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Tiie Evolutionary History and Diversity of Australian Mammals TIIE EVOLUTIONARY HISTORY AND DIVERSITY OF AUSTRALIAN MAMMALS MIcHAEL ARCHERT't.RlcK ARENAI, Ml\a BlssaRova.t, K{REN BLacKt,. JEi...tl BnlunaLl], BERNARDCOoK-Etr, Fstl CneaSER', KIRqTEN CROSBY',ANNA.GILLESPIE', HENK GODTUELn', MIRANDA GOTT''', SUZeNI,IE'J. HAND', BENJAN{fi\i KEAR', ALAN, KRIK}IANN', BRITN ANNEMussER', TRSY'MYER5', NEyILLE PLEDGE''', iiiiiKnis'l .WANGO,l^riry-irr"-naumnrant, YUANQING STEVEN WROE.* Archer. M. et a\..1999. The evolutionaryhistory and diversityof Australianmammals. Australian Mammalogy2l: 1-45. Palaeodiversity and relationships of all groups of Australian mammals are reviewed. The 'Dark fossil recordspanning this time is of variablequality. Ages'about which nothing is known in terms of Australianmammal evolution include the late Triassicto late Jurassic, late Cretaceousto late Paleoceneand middle Eoceneto middle Oligocene.Very little is known about the early Cretaceousand late Miocene. The late Oligoceneto middle Miocene record documentsthe highestlevels of biodiversity known for the continent,comparabie to that which characterises the lowland rainforests of Borneo and Brazil. Order Monotremata spans at least the last 110 million years and includes four families. The enigmatic Ausktribosphenos from I 15 million-year-old sediments in Victoria may ..piesent an archaic monotreme,specialised peramurid or previously undocumented order of mammals but is unlikely to representa placentalas suggestedin the initial description. Order Microbiotheria is representedin the early Eocene(-55 mya) by two generasimilar in morphology to early Eocene taxa from Argentina. Order Peramelemorphiaspans the early Eocine to Holocene and includes at least five families. Order Dasyuromorphia spans at least the late Oligocene to Holocene and includes at least three families. Other dasyuromorphian-like marsupials are indeterminate in terms of family-level affinities. Order Notoryctemorphia spans the early Miocene to Holocene rvith one family. Order Yalkaparidontia spans the late Oligocene to middle Miocene with one genus. Order Diprotodontia spans the late Oligocene to Holocene,represented throughout by three *ujot groups: Phalangerida(eight families), Vombatomorphia (seven families) and Macropodoidea (at least three families). A possibleplacental condylarth (Tingamarua) has been iecorded from the early Eocene. An archaeonycteridid bat (Australonycteris) is known from the early Eocene.Among bats, the late Oligocene to middle Miocene is dominated by rhinolophoids, many of which have European,Asian and African affinities. Mystacinids, megadermatids,hipposiderids and molossids are rvell-representedin the Oligocene to Miocene deposits. Vespertilionids are uncommon in the Oligocene to Miocene but become more diverse in the Pliocene to Holocene. Emballonurids and rhinolophids appearfor the first time in the Plio-Pleistocene.Pteropodids are unknown prior to the Holocene. Murids span the early Pliocene to Holocene. In the oldest assemblageat Riversleigh. one undescribedlineage resemblesarchaic forms otherwise only known from the fossil recordsof Africa and Eurasia. 2Centre tschool of Biotogicat Science, (tniversity ofNew South llales, Sydney 2052; for Research into the Evolution of Australia's Terrestrial Ecosystems,Australian Museum,6- tschool Sydney, NSII 2000; of nuatural Resource sciences, 8 College St, rEndangered.oueensland (,/niverity^ of Techiology, GPO Box 2434, Brisbane, Queensland 4001; Species Unit, NSW National Parl<s and Wildlife Service, PO Box 1967, Hurstville, NSW 5South 6lnstitute iZZO' Australian Museum, North Tnrrore, Adelaide, SA 5000; of Verteblate Paleontology and Paleoanthropology, Chinese Academv of Sciences' PO Box 64j Beijing 100014, China. Manuscript received 2l October 1998; accepted 23 June 1999. AUSTRALIANMAMMALOGY MODERN understanding about the evolutionary In the palaeodiversityfigures below, we have history of Australia's mammals has derived from amalgamated all taxa known from particular four main arenas of research: palaeontology, intervalsof time as follows: early Cretaceous(taxa comparative anatomy, cytology, and molecular spanning only the late early Cretaceousfrom I I 5 biology. While palaeontological and anatomical to 100 my); early Paleocene(only the interval 63 studies have been pursued since the early part of to 6l my and only in Argentina): early Eocene the 19th Century, neither cytology nor molecular (only one intervalat approximately 55 my); latest biology made major contributions to under- Oligocene(only the latest late Oligocenefrom 26 standing about the relationships of Australian to 23.3 my); early Miocene (all of it from 23.3 to mammalsuntil the late 1970s (e.g.,Kirsch 1977). 16.3 my); middle Miocene (all of it from 16.3 to Since then, molecular systematic studies i n 10.4 my); late Miocene (all of it from 10.4 to 5 particular have played an increasingly significant my); earlyPliocene (all of it from 5 to 3.3 my); late role in testing and refining understanding about Pliocene(all of it from 3.3 to 2 my): Pleistocene relationships at all levels from intraspecific to (all of it from 2 my to 10,000yrs): Holocene(all of interordinal (e.g., Richardson et al. 19 86; the last 10,000yrs). Allocation ofparticular faunal Westerman et al. 1990; Springer et al. 1994; Messer assemblagesfrom Riversleigh to these time etal.1998). In aburst ofactivity startingin 1983, periods follows Archer et al. (1997c) and Creaser fossil representativesof most families of living (1997). Hence. seven morphologically distinct Australian terrestrial, arboreal and volant mammals dasyuromorphians indicated by Wroe and have now been found, thereby providing minimal Muirhead (this work) for the eariy Miocene ages of family-level differentiation and much new representsminimum species diversity for this information about the interrelationships of group spanningthis total interval of time. In these otherwise disparate groups. figures, taxa known from one or more of these intervalsare shown as occurring throughout these hoped that what follows is a brief but It is intervals, despite the fact that in some cases they understanding about succinct overview of current may only be known from a single fossil locality. families the fossil record and relationshipsof all This has been adopted because of current terrestrial, arboreal and volant of Australian uncertainty about the precise ages (rather than mammals. would in increasingly As be expected an 'early Miocene' etc.) of each of the hundreds of active field research, much of this of assemblagesknown. However, because mammal understanding is in process tested the of being speciescommonly last 3 to 8 million years,these yet published. reason and somehas to be For this generalisationsare probably not unreasonable. and because of page-length restrictions, the monophyly of most taxa in the cladogramshas not been able to be rationalisedhere. Publishedand/or SYNOPSIS AND LIMTATIONS OF TT€ FOSSIL ongoing studies that do rationalise these RECORD OF AUSTRALIAN MAI,,[N4ALS hypotheses are indicated in the individuallv authored subsections that follow. In 1983, in a 432 page treatise over-viewing mammalianpalaeofaunas of the world, Don Savage Higher-level systematic nomenclature used and Don Russellfairly summarisedthe highlights here. unless otheruise indicated, follows Aplin of the Australian Cainozoic record in five pages and Archer (1987). Conceptsof biocorrelationand (accompanied an upside down map of gaps in the record of Australian Tertiary mammal- by Australia!)" At time there were about 75 bearing sediments follow Archer et al. (1997c). that Tertiary described.Subsequent growth in Abbreviations used here are as follows: my. species Australian palaeomammalogy led to many new million years;myo, million years old; mya or M4 u'hich have more than trebled that million years ago; K4 thousand years ago; LF, discoveries level of understanding.Key events in the histor, Local Fauna(: a contemporaneousassemblage of Australianmammals in Table l. creatures recovered from a single or closell of are summarised Correlation and age of significant Tertiary fossil associated series of fossil localities); lfs, local mammalassemblages in Australia are summarised faunas; Fm, Formation (= a body of rock strata, in Fisure1. commonly of similar rock rype). Dental homology in marsupials follows Luckeft (1993) for P3-M4 and Flower (1867) for ll-P2. ARCHERET AL. EVOLUTIONOF AUSTRALIANMAMMALS 'Dark age 1'for Australian mammalsbetween 220 to 115 mya; none are known despitebeing present in other areasof the world as early as 220 mya. Ausktribosphenosnyktos, -ll5 mya (e. Cret.), ?monotreme,peramurid or an unknown group (but almost certainly not a placental),Flat Rocks, Vic. Steropod'on galmani , Kollikodon ritchiei, ,-110 to 100 mya (e. Cret.),__monotypicmembers of the monotreme families Steropodontidaeand Kollikodontidae,Lightning Ridge, NSW. 'Dark age 2' for Australian mammalsfrom 100 to 55_mya. lvlonotrimatumsudamericanum,63 to 6l mya (e. Paleoc.),ornithorhynchid monotreme,Argentina (Patagonia), SAm (with oldest undoubted SAm marsupialsand placentals). Marsupiali probably reachedAustralia from SA, via Ant.. sometimeb9twe91 65 a{l-d.5.5 mya.^ . Thylaiotinga- bariholomaii, microbiotheriids, other marsupials with SAm affinities, a ?dasyuromorphian.._ perameloid, Tingamarra porterorum (possible placental) and Australonycteris clarkeae (archaeonycteridi bat), -55 mya (e. Eoc.), Murgon, SE Qld. 'Dark for Australian mammals extendsfrom 55 to 26 mya. ase 3' ^ 'spotty' After fiial separation from Ant. between
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