus ursinus grazer terrestrial McIlroy 1998 Wallabia bicolor browser/frug hopper Merchant 1998

194 CHAPTER 5

APPENDIX C6.1. continued…

NEW GUINEA (** indicates an author in Strahan (ed.) 1998) Species Diet Locomotion Reference Antechinus melanurus* insectivore? arboreal Flannery 1995 Dactylopsila trivirgata* insectivore arboreal Van Dyck 1998** Dasyurus albopunctatus carnivore scansorial Flannery 1995 Dendrolagus dorianus browser/frug arboreal Flannery 1995 Dendrolagus spadix browser/frug arboreal Flannery 1995 Distoechurus pennatus* omnivore arboreal Flannery 1995 Echymipera kalubu omnivore terrestrial Flannery 1995 Microperoryctes insectivore terrestrial? Flannery 1995; Menzies 1991, cited in Flannery 1995 longicaudata Murexia longicaudata* insectivore scansorial Flannery 1995 Myoictis melas* carnivore terrestrial Flannery 1995 Peroryctes raffrayana omnivore terrestrial? Flannery 1995 Petaurus breviceps* omnivore arboreal Suckling 1998** Phalanger carmelitae browser/frug arboreal Flannery 1995; Hume et al.1993 cited in Flannery 1995 Phalanger gymnotis browser/frug scansorial Flannery 1995; Hume et al.1993 cited in Flannery 1995 Phalanger orientalis browser/frug arboreal Flannery 1995 Phalanger sericeus browser/frug arboreal Flannery 1995; Hume et al.1993 cited in Flannery 1995 Phalanger vestitus browser/frug arboreal Flannery 1995; Hume et al.1993 cited in Flannery 1995 Pseudocheirops corinnae browser/frug arboreal Flannery 1995; Hume et al.1993 cited in Flannery 1995 Pseudocheirulus canescens* browser/frug arboreal Flannery 1995; Hume et al.1993 cited in Flannery 1995 Pseudocheirulus forbesi browser/frug arboreal Flannery 1995; Hume et al.1993 cited in Flannery 1995 Spilocuscus maculatus browser/frug arboreal Winter and Leung 1998** Tachyglossus aculeatus insectivore terrestrial Augee 1998** Zaglossus bruijn insectivore terrestrial Flannery 1995

CENTRAL AND SOUTH AMERICA Species Diet Locomotion Reference Caluromys derbianus* omnivore arboreal Medellin 1994; weight from Fleming 1973 Caluromys lanatus* omnivore arboreal Redford and Eisenberg 1992; weight, Nowak and paradiso 1983 Caluromys philander* omnivore arboreal Redford and Eisenberg 1992; weight, Nowak and paradiso 1983 Caluromysiops irrupta* omnivore arboreal Hutchins et al. (eds) 2003; weight inferred from Nowak and Paradiso 1983 Chironectes minimus carnivore terrestrial/aquatic? Medellin 1994 Didelphis albiventris insectivore scansorial Ojeda and Mares 1989 Didelphis marsupialis omnivore scansorial Medellin 1994 Didelphis virginiana omnivore scansorial Medellin 1994 Glironia sp. * omnivore arboreal Hutchins et al. (eds) 2003; weight inferred from Nowak and Paradiso 1983 Lutreolina crassicaudata* carnivore scansorial Ojeda and Mares 1989; weight from Nowak and Paradiso 1983 Marmosa canescens* omnivore arboreal Hutchins et al. (eds) 2003 Marmosa constantiae* insectivore scansorial Ojeda and Mares 1989; weight from Nowak and Paradiso 1983 Marmosa elegans* insectivore arboreal Nowak and Paradiso 1983; diet inferred Marmosa fuscata* insectivore arboreal Nowak and Paradiso 1983 and Hutchins et al. (eds) 2003 Marmosa mexicana* insectivore arboreal Medellin 1994 Marmosa murina* insectivore arboreal Nowak and Paradiso 1983 and Hutchins et al. (eds) 2003 Marmosa pusilla* insectivore arboreal Nowak and Paradiso 1983; diet inferred Marmosa robinsoni* insectivore arboreal Nowak and Paradiso 1983; weight inferred Hutchins et al. (eds) 2003 Marmosops noctivagus* insectivore scansorial weight based on others from this species Marmosops parvidens* insectivore scansorial weight based on others from this species Metachirus nudicaudatus omnivore scansorial Medellin 1994 Micoureus cinerea* insectivore scansorial Hutchins et al. (eds) 2003; weight inferred from other species Micoureus regina* insectivore scansorial Hutchins et al. (eds) 2003; weight inferred from other species Monodelphis adusta* carnivore scansorial Nowak and Paradiso 1983; weight inferred from other species Monodelphis americana* carnivore scansorial Redford and Eisenberg 1992; diet from Nowak and Paradiso 1983 Monodelphis brevicaudata* carnivore terrestrial Hutchins et al. (eds) 2003; weight inferred from other species Monodelphis dimidiata* insectivore scansorial Ojeda and Mares 1989; weight from Nowak and Paradiso 1983 Philander andersoni weight? insectivore scansorial Hutchins et al. (eds) 2003 Philander opossum omnivore scansorial Medellin 1994; weight from Eisenberg and Thorington 1973

195 CHAPTER 5

APPENDIX C6. 2. Trophic and locomotor adaptations for modern primate and edentate species; *denotes body mass < 500 g EDENTATES Species Diet Locomotion Reference Bradypus infuseatus browser/frugivore arboreal Fleming 1973 Bradypus tridactylus browser/frugivore arboreal Macdonald (ed), 2001 Bradypus variegatus browser/frugivore arboreal Macdonald (ed), 2001 Cabassous centralis insectivore terrestrial Macdonald (ed), 2001 Cabassous unicinctus insectivore terrestrial Medellin (1994) Chaetophractus vellerosus insectivore terrestrial Ojeda and Mares 1989 Chlamyphorus retusus insectivore terrestrial Ojeda and Mares 1989 Choloepus didactylus browser/frugivore arboreal Macdonald (ed), 2001 Choloepus hoffmanni browser/frugivore arboreal Fleming 1973 Cyclopes didactylus insectivore arboreal Fleming 1973 and Medellin 1994 Dasypus kappleri insectivore terrestrial Macdonald (ed), 2001 Dasypus novemcinctus insectivore terrestrial Fleming 1973; Ojeda and Mares 1989; Medellin 1994 Dasypus septemcinctus insectivore terrestrial Macdonald (ed), 2001; Nowak and Paradiso 1983 Euphractus sexcinctus insectivore terrestrial Macdonald (ed), 2001; Ojeda and Mares 1989 Myrmecophaga tridactyla insectivore terrestrial Macdonald (ed), 2001; Ojeda and Mares 1989 Priodontes maximus insectivore terrestrial Macdonald (ed), 2001; Ojeda and Mares 1989 Tamandua mexicana insectivore scansorial Medellin 1994 Tamandua tetradactyla insectivore scansorial Fleming 1973; Ojeda and Mares 1989 Tolypeutes matacus insectivore terrestrial Ojeda and Mares 1989

PRIMATES Species Diet Locomotion Reference Alouatta belzebul browser/frugivore arboreal Macdonald (ed), 2001 Alouatta caraya browser/frugivore arboreal Ojeda and Mares 1989 Alouatta palliata browser/frugivore arboreal Macdonald (ed), 2001 Alouatta pigra browser/frugivore arboreal Medellin 1994 Alouatta seniculus browser/frugivore arboreal Macdonald (ed), 2001 Alouatta villosa browser/frugivore arboreal Fleming 1973 Aotus lemurinus omnivore arboreal Macdonald (ed), 2001 Aotus nigriceps omnivore arboreal Macdonald (ed), 2001 Aotus trivirgatus omnivore arboreal Macdonald (ed), 2001 Ateles belzebuth browser/frugivore arboreal Macdonald (ed), 2001 Ateles geoffroyi browser/frugivore arboreal Medellin 1994 Ateles paniscus browser/frugivore arboreal Macdonald (ed), 2001 Callicebus moloch browser/frugivore arboreal Macdonald (ed), 2001 Callimico goeldii omnivore arboreal Hutchins et al. (eds) 2003 Cebuella pygmaea omnivore arboreal Macdonald (ed), 2001 weight? Cebus albifrons omnivore arboreal Macdonald (ed), 2001 Cebus apella granivore arboreal Ojeda and Mares 1989 Cebus capucinus omnivore arboreal Macdonald (ed), 2001 Cebus nigrivittatus omnivore arboreal Macdonald (ed), 2001 Chiropotes satanas granivore arboreal Macdonald (ed), 2001 Lagorthix lagorthricha browser/frugivore arboreal Macdonald (ed), 2001 Pithecia monachus granivore arboreal Macdonald (ed), 2001 Pithecia sp. granivore arboreal Macdonald (ed), 2001 Saguinus fuscicollis* omnivore arboreal Macdonald (ed), 2001; weight, De Bruine 2002 Saguinus geoffroyi omnivore arboreal Fleming 1973 Saguinus imperator omnivore arboreal Macdonald (ed), 2001 Saguinus midas omnivore arboreal Macdonald (ed), 2001 Saguinus oedipus omnivore arboreal Macdonald (ed), 2001 Saimiri boliviensis omnivore arboreal Macdonald (ed), 2001 Saimiri sciureus omnivore arboreal Macdonald (ed), 2001

196 CHAPTER 5

APPENDIX C6. 3. Trophic and locomotor adaptations for modern species of Carnivora, Perissodactyla, Artiodactyla and Lagomorphia from North, Central and South America; *denotes body mass < 500 g. CARNIVORA Species Diet Locomotion Reference Atelocynus microtis carnivore terrestrial Hutchins et al. (eds) 2003 Bassaricyon gabbii omnivore arboreal Hutchins et al. (eds) 2003 Bassariscus astutus carnivore scansorial Hutchins et al. (eds) 2003 Canis latrans carnivore terrestrial Hutchins et al. (eds) 2003 Canis lupus dingo carnivore terrestrial Corbett (in Strahan, ed., 1998) Cerdocyon thous carnivore terrestrial Ojeda and Mares 1989 Conepatus chinga insectivore terrestrial Ojeda and Mares 1989 Conepatus mesoleucus insectivore terrestrial Hutchins et al. (eds) 2003 Conepatus semistriatus omnivore terrestrial Medellin 1994 Dusicyon culpaeus carnivore terrestrial Ojeda and Mares 1989 Dusicyon griseus carnivore terrestrial inferred from Nowak and Paradiso 1983 Dusicyon gymnocercus carnivore terrestrial inferred from Nowak and Paradiso 1983 Eira barbara carnivore scansorial Ojeda and Mares 1989; Medellin 1994 Felis colocolo carnivore terrestrial Ojeda and Mares 1989 Felis concolor carnivore scansorial Fleming 1973 Felis geoffroyi carnivore arboreal inferred from Nowak and Paradiso 1983 Felis tigrina carnivore arboreal inferred from Nowak and Paradiso 1983 Felis weidii carnivore arboreal Nowak and Paradiso 1983 Felis yagouaroundi carnivore terrestrial Medellin 1994 Galactis allamandi carnivore terrestrial Fleming 1973 Galictis cuja carnivore terrestrial Ojeda and Mares 1989 Galictis vittata carnivore scansorial Nowak and Paradiso 1983 Gulo luscus omnivore terrestrial Fleming 1973 Leopardus pardalis carnivore scansorial Medellin 1994 Leopardus tigrinus carnivore scansorial Hutchins et al. (eds) 2003 Leopardus wiedii carnivore scansorial Medellin 1994 Lontra longicaudus carnivore aquatic Medellin 1994 Lutra annectens carnivore aquatic Fleming 1973 Lutra canadensis carnivore aquatic Fleming 1973 Lutra longicaudis piscivore semi-aquatic Ojeda and Mares 1989 Lyncodon patagonicus insectivore terrestrial Ojeda and Mares 1989 Lynx canadensis carnivore scansorial Fleming 1973 Lynx rufus carnivore scansorial Fleming 1973 Martes americna omnivore scansorial Fleming 1973 Martes pennanti carnivore scansorial Fleming 1973 Mephitis macroura carnivore terrestrial Hutchins et al. (eds) 2003 Mephitis mephitis carnivore terrestrial Fleming 1973 Mustela africana* carnivore scansorial inferred Hutchins et al. (eds) 2003; Nowak and Paradiso 1983 Mustela erminea* carnivore terrestrial Fleming 1973 Mustela frenata* carnivore terrestrial Fleming 1973 Mustela rixosa* carnivore terrestrial Fleming 1973 Mustela sp.* carnivore scansorial inferred Hutchins et al. (eds) 2003; Nowak and Paradiso 1983 Mustela vison carnivore terrestrial Fleming 1973 Nasua narica omnivore scansorial Fleming 1973 Nasua nasua insectivore scansorial Ojeda and Mares 1989 Panthera onca carnivore scansorial Medellin 1994 Potos flavus omni-fen arboreal Medellin 1994 Procyon cancrivorus piscivore scansorial Ojeda and Mares 1989 Procyon lotor omni-fen terrestrial Medellin 1994 Pteronura brasiliensis carnivore aquatic Hutchins et al. (eds) 2003 Puma concolor carnivore terrestrial Medellin 1994 Speothos venaticus carnivore terrestrial/aquatic Hutchins et al. (eds) 2003 Spilogale pygmaea omnivore scansorial? Hutchins et al. (eds) 2003 Urocyon carnivore terrestrial Fleming 1973; Medellin 1994 cinereoargenteus Ursus americana omnivore terrestrial Fleming 1973 Ursus horribilis omnivore terrestrial Fleming 1973 Vulpes fulva carnivore terrestrial Fleming 1973

197 CHAPTER 5

APPENDIX C6.3. continued…

PERISSODACTYLA AND ARTIODACTYLA Species Diet Locomotion Reference Alces alces grazer terrestrial Fleming 1973 Blastoceros dichotomus grazer terrestrial Ojeda and Mares 1989 Catagonus wagneri omnivore terrestrial Ojeda and Mares 1989 Cervus canadensis grazer terrestrial Fleming 1973 Lama guanicoe grazer terrestrial Ojeda and Mares 1989 Mazama americana browser/frugivore terrestrial Medellin 1994 Mazama gouazoubira grazer terrestrial Hutchins et al. (eds) 2003 Odocoileus virginianus browser/frugivore terrestrial Medellin 1994 Ozotoceros bezoarcticus grazer terrestrial Ojeda and Mares 1989 Tapirus bairdii browser/frugivore terrestrial Medellin 1994 Tapirus terrestris browser/frugivore terrestrial Hutchins et al. (eds) 2003 Tayassu pecari browser/frugivore terrestrial Medellin 1994 Tayassu tajacu omnivore terrestrial Fleming 1973; Ojeda and Mares 1989 Vicugna vicugna grazer terrestrial Ojeda and Mares 1989

LAGOMORPHIA Species Diet Locomotion Reference Lepus americanus grazer terrestrial Fleming 1973 Lepus othus grazer terrestrial Fleming 1973 Sylvilagus floridanus grazer terrestrial Fleming 1973 Sylvilagus.brasilliensis grazer terrestrial Medellin 1994 Sylvilagus.cunicularius grazer terrestrial Hutchins et al. (eds) 2003

198 CHAPTER 5

APPENDIX C6. 4. Trophic and locomotor adaptations for modern rodent species, listed by area; *denotes body mass < 500 g; ? indicates uncertainty

AUSTRALIA (** indicates an author in Strahan, ed. 1998) Species Diet Locomotion Reference Hydromys chrysogaster carnivore aquatic Olsen 1998** Leggadina forresti* omnivore terrestrial Reid and Morton 1998**;Watts and Aslin 1981 Mastacomys fuscus* grazer terrestrial? Happold 1998** Melomys burtoni* omnivore arboreal Kerle 1998** Melomys cervinipes* browser/frugiv arboreal Redhead 1998** Melomys littoralis/burtoni* omnivore arboreal Kerle 1998** Mesembriomys gouldii browser/frugiv arboreal Friend and Calaby 1998** Notomys alexis* omnivore hopper Breed 1998** Pogonomys mollipilosus* browser/frugiv arboreal Winter and Whitford 1998** Pseudomys delicatulus* granivore terrestrial Flannery (1995) Pseudomys desertor* granivore terrestrial Kerle 1998**; Watts and Aslin (1981) Pseudomys fumeus* granivore terrestrial Dixon 1998**; Nowak and Paradiso 1983 Pseudomys gracilicaudatus* omnivore terrestrial Fox 1998**; Nowak and Paradiso 1983 Pseudomys granivore terrestrial diet, Watts and Aslin (1981); Breed 1995** hermannsburgensis* Pseudomys higginsi* omnivore terrestrial Green 1998** Pseudomys nanus* grazer terrestrial Robinson 1998**; Nowak and Paradiso 1983 Pseudomys novaehollandiae* omnivore terrestrial Kemper 1998**; Nowak and Paradiso 1983 Pseudomys oralis* granivore scansorial Kirkpatrick 1998** Pseudomys shortridgei* granivore terrstrial Cockburn 1998**; Watts and Aslin (1981) Rattus fuscipes* omnivore terrstrial Lunney 1998**; Watts and Aslin (1981) Rattus leucopus* omnivore terrestrial Moore and Leung 1998** Rattus lutreolus* omnivore terrstrial Lunney 1998**; Watts and Aslin (1981) Rattus sordidus* omnivore terrstrial Redhead 1998**; Watts and Aslin (1981) Rattus tunneyi* grazer/granivo terrestrial diet, Watts 1977; Watts and Aslin (1981) re Rattus villosissimus* grazer/granivo terrastrial Watts and Aslin (1981) re Uromys caudimaculatus omnivore scansorial diet, Flannery (1995); Watts and Aslin (1981) Uromys hadrourus* omnivore scansorial Winter and Moore 1998** Zyzomys argurus* grazer scansorial diet, Watts 1977; Watts and Aslin (1981)

NEW GUINEA Species Diet Locomotion Reference Anisomys imitator granivore scansorial diet, Flannery 1995; Nowak and Paradiso 1983 Chiruromys vates* grazer arboreal weight, Flannery 1995; Nowak and Paradiso 1983 Crossomys moncktoni* carnivore aquatic Flannery 1995 Leptomys elegans* insectivore terrestrial Flannery 1995 Lorentzimys nouhuysi* omnivore arboreal Flannery 1995 Mallomys aroaensis browser/frugiv scansorial Flannery 1995 Melomys levipes* granivore scansorial Watts and Aslin 1981; Flannery 1995 Melomys lutillus* browser/frugiv scansorial Flannery 1995; Nowak and Paradiso 1983 Melomys rubex* browser/frugiv scansorial Flannery 1995; Nowak and Paradiso 1983 Microhydromys richardsoni* insectivore terrestrial Flannery 1995; Nowak and Paradiso 1983 Parahydromys asper insectivore terrestrial/aqua Flannery 1995 tic Pogonomys loriae* grazer arboreal Flannery 1995 Pogonomys macrourus* grazer arboreal Flannery 1995; Nowak and Paradiso 1983 Rattus leucops* omnivore terrestrial Flannery 1995 Rattus steini* ? ? weight inferred from other members of genus Stenomys niobe* ? ? weight inferred from other members of genus Stenomys verecundus* omnivore ? Flannery 1995 Uromys caudimaculatus omnivore scansorial diet, Flannery 1995; Watts and Aslin 1981 Xenuromys barbatus omnivore terrestrial Flannery 1995

199 CHAPTER 5

APPENDIX C6.4. continued…

NORTH AND SOUTH AMERICA Species Diet Locomotion Reference Abrocoma cinerea* grazer terrestrial Ojeda and Mares 1989 Agouti paca browser/frug terrestrial Fleming 1973; Medellin 1994 Akodon albiventer* insectivore terrestrial Ojeda and Mares 1989 weight Akodon andinus* omnivore terrestrial Nowak and Paradiso 1983 Akodon boliviensis* omnivore terrestrial Nowak and Paradiso 1983 Akodon urichi* omnivore terrestrial Redford and Eisenberg (1992) Akodon varius* omnivore terrestrial Nowak and Paradiso 1983 Andinomys edax* grazer terrestrial Ojeda and Mares 1989 Auliscomys sublimis* grazer terrestrial Ojeda and Mares 1989 Baiomys musculus* grazer terrestrial Nowak and Paradiso 1983 Calomys callosus* omnivore scansorial Ojeda and Mares 1989 Calomys laucha* grazer scansorial Nowak and Paradiso 1983 Chinchilla brevicaudata grazer terrestrial Ojeda and Mares 1989 Citellus undulatus omnivore terrestrial Fleming 1973 Clethrionomys gapperi* grazer scansorial Fleming 1973 Clethrionomys rutilus* grazer terrestrial Fleming 1973 Coendou bicolor browser/frug arboreal Nowak and Paradiso 1983 Coendou koopmani browser/frug arboreal Nowak and Paradiso 1983 Coendou prehensilis browser/frug arboreal Redford and Eisenberg 1992 Coendu rothschildi browser/frugivore arboreal Fleming 1973 Ctenomys frater* grazer terrestrial Ojeda and Mares 1989 Ctenomys mendocinus* grazer terrestrial Nowak and Paradiso 1983 Ctenomys opimus* grazer terrestrial Nowak and Paradiso 1983 Ctenomys saltarius* grazer terrestrial Nowak and Paradiso 1983 Dactylomys dactylinus browser/frug arboreal Nowak and Paradiso 1983 weight? Dasyprocta agouti browser/frug terrestrial Nowak and Paradiso 1983 Dasyprocta prymnolopha browser/frug terrestrial Nowak and Paradiso 1983 Dasyprocta punctata granivore terrestrial Fleming 1973; Ojeda and Mares 1989 Dasyprocta variegata browser/frug terrestrial Nowak and Paradiso 1983 Dicrostonyx groenlandicus* grazer terrestrial Fleming 1973 Dinomys branickii browser/frug scansorial Nowak and Paradiso 1983 Diplomys darlingi* browser/frug scansorial Fleming 1973 Diplomys labilis* browser/frug arboreal Nowak and Paradiso 1983 Echimys "armatus" weight? ? arboreal Redford and Eisenberg 1992 Echimys "chrysurus" ? arboreal Redford and Eisenberg 1992 Echimys semivillosus* browser/frug arboreal Redford and Eisenberg 1992; Teeter 2000 Echimys sp.weight? ? arboreal Redford and Eisenberg 1992 Eligmodontia moreni omnivore terrestrial Ojeda and Mares 1989 weight? Eligmodontia puerulus insectivore terrestrial Nowak and Paradiso 1983 weight? Erethizon dorsatum browser/frug arboreal Fleming 1973 Galea musteloides* grazer terrestrial Ojeda and Mares 1989 Glaucomys sabrinus* omnivore scansorial Fleming 1973 Glaucomys volans* omnivore scansorial Fleming 1973 Graomys domorum* omnivore scansorial Ojeda and Mares 1989 Graomys griseoflavus* granivore scansorial Nowak and Paradiso 1983 Heteromys anomalus* granivore terrestrial Redford and Eisenberg 1992 Heteromys granivore terrestrial Fleming 1973; Medellin 1994 desmarestianus* Holochilus brasiliensis* browser scansorial Redford and Eisenberg 1992 Hoplomys gymnurus* frugivore terrestrial Fleming 1973 Hydrochaeris hydrochaeris grazer semi-aquatic Ojeda and Mares 1989; Redford and Eisenberg 1992 Isothrix bistriata weight? ? arboreal Redford and Eisenberg 1992 Lagidium viscacia grazer terrestrial Ojeda and Mares 1989 Lagostomus maximus grazer terrestrial Ojeda and Mares 1989 Lemmus trimucronatus* grazer terrestrial Fleming 1973 Liomys adspersus* omnivore terrestrial Fleming 1973 Liomys pictus* granivore terrestrial Nowak and Paradiso 1983 Marmota monax grazer terrestrial Fleming 1973

200 CHAPTER 5

APPENDIX C6.4. continued…

NORTH AND SOUTH AMERICA Species Diet Locomotion Reference Mesomys hispidus weight? ? arboreal Nowak and Paradiso 1983 Microcavia australis* browser/frugi terrestrial Ojeda and Mares 1989; Nowak and Paradiso 1983 Microcavia shiptoni* browser/frugi terrestrial Nowak and Paradiso 1983 Microsciurus alfari* granivore arboreal Nowak and Paradiso 1983; Redford and Eisenberg 1992 Microtus miurus* grazer terrestrial Fleming 1973 Microtus oeconomus* grazer terrestrial Fleming 1973 Microtus pinetorum* grazer terrestrial Fleming 1973 Myocastor coypus grazer semi-aquatic Ojeda and Mares 1989) Myoprocta pratti browser/frug terrestrial Nowak and Paradiso 1983 Napeozapus insignis* omnivore terrestrial Fleming 1973 Neacomys spinosus* omnivore terrestrial based on similarity of molars to Oryzomys Neacomys tenuipes* omnivore terrestrial based on similarity of molars to Oryzomys Nectomys alfari* omnivore? terrestrial Fleming 1973 Nectomys squamipes* omnivore aquatic Redford and Eisenberg 1992 Neotoma alleni* grazer scansorial Nowak and Paradiso 1983 Nyctomys sumichrasti* granivore arboreal Fleming 1973; Medellin (1994) Octodontomys gliroides* grazer terrestrial Ojeda and Mares 1989; Nowak and Paradiso 1983 Oecomys bicolor* omnivore scansorial based on similarity to Oryzomys; Nowak and Paradiso 1983 Oecomys concolor* browser/frugi arboreal Vieira et al. 2003 Oecomys superans* omnivore ? based on similarity to Oryzomys; Nowak and Paradiso 1983 Oecomys tapajinus* omnivore ? based on similarity to Oryzomys; Nowak and Paradiso 1983 Oligoryzomys microtis* grazer/gran ? Nowak and Paradiso 1983 Ondatra zibethica omnivore aquatic Fleming 1973 Orthogeomys cherriei grazer terrestrial Nowak and Paradiso 1983 Orthogeomys hispidus grazer terrestrial Medellin 1994 Oryzomys albigularis* grazer/gran terrestrial Nowak and Paradiso 1983 Oryzomys alfaroi* granivore terrestrial Medellin 1994 Oryzomys bicolor* omnivore? scansorial Fleming 1973 Oryzomys bombicinus* grazer/gran ? Nowak and Paradiso 1983 Oryzomys caliginosus* grazer/gran ? Nowak and Paradiso 1983 Oryzomys capito* omnivore scansorial Redford and Eisenberg 1992; Eisenberg and Thorington 1973 Oryzomys concolor* omnivore? scansorial Fleming 1973 Oryzomys couesi* grazer/gran ? Nowak and Paradiso 1983 Oryzomys delicatus* grazer/gran ? Nowak and Paradiso 1983 Oryzomys fulvescens* granivore terrestrial Medellin (1994) Oryzomys legatus* omnivore scansorial Ojeda and Mares 1989; Nowak and Paradiso 1983 Oryzomys longicaudatus* omnivore scansorial? Ojeda and Mares 1989; Nowak and Paradiso 1983 Oryzomys macconelli* grazer/gran terrestrial Nowak and Paradiso 1983 Oryzomys melanotis* granivore terrestrial Medellin 1994 Oryzomys nitidus* grazer/gran ? Nowak and Paradiso 1983 Oryzomys yunganus* grazer/gran ? Nowak and Paradiso 1983 Osgoodomys banderanus* granivore terrestrial Nowak and Paradiso 1983 Ototylomys phyllotis* granivore scansorial Medellin 1994 Oxymycterus paramensis* insectivore terrestrial Ojeda and Mares 1989; Nowak and Paradiso 1983 Oxymycterus sp.* insectivore terrestrial Redford and Eisenberg 1992; Nowak and Paradiso 1983 Pappogeomys bulleri* grazer terrestrial Nowak and Paradiso 1983; Redford and Eisenberg 1992 Pediolagus salinicola grazer terrestrial Ojeda and Mares 1989 Peromyscus leucopus* omnivore scansorial Fleming 1973 Peromyscus maniculatus* omnivore scansorial Fleming 1973 Peromyscus mexicanus* granivore terrestrial Medellin 1994 Peromyscus perfulvus* granivore terrestrial Nowak and Paradiso 1983 Phyllotis darwini* granivore scansorial Ojeda and Mares 1989; Nowak and Paradiso 1983 Proechimys "guyanensis"* roots/fungi terrestrial Redford and Eisenberg 1992; Nowak and Paradiso 1983 Proechimys brevicauda* roots/fungi terrestrial Redford and Eisenberg 1992; Nowak and Paradiso 1983 Proechimys semispinosus* roots/fungi terrestrial Redford and Eisenberg 1992; Nowak and Paradiso 1983 Proechimys simonsi* roots/fungi terrestrial Redford and Eisenberg 1992; Nowak and Paradiso 1983 Proechimys steerei* roots/fungi terrestrial Redford and Eisenberg 1992; Nowak and Paradiso 1983

201 CHAPTER 5

APPENDIX C 6.4. continued…

NORTH AND SOUTH AMERICA Species Diet Locomotion Reference Reithrodontomys granivore scansorial Nowak and Paradiso 1983 fulvescens* Rhipidomys couesi weight? ? arboreal Nowak and Paradiso 1983 Rhipidomys leucodactylus omnivore arboreal Ojeda and Mares 1989 weight? Rhipidomys mastacalis ? arboreal Nowak and Paradiso 1983 weight? Rhipidomys sp. weight? ? arboreal Nowak and Paradiso 1983 Rhipidomys sp2 weight? ? arboreal Nowak and Paradiso 1983 Rhipidomys venezuelae ? arboreal Nowak and Paradiso 1983 weight? Sciurus aureogaster granivore arboreal Medellin 1994 weight? Sciurus carolinensis omnivore scansorial Fleming 1973 Sciurus colliaei weight? granivore arboreal Nowak and Paradiso 1983 Sciurus deppei* granivore arboreal Medellin 1994 Sciurus gilvigularis weight? granivore arboreal Nowak and Paradiso 1983 Sciurus granatensis* granivore scansorial Fleming 1973 Sciurus ignitus weight? granivore scansorial Ojeda and Mares 1989 Sciurus niger omnivore scansorial Fleming 1973 Sciurus sanborni weight? granivore arboreal Nowak and Paradiso 1983 Sciurus spadiceus* granivore arboreal Nowak and Paradiso 198; Fleming 1973 Sciurus variegatoides granivore arboreal Nowak and Paradiso 1983 Sciurus yucatanensis granivore arboreal Medellin 1994 weight? Sigmodon alstoni* omnivore terrestrial Nowak and Paradiso 1983 Sigmodon hispidus* grazer terrestrial Medellin 1994 Sigmodon mascotensis* omnivore terrestrial Nowak and Paradiso 1983 Sphiggurus mexicanus browser/frug arboreal Medellin 1994 Sphiggurus rothschildi browser/frug arboreal Nowak and Paradiso 1983 Tamias striatus* omnivore scansorial Fleming 1973 Tamiastriatus hudsonicus* omnivore scansorial Fleming 1973 Tylomys nudicaudatus* granivore arboreal Medellin 1994; Nowak and Paradiso 1983 Tylomys panamensis* omnivore? terrestrial Fleming 1973 Tylomys watsoni* granivore arboreal Nowak and Paradiso 1983 Xenomys nelsoni* ? arboreal Nowak and Paradiso 1983 Zygodontomys "lasiurus"* granivore terrestrial Nowak and Paradiso 1983 Zygodontomys brevicauda* granivore terrestrial Nowak and Paradiso 1983 Zygodontomys microtinus* omnivore terrestrial Fleming 1973

202 CHAPTER 5

APPENDIX C7.1. Split-sample cluster analysis for diet categories for modern sites

Group A

1 Prepuna Monte UppMontan V.GrGW Puna Tundra Law nHill Mos s man Atherton Dtf MTf Ingham V.GipMW CapeY ork Lamington V.GipRW V.GrOF TasWS V.GipDLF Kuringai V.GipWLF V.GipCTR V.GipRF LeeUp V.GipWTF New Guinea V .GrHRW V.GrWDF Jervis CentA ust Transitio V.GipCH V.GipHW SouthCoas V.GipSGH TasH TasS TasR Low Montan Wsp Dbm Vhf

0.9

0.8

0.7

0.6

0.5

Similarity

0.4

0.3

0.2

0.1

10 20 30 40

Group B

1 Rw enz oriN GuineaWoo NatalWood KrugerNP KafueNP Sudan WLungaNP LakeMw eru Serengeti Serengeti Rukw aVall SSWoodlan Tarangire AmboseliN LinyantiS SWarid Okavango Chobe KafueFlat SSGrassla Sahel Namibdese KalahariT Serengeti Chamela Chaco MontChac o MtAzules LaSelva Masaguara Barro Manu Guatapo Cuz co Belem ENiger Kilimanja ECr os s Makakou WNiger CongoBasi Knysna

0.9

0.8

0.7

0.6

0.5 Similarity

0.4

0.3

0.2

0.1

10 20 30 40 203 CHAPTER 5

APPENDIX C7.2. Split-sample cluster analyses for diet and locomotion categories for modern sites excluding African sites

Group A

Mos s man Atherton Ingham New Guinea Dtf V.GrHRW V.GipCH V.GipHW V.GipSGH V.GipRW V.GipMW V.GipDLF V.GipRF V.GipWLF V.GipCTR V.GipWTF TasWS TasH TasS Low Montan TasR CentAust V.GrGW Law nHill Puna Tundra

0.9

0.8

Similarity 0.7

0.6

10 20

Group B

Prepuna Monte Chamela MontChac o Chaco UppMontan Dbm Wsp Vhf MTf LaSelva Barro Cuz co Manu LeeUp Guatapo Kuringai Lamington CapeY ork SouthCoas V.GrOF V.GrWDF Jervis Transitio Belem MtAzules Masaguara

0.9

0.8

Similarity

0.7

0.6

10 20

204 CHAPTER 5

APPENDIX C8.1. Zskewness values for variables from the ‘Diet Only’ data set (sample size = 84)

Variable Zskewness Grazer 2.43 Browser 0.685 Omnivore 0.0198 Insectivore 1.928 Carnivore 0.483 Log10root/fungus 1.182

APPENDIX C8.2. Zskewness values for variables from the ‘Diet and Locomotion’ data set (sample size = 53)

Variable Zskewness Log10root/fungus 2.554 Log10granivore 0.429 Log10grazer -0.455 Log10small mixed -0.335 Log10terrestrial+hoppers -0.241 Browser -1.304 Omnivore -1.482 Insectivore 1.494 Carnivore 1.798 Arboreal 0.512 Scansorial 2.057 Other 0.795

205

CHAPTER 6

AUSTRALIA’S CHANGING CLIMATE THROUGH THE TERTIARY

CHAPTER 6

AUSTRALIA’S CHANGING CLIMATE THROUGH THE TERTIARY: REVIEW OF DIFFERENT LINES OF EVIDENCE

6.1. Climate change

Continental movements, tectonic events, and sea level and temperature changes all act to affect climatic conditions and, therefore, impact on vegetation structure and composition and faunal communities. High and uniform rainfall on mainland Australia is proposed by Truswell (1990) for the Late Cretaceous and into the Tertiary. The general trend through the Tertiary was high rainfall and widespread, dense vegetation, gradually decreasing, with vegetation becoming predominantly open in structure and pronounced seasonality developing (Martin 1994; Kershaw et al. 1994). The rainforests that covered much of Australia started to contract by the late Tertiary in response to climatic cooling and drying (Truswell 1990). Rainfall decreased through the Miocene, with the general lowering of sea surface temperatures and, by the late Miocene, there was an increase in the frequency of dry, anticyclonic circulation (Kemp 1984).

The causes of aridity in Australia were a combination of continental drift and world climatic changes. Climatic changes with a trend towards aridity through the Tertiary, were a global phenomenon (Janis 1993). For example, there is evidence for middle and late Miocene environmental change towards drier, more seasonal climates and the appearance of extensive grasslands in Northern Pakistan (Barry et al. 2002), East Africa

(Van Couvering 1999) and North America (Janis et al. 2002). Palaeoclimate can be inferred from many lines of evidence and there are many factors to be taken into account for a complete picture. It is worthwhile reviewing some of the lines of evidence and the particular results obtained for Australia as a way of verifying palaeohabitat

206 CHAPTER 6

conclusions reached in Chapter 5 and of enhancing interpretations with independent evidence.

Table 6.1 shows the geological time scale and names of eras, periods and epochs used below.

Mya Era Period Epoch 2 QUATERNARY Pleistocene 5.1 Pliocene late 10.4-5 mya

Miocene middle 16.3-10.4 mya

24.6 NEOGENE early 23.3-16.3 mya

Oligocene

38 CAINOZOIC TERTIARY Eocene 54.9 PALEOGENE Paleocene 65

Table 6.1. Geological time scale (adapted from Adam 1994; divisions of the Miocene as in Archer et al. 1998). mya = million years ago.

6.2. Evidence of climate change

6.2.1. Moving continents and changing currents

Continental drift, the relative positions of continents and associated oceanic currents and sea-level changes are important factors for climate determination. Tectonic events, such as Australia’s northward progression through the Cainozoic, have affected marine circulation patterns, and in turn, climate (Rosen 1984, Rögl 1984). The northward drift of Australia is partly associated with the increasing seasonality of moisture regimes and the evolution of vegetation on the continent (Macphail et al. 2000 in McGowran et al.

2000).

207 CHAPTER 6

Veevers (1984) outlines the timing and sequence of Australia’s separation from

Antarctica and New Zealand. Separation of Australia and Antarctica by deep waters

(about 38 million years ago; Kemp 1984) and the opening of the Drake Passage between

South America and Antarctica (about 23 million years ago; Barker and Burrell 1977, cited in Kemp 1984) created a cold current around Antarctica with subsequent ice accumulation. The size of the Antarctic ice cap has been a major influence on

Australia’s climate (Kemp 1984, Adam 1994), gradually resulting in a drying out of

Australia (White 1993; Frakes 1997, cited in Archer et al. 1998). The drier climate is to a large extent associated with precipitation levels, which are influenced by sea surface temperatures, which in turn depend on oceanic currents and meltwater from Antarctic ice (Kemp 1984; Hill et al. 1999: 288).

Australia drifted northwards into warmer, wetter latitudes while global temperatures declined from the early Palaeogene to the late Neogene (McGowran et al. 2000). This did not necessarily result in a stable climate, in fact there was a complex interaction of factors involved (Kemp 1984). These include fluctuating temperature and rainfall due to alternating greenhouse/icehouse climatic shifts throughout the Cainozoic (Archer et al.

1998 and references within) and significant sea level changes (White 1993). High sea levels produce warm, shallow seas on continental shelves which result in increased evaporation and subsequently in high precipitation levels; during times of low sea levels, continental shelves are exposed and surrounded by deep, cold seas and evaporation and precipitation are therefore low (Martin 1998). In the early Miocene, there were extensive shallow seas with warm surface waters across Australia (Kemp

1984; Hill et al. 1999; and references cited by Galloway and Kemp 1984: 87). It is expected, therefore, that precipitation was high at this time (Kemp 1984). In the middle

208 CHAPTER 6

Miocene, about 15 million years ago, the drying of Australia commenced, associated with the build up of the Antarctic ice cap. This extension of the ice-cap influenced southern hemisphere wind circulation patterns (Adam 1994) as well as causing lowering of sea levels and sea surface temperatures, which resulted in reduced precipitation

(Kershaw et al. 1994).

6.2.2. Palynology and plant macrofossils

Truswell (1990), Adam (1994), Macphail et al. (1994), Greenwood (1996), Martin

(1998) and Hill et al. (1999), among others, describe the use of pollen, spores and plant macrofossils in reconstructing palaeovegetation and deducing past climates.

Temperatures and annual rainfall for the Palaeocene and Eocene Eyre Formation in south central Australia have been estimated by Sluiter (1991, cited in Hill et al. 1999:

289), on the basis of palynological data, with mean temperatures being approximately

18-19oC, and annual precipitation over 1 400 mm. Greenwood (1996) has suggested that the Eocene-Oligocene Lake Eyre region possibly consisted of monsoonal forest communities, with seasonal rainfall of about 1 000 mm per year. Greenwood’s interpretation is based on findings of fossil Eucalyptus leaves.

Most plant fossil localities for the Oligocene and Miocene are confined to southeastern

Australia (Truswell 1990). The Oligocene was apparently a relatively wet period in the

Tertiary (annual rainfall >1800 mm), at least for southern Australia, with predominantly rainforests which started to decline in the middle Miocene (Martin 1998). Estimates of annual rainfall for the late Oligocene-early Miocene Lachlan River region in NSW, based on the requirements of plant taxa found there, are over 1 500 mm, which is the lower limit for widespread rainforest in the region today (Martin 1994 and references

209 CHAPTER 6

within). This dropped to 1 000 – 1 500 mm by the late Miocene, reaching the lower limit for wet sclerophyll forest (Ashton 1981, cited in Martin 1994). Galloway and

Kemp (1984) describe the wide distribution of Nothofagus brassi in the Miocene, based on the pollen record, and suggest that this indicates high levels of precipitation because presently this group requires 1 500 – 1 800 mm of rainfall per year. Nothofagus forests of the Oligocene were perhaps more structurally complex than the extant

Tasmanian Nothofagus forests (Truswell 1990: 15). The widespread closed Nothofagus forest of south-eastern Australia in the early Miocene included some gymnosperms as major elements (Adam 1994).

Compared to south-eastern Australia, central Australia had more sclerophyllous vegetation in the ?early-mid Miocene (Martin 1998). A Lake Frome (central Australia) late Oligocene-early Miocene assemblage contains rainforest gymnosperms with some

Nothofagus (indicative of temperate rainforest – Adam, 1994: 150) and many swamp taxa but there is an apparent change to entirely sclerophyllous vegetation there by the

?early Pliocene (Martin 1998). Palynological deposits from regions near Alice Springs and in the Lake Eyre and Lake Frome basins, dated as approximately middle Miocene, indicate at least localised development of rainforest and suggest “deep inland penetration by rain-bearing winds” (Galloway and Kemp 1984: 87).

This central region of Australia is one of the geographically closest areas to the

Riversleigh region that has produced palaeobotanical elements. Quantum Leap Site at

Riversleigh is approximately late Oligocene in age; Camel Sputum, Mike’s Menagerie and Ringtail Sites span the early to middle Miocene; and Encore Site may be early late

Miocene in age. The climatic signals at these sites, from mammalian

210 CHAPTER 6

palaeocommunities, are not inconsistent with the evidence for central Australia from the palynological record.

From palynoflora records from deep sea sites off Cairns and Mackay it is evident that rainforest, and hence high precipitation levels, were maintained in the north-east of the continent throughout the Neogene – Miocene and Pliocene epochs (Martin 1998).

Today in northern Australia, there is a number of species, closely related to rainforest taxa, interspersed in open sclerophyll communities (Webb and Tracey 1981a and

Gillison 1985, 1987, cited in Adam 1994: 156). These species possibly evolved from closed forest ancestors in response to drying in the late Tertiary (Adam 1994), indicating the presence of closed forest in northern Australia in the Tertiary. Today the monsoon tropics, compared to the south-east, have a greater interspersion of rainforest elements in the open woodlands due to the predictably high rainfall in the wet season

(Adam 1994). In the Miocene, a variation in fern and angiosperm taxa from north to south is apparent, with more groups known today from the humid tropics occurring in the north (Truswell 1990).

Evidence for a Miocene climatic change is presented by Martin (1998). Her palaeobotanic studies indicate that rainforest angiosperms were present in the late

Oligocene-early Miocene assemblages of central Australia, but in the ?early Pliocene the vegetation was entirely sclerophyllous. Martin (1998) also found that a ?late

Pliocene-Pleistocene assemblage in the Lake Eyre Basin had open shrublands. The north-west Australian vegetation record for the late Miocene shows no unequivocal rainforest taxa and a relatively dry climate (Martin 1998). A deep sea core from the north-west indicates a small increase in grass pollen in the late middle Miocene (Martin

211 CHAPTER 6

1994) and there is further increase of grass pollen from the late Miocene, through the

Pliocene and into the Pleistocene (Martin 1998). In the early Pliocene there was a strong trend in north-west and central Australia towards Acacia, Casuarinacea, Myrtacea and low open communities (Macphail et al. 2000 in McGowran et al. 2000). This indicates progressive drying of the Australian continent. Martin (1998) finds no evidence of early

Miocene grasslands but an increase in grass begins from the middle Miocene based on the appearance of phytoliths (silica bodies from the cells of grasses). No data are available for the Pliocene, from northern Australia, that may be linked with the modern savannah vegetation of the region (Hill et al. 1999). However, an early Pliocene assemblage from Lake Tay, Western Australia, has been interpreted, based on pollen, to represent dry sclerophyllous forest (Martin 1994).

Wolfe (1979) describes the use of fossil leaves in determining palaeotemperatures. This involves the correlation of leaf shape and other morphological features with a latitudinal range of temperatures for an estimate of mean annual temperature. Truswell (1990) discusses leaf morphology (size, margins, texture) and how it can be related, in a general way, to climatic factors. For example, the humid tropics have vegetation with large, smooth-margined leaves, and cool temperate zones have vegetation with small, serrated leaves. In terms of preservation and dating, the best leaf fossil bearing sites for

Australia are near Adelaide (sites at Maslins Bay, middle Eocene in age), Victoria (sites at Anglesea and the Latrobe Valley, Eocene-Pliocene in age; and Bacchus Marsh, early

Miocene in age) and Nerriga (middle Eocene in age) in southern NSW (Truswell 1990,

Greenwood 1994, Frakes 1997). Also, the late Oligocene-early Miocene Pioneer and

Monpeelyata macroflora from Tasmania are well documented (Greenwood 1994 and

Hill and Macphail 1983, cited within). A good summary of the macroflora evidence and

212 CHAPTER 6

interpretation is provided by Greenwood (1994), but is not explored here because results are very specific to the local region, making extrapolations to the rest of the continent very difficult.

Fossil leaves and fruits have been found at Melville Island, north of Darwin (Pole and

Bowman 1996). These are the only plant macrofossils studied from the north of

Australia for the Tertiary, but their age is not known more specifically than this (Pole and Bowman 1996). There is no indication of rainforest and, in fact, an open-canopy vegetation with seasonally limited moisture (probably monsoonal) is suggested for the deposit (Pole and Bowman 1996). The authors discuss this with respect to Riversleigh, suggesting that if the Riversleigh and Melville Island fossil assemblages overlap in age, the Melville Island climatic conditions would be similar to those expected at

Riversleigh, with a corresponding vegetation structure. In fact, they suggest that

Riversleigh, being further inland, would be subject to even drier conditions. Their interpretation of the possible Riversleigh vegetation in this case is gallery forest with more extensive open-canopied vegetation. Bearing in mind that the precise age of the

Melville Island fossil assemblage is unknown, if it correlates with younger Riversleigh deposits (late Miocene-Pliocene), then interpretations would not be in conflict with faunal evidence from Riversleigh.

Another potential source of information on terrestrial palaeoclimates is growth rings in fossil wood (Truswell 1990). Very weakly defined growth rings are consistent with growth in moist, tropical palaeolatitudes, whereas woods from temperate palaeolatitudes show marked growth rings (Creber and Chaloner 1984). Work in progress, by botanists at the University of Adelaide, on fossil wood from Riversleigh (R. Arena pers. comm.

213 CHAPTER 6

2005) may be informative. Growing season length, temperature, rainfall, and climatic stability all influence the condition of the cells in wood, their size and the magnitude of ring widths (Creber and Chaloner 1984). Estimates of seasonal variations in temperature have been made on the basis of tree ring widths for south-eastern Australia (Taylor et al.

1990). This type of information provides more direct evidence of patterns of seasonality than the palynological record (Macphail et al. 1994).

In addition to plants, epiphyllous fungi have the potential to be used to estimate rainfall

(Lange 1978; Wells and Hill 1993; Hill et al. 1999). Only preliminary research has been carried out so far (Hill et al. 1999). The Eocene environment of Nelly Creek, in central

Australia, has been estimated to have had a mean annual precipitation of under 1 200 mm (Lange 1978).

6.2.3. Oxygen isotopes

Oxygen isotopic (18O/16O) measurements of oceanic sediments provide direct evidence of palaeotemperatures for oceans (Brenchley 1984). Oceanic temperatures, through their relationship to precipitation levels, can have an influence deep within continental masses (Martin 1994). Higher sea temperatures result in higher evaporation and, therefore, a more humid climate (Martin 1998), which is relevant to water availability for vegetation. For continents, oxygen isotope measurements of fossil vertebrate bones can be used for determining temperatures (Kolodny and Raab, cited in Frakes 1997:

361). Elizabeth Price (University of NSW, Australia) is currently investigating the potential of undertaking oxygen isotope analysis for Riversleigh fossil bones and calcites.

214 CHAPTER 6

Sea surface temperatures off north-eastern Australia for the Cainozoic have been estimated through extrapolation of oxygen isotopic data from sites in the western Pacific

(Feary et al. 1991). For the late Oligocene, temperatures are between approximately 11o and 15o C; for the early Miocene, temperatures are between 16o and 20o C; and for the middle and late Miocene, temperatures are between 18o and 21o C. This indicates increasing temperatures through the Oligo-Miocene for areas around the north of

Australia. For the south-east of Australia, sea surface temperatures were warmer in the early Miocene than in the Oligocene (Galloway and Kemp 1984). The Tasman Sea had a rise of about 3o C in the early Miocene (Kemp 1984: 65). The middle Miocene saw a fall in temperatures around the south-east of Australia and an associated ice accumulation on Antarctica (Galloway and Kemp 1984). This trend became more pronounced in the late Miocene (Shackleton and Kennett 1975, Hayes and Frakes 1975, cited in Galloway and Kemp 1984: 87) and probably resulted in an increase in the frequency of dry, anticyclonic circulation in Australia (Galloway and Kemp 1984).

6.2.4. Sea creatures

Climate change may also be reflected in the biogeographic changes in oceanic neritic and pelagic biotas. Large warm-water foraminiferids were present in the late Oligocene around north-western Australia (Chaproniere 1984, cited in Quilty 1994), indicative of tropical conditions across northern Australia (Quilty 1994). In the late Oligocene to middle Miocene, numerous warm-water molluscan genera appeared off southern

Australia, which is indicative of a significant warming across the south of the continent, with a thermal maximum in the early middle Miocene (Maxwell and Darragh 2000 in

McGowran et al. 2000). This is also reflected in immigrations by large species of planktonic foraminifera to the south of Australia (McGowran and Li 2000 in McGowran

215 CHAPTER 6

et al. 2000). Support for warmer waters in the north of the continent during the early

Miocene is provided by a putative warm-water species of organic phytoplankton,

Homotryblium floripes, found at a site off eastern Queensland (Partridge 2000 in

McGowran et al. 2000). By the late Miocene, many of the warm-water molluscan genera had disappeared (Maxwell and Darragh 2000 in McGowran et al. 2000), in agreement with evidence of sea surface temperature changes for this period (Galloway and Kemp 1984). In the middle and late Miocene four genera of terebratellines in calcilutites (Brachiopoda) became extinct in response to a reduction in the extent and range of shallow marine environments (Richardson 2000 in McGowran et al. 2000), indicating lowering of sea levels and drier conditions.

6.2.5. Grossplots

Using palaeotemperature data obtained from methods described above, Frakes (1997) has produced ‘grossplots’ to show global temperature conditions and their variation with latitude from the Cretaceous through to the end of the middle Miocene. Grossplots

“can be used to estimate the temperature at any latitude, at any time" (Frakes 1997: 361) and as such, provide a very useful tool because direct palaeotemperature evidence is available only from a limited number of sites.

The Riversleigh region covered a latitudinal range of approximately 32o - 25o through the late Oligocene to the middle Miocene (estimates from Embleton 1984, in Veevers

1984). Estimates of oceanic palaeotemperatures are approximately 10o C in the

Oligocene, approximately 15o in the early Miocene, and 15o - 20o C in the mid-late

Miocene (Frakes 1997) at latitudes covered by Riversleigh through this time. This is roughly consistent with (but at the lower end of) temperature values obtained from

216 CHAPTER 6

extrapolation of oxygen isotope data for north-eastern Australia (see above). Grossplots for continental temperatures indicate that, at Riversleigh’s latitudinal range for the

Oligo-Miocene, temperatures on land may have been over 20o C, and possibly

20o - 25o C (Frakes 1997). Frakes explains that continental warming and cooling rates would have been less than for oceans, with only slight, short changes.

Frakes also estimates precipitation using the continental grossplot of temperature and latitude. For temperatures of 20o C, rainfall is about 500 mm per year; for temperatures of 25o C, rainfall is about 1 580 mm per year. Temperatures at Riversleigh are estimated to have been over 20o C, therefore annual rainfall would have been over 500 mm, and possibly up to 1 580 mm. Rainfall values obtained for the five study sites, in the regression analysis in Chapter 5, are over the estimated upper limit of 1 580 mm.

However, because temperature is not the only condition that affects rainfall (the degree of continentality and distance travelled by air masses over land, and the position of major water bodies are other factors), Frakes (1997) predicts that coastal areas would have received more precipitation than the estimated mean. At present, Riversleigh is not far from the north coast of Australia, at approximately 200 km distance from the Gulf of

Carpentaria. Veevers (2000a and 2000b) indicates that through the Miocene, the shoreline of this region would not have been very different from the present, with

Riversleigh being approximately twice the distance from the water as it is today.

6.2.6. Terrestrial faunas

Results obtained in Chapter 5 indicate that Riversleigh early-middle Miocene environments were generally densely forested or at least had tropical forest components and annual rainfall was accordingly high. A decrease in rainfall appears to have started

217 CHAPTER 6

by the late Miocene (on the basis of rainfall values obtained for Encore Site). This conclusion is based on aspects of community structure of the terrestrial mammalian fauna. Younger sites such as Rackham’s Roost (Pliocene in age) could not be included in these palaeoecological analyses due to sample size limitations and the quantitative nature of the analyses. Younger sites would provide an extended view of the trends. The early Miocene deposits of Riversleigh have also been interpreted to represent dense rainforest palaeoenvironments by Archer et al. (1989 and 1997) for the following reasons: high species diversity; high proportion of folivores; finely-partitioned feeding guilds; presence of taxa known only from rainforest environments; complete absence of grazers.

Woodburne et al. (1985) refer to a change in the climate of interior Australia, 10-5 million years ago (late Miocene), from relatively humid to more arid conditions.

Evidence for this includes the discovery, in central Australia, of late Oligocene to middle Miocene taxa and faunal assemblages, whose closest surviving analogues now occur mainly in humid northern Australia or New Guinea (Woodburne et al. 1985). For example, diverse pseudocheirid communities from the late Oligocene to middle

Miocene occurred in central Australia and Riversleigh, whereas diverse modern pseudocheirid communities are found in humid, dense upland forests of Australia and

New Guinea. Woodburne et al. (1985) add that, in contrast, many taxa from early

Pliocene (5.2 - 3 million year old) assemblages of central Australia have their closest living representatives in much drier regions.

Rackham’s Roost Site (at Riversleigh) is Pliocene in age and provides evidence of dry open forest conditions (Archer et al. 1997). At about this time arboreal marsupials

218 CHAPTER 6

became less common in inland regions of Australia, presumably due to increasing aridity and natural deforestation (Archer 1992; Flannery 1994). Other fossil faunal assemblages of the late Miocene and younger indicate non-rainforest environments. The habitat at Alcoota (Northern Territory) was probably very open sclerophyll forest from

10 million years ago (Murray and Megirian 1992). This conclusion is based on the almost complete absence of arboreal mammals from the Alcoota Local Fauna and on the diversity of browsing mammals (Archer et al. 1995). The Local Faunas of Bluff

Downs (eastern Queensland) and Curramulka (southern South Australia), both 5.2-3.4 million years old, also seem to indicate open sclerophyll forest and woodlands (Archer et al. 1995). The Hamilton Local Fauna (4.46 million years old), which is generally regarded to represent a rainforest community, appears to have been an exception among the other early Pliocene assemblages. However, Archer et al. (1995) suggest that rainforest refugia probably persisted in other coastal areas (e.g. north-eastern

Queensland) because of the current presence there of relictual, endemic rainforest groups, such as ringtails of the genus Pseudochirops.

6.2.7. Other factors relevant to climate and vegetation

Further impacts of tectonism are seen in topographic changes and their effects. With the collision of the Australian and south-east Asian plates, 15 million years ago, the mountains of New Guinea and the Great Divide of Australia rapidly rose in height, resulting in a rain-shadow that spread across the interior of Australia with a consequent increase in aridity (Archer 1984). Mountain ranges stop or deflect winds upwards and as a result, clouds drop most of their moisture over the mountains (Gressitt 1982).

219 CHAPTER 6

Climatic evidence can also be extracted from soil characteristics. Laterisation fixes phosphorous in soluble iron and aluminium complexes in soil (Turnbull 1997). In the early Tertiary, Australia developed lateritic soils, which still persist in tropical areas, and are an indication of a uniformly warm and humid climate (Brown et al. 1997;

Turnbull 1997; Martin 1998). Evidence for widespread deep weathering and laterised surfaces in the late Oligocene-early Miocene suggests wet and maybe relatively warm climates over much of the continent (Galloway and Kemp 1984; Kemp 1984).

Climatic conditions influence sediments and Martin (1994) found that lignites from late

Eocene-Oligocene deposits, which require very wet conditions to form, support climatic conclusions deduced from the vegetation record. She found that year-round, high humidities were maintained in the early-middle Miocene in southeastern Australia, as indicated by pale grey clays, and that these clays became restricted by the late Miocene-

Pliocene, with the increasing presence of red and brown clays indicating a well-marked dry season.

Dust blown off the continent into the ocean is an indicator of the extent of vegetation.

Wind erosion increases with a dryer climate and disturbed vegetation resulting in wind blown dust settling in the ocean (Martin 1998). This is a useful source of evidence for aridity. Several deep-sea sites on the Lord Howe Rise provide dust particle information that shows a decrease in smectite/illite ratios in the middle Miocene (10-15 million years ago), indicating increased aridification in the northern to central parts of the continent (Martin 1998). Also, the trend towards aridity is apparent from the middle

Miocene at Lake Frome, from clay mineralogy (Martin 1998; Galloway and Kemp 1984 and Callen and Tedford 1976, cited within). Although the late Miocene Encore Site

220 CHAPTER 6

local fauna suggests a drier climate relative to the early Miocene sites (Camel Sputum and Mike’s Menagerie) and the middle Miocene Ringtail Site, the signal is not as strong as for the wind blown dust particles and clay mineralogy. Possible explanations are: the age of the Encore Site deposit may be older than late Miocene and therefore represents a period before significant aridification took place; the palaeoenvironment of the Encore

Site deposit may have had a strongly seasonal climate; or the Encore Site deposit may be time averaged over a longer period that incorporates more fluctuating environments.

Martin (1998) explains that a distinct dry season would enable the transport of dust out to sea and it is possible that strong seasonality patterns marked the early stages of aridification (Callen 1977, cited in Martin 1998).

6.3. Concluding remarks

Evidence for palaeoclimate is geographically and temporally scattered.

Palaeovegetation and temperature interpretations must be extrapolated cautiously for

Riversleigh from other parts of the continent because there are few palaeobotanical records from the Riversleigh region (Martin 1998) and isoptope data are only available from the north-west and north-east of Australia. Generally, early Miocene was a warmer, wetter period for Australia than later times, with warm, shallow seas and widespread rainforests. The build up of the Antarctic ice cap, formation of mountains in

New Guinea and in Australia, and decreasing sea temperatures and levels from the middle Miocene resulted in reduced precipitation levels, contraction of rainforests and an increase in grasslands. By Pliocene times, there is increasing evidence of dry open forests on the continent (Table 6.2).

221 CHAPTER 6

EPOCH CLIMATE/ENVIRONMENT • start of major greenhouse event 23 million years ago warm-water marine taxa off southeastern Australia Late Oligocene • (~25 mya) • north-west Australia surrounded by warm oceanic temperatures • widespread wet, relatively warm climate suggested by deep weathering

• extensive shallow seas over Australia, resulting in high precipitation • increase in sea temperatures in the south-east • warm-water marine taxa off south eastern Australia • warm sea temperatures in northeastern Australia Early Miocene widespread deep weathering profiles an laterised surface confirm (23.3-16.3 mya) • abundant precipitation • widespread Nothofagus forests on mainland • rainforest angiosperms present in central Australia • Riversleigh terrestrial fossil fauna indicates dense forest • start of major icehouse event 15 million years ago • increase in grass pollen; occurrence of phytoliths • drying of continent commences; build up of Antarctic ice cap • rainforest decline • decrease in sea temperatures in the south-east Middle Miocene • lowering of sea levels begins (16.3-10.4 mya) • decrease in warm-water marine taxa • mountains of New Guinea and the Australian Great Divide form a rainshadow about 15 million years ago • dust blown out to sea indicates increase in aridification in northern and central Australia • clay mineralogy indicates increase in aridification in central Australia • extinction of brachiopod genera in response to retreat of marine water from continent • warm-water molluscan taxa disappear from southern Australia Late Miocene • increase in grass pollen for north-west Australia (10.4-5 mya) • well-marked dry season indicated for southern Australia by sediment clays • Alcoota Local Fauna (Northern Territory) probably open sclerophyll forest • entirely sclerophyllous vegetation in central Australia • open shrubland in the Lake Eyre Basin • Lake Tay assemblage (Western Australia) interpreted as dry Pliocene sclerophyll forest (5-1.6 mya) • dry open forest conditions at Riversleigh suggested by Rackham’s Roost Local Fauna • Bluff Downs (Queensland) and Curramulka (South Australia) Local Faunas indicate open sclerophyll forest and woodland

Table 6.2. Summary of evidence for climate and vegetation change in Australia during the late Tertiary (mya = million years ago).

The palaeohabitat interpretations of Riversleigh sites, made on the basis of mammalian community structure in Chapter 5, are consistent with the general climatic trends for the continent through the Tertiary. High sea temperatures and levels and widespread deep weathering profiles and laterised surfaces through the early Miocene indicate abundant

222 CHAPTER 6

precipitation (Kemp 1984) which supported widespread Nothofagus forest and rainforest angiosperm taxa. Relative estimates of annual rainfall for the five Riversleigh study sites indicate high rainfall, gradually decreasing through the Miocene, however, it is not possible to verify the absolute figures for the estimated annual rainfall. All lines of evidence support the beginning of aridification in Australia from the middle-late

Miocene (~15 to 5 mya). The palaeoenvironment of the Encore Site deposit (late

Miocene) was interpreted as forested but could not be described more precisely in

Chapter 5. Taking into account all evidence for climate change in Australia, it seems reasonable to suggest that the late Miocene of Riversleigh was drier than the preceding periods of the Tertiary and perhaps subject to greater seasonality.

Truswell (1990) suggests that rainfall in the Tertiary was probably high and uniform, except perhaps for the southern parts of the continent, where high latitudes would have imposed seasonality to some extent. No definite dry period existed in the late Oligocene

Lachlan River region of NSW, but by the late Miocene, a definite dry period had developed and burning was a regular event (Martin 1994). Evidence of drainage, significant vegetation and the existence of lakes and water courses in central Australia, in the middle Miocene, indicates humid, equable conditions (Quilty 1994). A change to seasonal conditions (in central Australia) from the middle Miocene is noted by Martin

(1989) and Macphail and Truswell (1989, cited in Quilty 1994). These broad statements support the conclusions reached about seasonality at Riversleigh (Chapter 5): that the early Miocene palaeoenvironment was one with even rainfall throughout the year. The situation for the Encore Site local fauna is less clear but it is possible that it experienced a potentially strong seasonal climate.

223 CHAPTER 6

6.4. References

Adam, P. 1994. Australian Rainforests, Oxford University Press, Oxford.

Archer, M. 1984. Evolution of arid Australia and its consequences for vertebrates. Pp. 97-108 in Vertebrate Zoogeography and Evolution in Australia, eds M. Archer and G. Clayton. Hesperian Press, Perth.

Archer, M. 1992. Ringtail possums (Pseudocheiridae, Marsupialia) from the Tertiary deposits of Riversleigh. The Beagle, Records of the Northern Territory Museum of Arts and Sciences 9: 257.

Archer, M., Burnley, I., Dodson, J., Harding, R., Head, L. and Murphy, A. 1998. From Plesiosaurs to People: 100 Million Years of Australian Environmental History. Australia: State of the Environment Technical Paper Series, Environment Autralia, Canberra.

Archer, M., Godthelp, H., Hand, S. J. and Megirian, D. 1989. Fossil mammals of Riversleigh, northwestern Queensland: preliminary overview of biostratigraphy, correlation and environmental change. Australian Zoologist 25: 29-65. Archer, M., Hand, S. J. and Godthelp, H. 1995. Tertiary environmental and biotic change in Australia. Pp. 77-90 in ‘Paleoclimate and evolution, with emphasis on human origins’ ed. E. S. Vrba, G. H. Denton, T. C. Partridge, L. H. Burckle.Yale University Press: New Haven. Archer, M., Hand, S. J., Godthelp, H. and Creaser, P. 1997. Correlation of the Cainozoic sediments of the Riversleigh World Heritage Fossil Property, Queensland, Australia. Pp. 131-52 in ‘Actes du Congres BiochroM’97’ ed. J. P. Aguilar, S. Legendre, J. Michaux. Mem. Trav. E.P.H.E., Inst. Montpellier.

Barry, J. C., Morgan, M. E., Flynn, L. J., Pilbeam, D., Behrensmeyer, A. K., Raza, S. M., Khan, I. A., Badgley, C., Hicks, J. and Kelley, J. 2002. Faunal and environmental change in the late Miocene Siwaliks of northern Pakistan. Paleobiology Memoirs Supplment to 28:1-71.

Brenchley, P. J. 1984. Introduction. Pp. xi-xv in Fossils and Climate. Ed. P. J. Brenchley. John Wiley and Sons Ltd, Chichester.

Creber, G. T. and Chaloner, W. G. 1984. Climatic indications from growth rings in fossil woods. Pp. 49-74 in Fossils and Climate. Ed. P. J. Brenchley. John Wiley and Sons Ltd, Chichester.

Feary, D. A., Davies, P. J., Pigram, C. J. and Symonds, P. A. 1991. Climatic evolution and control on carbonate deposition in northeast Australia. Palaeogeography, Palaeoclimatology, Palaeoecology 89: 341-361.

224 CHAPTER 6

Flannery, T. F. 1994. Possums of the World. Geo Productions Pty Ltd, NSW, Australia.

Frakes, L. A. 1997. Grossplots: a method for estimating the temperature state of the Earth and of Australia, Cretaceous to middle Miocene. Australian Journal of Botany 45: 359-372.

Galloway, R. and Kemp, E. 1984. Late Cainozoic environments in Australia. Pp. 83-95 in Vertebrate Zoogeography and Evolution in Australia, eds M. Archer and G. Clayton. Hesperian Press, Perth.

Greenwood, D. R. 1994. Palaeobotanical evidence for Tertiary climates. Pp. 44-59 in History of the Australian Vegetation: Cretaceous to Recent. Ed. R. S. Hill. Cambridge University Press, Great Britain.

Greenwood, D. R. 1996. Eocene monsoon forests in central Australia? Australian Systematic Botany 9: 95-112.

Gressitt, J. L. 1982. General introduction. Pp. 3-13 in Biogeography and Ecology of New Guinea. Ed. J. L. Gressitt. Dr W. Junk Publishers, The Hague.

Hill, R. S., Truswell, E. M., McLoughlin, S. and Dettman, E. 1999. Evolution of the Australian flora: fossil evidence. Pp. 251-320 in Flora of Australia Volume 1, Introduction. Eds A. E. Orchard and H. S. Thompson. ABRS/CSIRO, Australia.

Janis, C. M. 1993. Tertiary mammal evolution in the context of changing climates, vegetation, and tectonic events. Annual Review of Ecology Systems 24: 467-500.

Kemp, E. 1984. Tertiary palaeogeography and the evolution of Australian climate. Pp. 61-67 in Vertebrate Zoogeography and Evolution in Australia, eds M. Archer and G. Clayton. Hesperian Press, Perth.

Kershaw, A. P., Martin, H. A. and McEwen Mason, J. R. C. 1994. The Neogene: a period of transition. Pp. 299-327 in History of the Australian Vegetation: Cretaceous to Recent. Ed. R. S. Hill. Cambridge University Press, Great Britain.

Lange, R. T. 1978. Southern Australian Tertiary epiphyllous fungi, modern equivalents in the Australasian region, and habitat indicator value. Canadian Journal of Botany 56: 532-541.

Macphail, M. K., Alley, N. F., Truswell, E. M. and Sluiter, I. R. K. 1994. Early Tertiary vegetation: evidence from spores and pollen. Pp. 189-261 in History of the Australian Vegetation: Cretaceous to Recent. Ed. R. S. Hill. Cambridge University Press, Great Britain.

Macphail, M. K. and Truswell, E. M. 1989. Palynostratigraphy of the central west Murray Basin. BMR Journal of Australian Geology and Geophysics 11: 301-331.

225 CHAPTER 6

McGowran, B., Archer, M., Bock, P., Darragh, T. A., Godthelp, H., Hageman, S., Hand, S. J., Hill, R., Li, Q., Maxwell, P. A., McNamara, K. J., Macphail, M., Mildenhall, D., Partridge, A. D., Richardson, J., Shafik, S., Truswell, E. M. and Warne, M. 2000. Australasian palaeobiogeography: the Palaeogene and Neogene record. Memoir of the Association of Australasian Palaeontologists 23: 405-470.

Martin, H. A. 1989. Vegetation and climate of the late Cainozoic in the Murray Basin and their bearing on the salinity problem. BMR Journal of Australian Geology and Geophysics 11: 291-299.

Martin, H. A. 1994. Australian Tertiary phytogeography: evidence from palynology. Pp. 104-142 in History of the Australian Vegetation: Cretaceous to Recent. Ed. R. S. Hill. Cambridge University Press, Great Britain.

Martin, H. A. 1998. Tertiary climatic evolution and the development of aridity in Australia. Proceedings of the Linnean Society of New South Wales 119: 115-136.

Murray, P. and Megirian, D. 1992. Continuity and contrast in the middle and late Miocene vertebrate communities from the Northern Territory. The Beagle 9: 195-218.

Pole, M. S. and Bowman, D. M. J. S. 1996. Tertiary plant fossils from Australia’s ‘Top End’. Australian Systematic Botany 9: 113-126.

Quilty, P. G. 1994. The background: 144 million years of Australian palaeoclimate and palaeogeography. Pp. 14-43 in History of the Australian Vegetation: Cretaceous to Recent. Ed. R. S. Hill. Cambridge University Press, Great Britain.

Rögl, F. 1984. Neogene Paratethys, Mediterranean and Indo-Pacific seaways – implications for the paleobiogeography of marine and terrestrial biotas. Pp. 171-200 in Fossils and Climate. Ed. P. J. Brenchley. John Wiley and Sons Ltd, Chichester.

Rosen, B. R. 1984. Reef coral biogeography and climate through the late Cainozoic: just islands in the sun or a critical pattern of islands? Pp. 201-262 in Fossils and Climate. Ed. P. J. Brenchley. John Wiley and Sons Ltd, Chichester.

Taylor, G., Truswell, E. M., McQueen, K. G. and Brown, M. C. 1990. Early Tertiary palaeogeography, landform evolution and palaeoclimates of the Southern Monaro, NSW, Australia. Palaeogeography, Palaeoclimatology, Palaeoecology 78: 109-134.

Truswell, E. M. 1990. Australian rainforests: the 100 million year record. Pp. 7-22 in Australian Tropical Rainforests. Eds L. J. Webb and J. Kikkawa. CSIRO Publications, Melbourne, Australia.

Turnbull, J. W. 1997. Australian vegetation. Pp. 19-37 in Australian Trees and Shrubs: Species for Land Rehabilitation and Farm Planting in the Tropics. Eds J. C. Doran and J. W. Turnbull. Australian Centre for International Agriculture Research, Canberra.

Veevers, J. J. 1984. Phanerozoic Earth History of Australia. Clarendon Press, Oxford.

226 CHAPTER 6

Veevers, J. J. 2000a. Morphotectonics of the divergent margins. Pp. 34-51 in Billion- year History of Australia and Neighbours in Gondwanaland. Ed. J. J. Veevers. Gemoc Press, Sydney.

Veevers, J. J. 2000b. Quantitative estimates of the vertical motions of the Australian platform since the Jurassic. Pp. 91-101 in Billion-year History of Australia and Neighbours in Gondwanaland. Ed. J. J. Veevers. Gemoc Press, Sydney.

Wells, A. K. and Hill, R. S. 1993. Epiphyllous microorganims as palaeoclimate estimators: the developmental sequence of fungal ‘germlings’ on their living host. Australian Systematic Botany 6: 377-386.

White, M. E. 1993. The Nature of Hidden Worlds. Reed, Sydney.

Wolfe, J. A. 1979. Temperature parameters of humid to mesic forests of eastern Asia and relation to forests of other regions of the northern hemisphere and Australasia. Geological Survey Professional Paper 1106, Washington, DC.

227

CHAPTER 7

DIETS CONCLUSIONS

CHAPTER 7

CONCLUSIONS AND FURTHER WORK

The rich fossiliferous deposits of Riversleigh are very appropriate for palaeoecological studies of northwestern Queensland through the Tertiary. Faunal palaeocommunities from this region were examined for evidence of past habitats and the potential for climatic inference.

Taphonomic analyses of mammalian fossil remains from six Riversleigh fossil sites

(Chapter 2) reveal no biases in taxonomic or specimen representation due to predation, hydraulic transport or spatial mixing of faunal elements from different communities, thereby indicating autochthonous assemblages. Additionally, time averaging is considered to be in the order of hundreds or thousands of years, with consistent preservation quality at the sites and no mixing of animals at different evolutionary stages. Minimal weathering of bones, combined with the extent of disarticulation of skeletal elements, indicates moist depositional conditions, either underwater or in wet microenvironments for the six sites studied. Further considerations of possible biases in community representation by the Camel Sputum, Mike’s Menagerie, Quantum Leap,

Ringtail and Encore Site local faunas are presented in Chapter 5. The numbers of individuals identified for described species from these sites were found to be skewed, with much lower numbers of small species than expected. However, the adaptive structure of the fossil faunas was found to be mostly within the range for modern communities, indicating that the fossil communities may be sufficiently represented in terms of the ecological diversity characteristics derived from identified taxa.

228 CHAPTER 7

Extensive taxonomic and phylogenetic research by previous authors (refer to Table

2.3.1, Chapter 2 for species lists and references) provides the groundwork for sound palaeoecological analyses at the community level for Riversleigh fossil sites. Ecological components of the Riversleigh sites were quantified based on identified taxa. Trophic and locomotor adaptive structure of the fossil faunas from Camel Sputum, Mike’s

Menagerie, Quantum Leap, Ringtail and Encore Sites was used in palaeoecological analyses and habitat interpretations. A method for inferring locomotor behaviour of fossil marsupials was developed with consideration of the particular limits imposed by the available evidence (Chapter 3). These limits include restriction to the use of a single element due to disassociation of the majority of skeletal elements, and the inability to match post-cranial elements with cranio-dental elements, on which most Riversleigh species descriptions are based. Much research has been carried out on the use of morphometrics in determining locomotor behaviour for placental mammals. Marsupials, on the other hand, have rarely been considered in such quantitative approaches.

General habitat descriptions were ascribed to the fossil communities from Camel

Sputum, Mike’s Menagerie, Quantum Leap, Ringtail and Encore Sites using the ecological diversity profiles derived for each site. The Quantum Leap Site local fauna is tentatively interpreted to have inhabited a heterogeneous environment consisting of a mosaic of forested and structurally open areas. The palaeohabitats of Camel Sputum,

Mike’s Menagerie and Ringtail Site local faunas are interpreted to have been densely forested, with accordingly high estimates of rainfall. The Encore Site local fauna represents an apparently drier palaeohabitat, with lower rainfall. These Miocene palaeohabitats fit the general pattern of vegetation and climate change during this period for the Australian continent as a whole.

229 CHAPTER 7

Rainfall estimates for these sites are high, providing further support for forested environments. Even when the minimum value, at the 95% confidence interval, is considered for each site, average annual rainfall estimates are either over 1 000 mm (for

Camel Sputum Site in both data set analyses, and for Mike’s Menagerie and Quantum

Leap Sites in the Diet Only data set analysis), or not far below 1 000 mm (Table 5.3.9,

Chapter 5). Aside from the estimated values, the general trend in rainfall is informative.

The decrease in rainfall at Riversleigh through the Miocene is consistent with other evidence of climatic conditions for Australia during this time (see Chapter 6).

Habitat groupings for the Riversleigh fossil sites in this study serve as approximate descriptions of palaeoenvironment. This is the extent to which the available evidence can be interpreted at this stage. Fossil communities did not closely resemble the habitat types formed in the Diet and Locomotion data set analysis. These fossil communities may have been substantially different to modern communities in some respect, or a preservation bias may have affected the occurrence of calcanea (on which locomotor behaviour is based) and resulted in an apparent difference in community structure. This analysis is more sensitive in terms of distinguishing forest types, compared to the Diet

Only data set analysis. There was enough generality in the habitat types formed in the

Diet Only data set analysis to accommodate the fossil communities. The generality of the interpretations may, in part, be due to the fact that mammalian communities provide indirect evidence of vegetation structure. Alternatively, certain factors, if left unaccounted for, may affect the sites of the modern communities used as analogues.

These factors may influence the ecological diversity profiles for the communities, causing incomplete resolution of habitat types (keeping in mind that habitat types in this

230 CHAPTER 7

study are not determined a priori, but are based on the ecological diversity profiles of the communities). A possible consequence of this, for fossil communities, may be ambiguity in their grouping into broad habitat types, especially for the more sensitive analysis where the effects may be exaggerated.

Such factors are described as follows: a community of plants and/or animals does not have strict boundaries and there is always a gradual change from one community type to another, with an area of potential overlap or transition. The boundaries created for the modern communities in this analysis are arbitrary, either with respect to mammal distribution, (for example the boundaries of the communities from Victoria, Australia are determined by characteristics of vegetation) or to both mammal and plant distributions. The effects of adjacent areas on faunal composition at a study site are not considered, although they can be identified in some cases (usually when the area of a site used in the analysis is small and the site is surrounded by a variety of different vegetation types, as for some of the sites in Victoria, Australia). Heterogeneity within a habitat may not be recognised if one type of vegetation structure is dominant at the site

(as for Lawn Hill National Park) and conversely, habitat homogeneity may be masked by the effects of adjacent habitats with different vegetation structure (if, for instance, animals are able to cross between habitats). The scale at which habitat homogeneity/heterogeneity is considered is very important in this respect. For studies of the sort undertaken here, the scale needs to be determined by the potential spatial range of animal species. For example, restricting the analyses only to small species with relatively small habitat ranges may result in better resolution between habitat types. In summary, it is difficult to obtain consistency between the modern analogues used in such studies. There are factors, other than mammalian community structure and

231 CHAPTER 7

vegetation structure of the sites that must be considered. These include the area of the site; proximity to different vegetation types, or extent of vegetation structure heterogeneity at the site; and the extent of faunal exchange between a study site and other adjacent sites. Work in progress by Kenny Travouillon (University of New South

Wales) addresses the possible effects of the area of a site, its faunal composition and potential extent of faunal interchange with adjacent sites, on habitat interpretations of the site. This work may provide a means by which to quantify these effects and refine distinctions between habitats.

It is best to use multiple lines of evidence, especially in the case of palaeontology and palaeohabitat studies, because evidence is so fragmentary. A project that will be immediately pursued is the analysis of the Bitesantennary bat community as it relates to palaeoenvironment. The Bitesantennary Site local fauna is of a similar age to the Camel

Sputum and Mike’s Menagerie Site local faunas (early Miocene) and will therefore, provide further independent evidence for Riversleigh palaeohabitats at that time.

There is the potential for consolidating the palaeoenvironmental and climatic interpretations made here with the use of oxygen isotopic evidence from calcites, which provides temperature data for water bodies (to be undertaken by Elizabeth Price,

University of New South Wales). Also, carbon isotopic analyses of the tooth enamel apatite of Riversleigh fossil wombats are being carried out by Gavin Prideaux (Western

Australian Museum) and potentially will provide information on the relative abundance of C3 to C4 plants in the diet of these animals and on seasonality patterns. Additional evidence for climatic stability/seasonality, temperature and rainfall may become

232 CHAPTER 7

available from the studies being carried out (University of Adelaide) on fossil wood from Riversleigh.

Of particular importance to all studies of Riversleigh fossil material will be the results of radiometric dating. Determination of the precise ages of Riversleigh fossil deposits is essential for a clearer understanding of the temporal relationships between palaeofaunas at Riversleigh and for specification of the extent of time averaging for deposits. The use and interpretation of the fossil evidence from Riversleigh will consequently be carried out with greater confidence.

233