An emerging consensus in the evolution, phylogeny, and systematics of marsupials and their fossil relatives (Metatheria)

Marsupials and their fossil relatives, which collectively comprise Metatheria, have been of scientific interest for centuries, with many aspects of their evolution and systematics subject to intense research and debate. Here, we review progress over the last 25 years, which has included the description of many new species (modern and fossil), and major improvements in understanding of their phylogenetic relationships, as well as the overall evolutionary history and biogeography of Marsupialia (crown-clade) and Metatheria (total-clade). Significant advances have included the deployment of increasingly sophisticated molecular, morphological, and total evidence analyses, which have resolved most previously disputed relationships among and within the modern marsupial orders. A broad systematic consensus is now emerging, although several major areas of contention remain, particularly among fossil metatherians. New modern species continue to be described at an impressive rate, with almost 50 named in the last 25 years, and many more await discovery. There has also been an explosion in the discovery and description of fossil marsupials and non-marsupial metatherians (~270 species), principally from Australasia and the Americas but also from Antarctica, Europe, and Asia. Most are represented by dental specimens only, but some consist of complete and well-preserved material, which has led to major improvements in our understanding of the evolution of cranial and postcranial morphology. Improvements in the fossil record and advances in methods for inferring divergence times have helped clarify when and where key events occurred in metatherian evolution, and the patterns of subclade diversification. We also have improved understanding of biogeographical relationships among metatherians on different landmasses. Despite enormous progress, numerous key uncertainties remain due to major gaps in the fossil record (e.g., Antarctica, Late Cretaceous, and early Paleogene of Australia) and a comparative lack of studies that directly combine molecular and fossil data. Future advances will largely depend on improvements in the fossil record and studies that better integrate neontological and paleontological evidence

A Phylogeny and Timescale for the Evolution of Pseudocheiridae (Marsupialia: Diprotodontia) in Australia and New Guinea

Pseudocheiridae (Marsupialia: Diprotodontia) is a family of endemic Australasian arboreal folivores, more commonly known as ringtail possums. Seventeen extant species are grouped into six genera (Pseudocheirus, Pseudochirulus, Hemibelideus, Petauroides, Pseudochirops, Petropseudes). Pseudochirops and Pseudochirulus are the only genera with representatives on New Guinea and surrounding western islands. Here, we examine phylogenetic relationships among 13 of the 17 extant pseudocheirid species based on protein-coding portions of the ApoB, BRCA1, ENAM, IRBP, Rag1, and vWF genes. Maximum parsimony, maximum likelihood, and Bayesian methods were used to estimate phylogenetic relationships. Two different relaxed molecular clock methods were used to estimate divergence times. Bayesian and maximum parsimony methods were used to reconstruct ancestral character states for geographic provenance and maximum elevation occupied. We find robust support for the monophyly of Pseudocheirinae (Pseudochirulus + Pseudocheirus), Hemibelidinae (Hemibelideus + Petauroides), and Pseudochiropsinae (Pseudochirops + Petropseudes), respectively, and for an association of Pseudocheirinae and Hemibelidinae to the exclusion of Pseudochiropsinae. Within Pseudochiropsinae, Petropseudes grouped more closely with the New Guinean Pseudochirops spp. than with the Australian Pseudochirops archeri, rendering Pseudochirops paraphyletic. New Guinean species belonging to Pseudochirops are monophyletic, as are New Guinean species belonging to Pseudochirulus. Molecular dates and ancestral reconstructions of geographic provenance combine to suggest that the ancestors of extant New Guinean Pseudochirops spp. and Pseudochirulus spp. dispersed from Australia to New Guinea ∼12.1–6.5 Ma (Pseudochirops) and ∼6.0–2.4 Ma (Pseudochirulus). Ancestral state reconstructions support the hypothesis that occupation of high elevations (>3000 m) is a derived feature that evolved on the terminal branch leading to Pseudochirops cupreus, and either evolved in the ancestor of Pseudochirulus forbesi, Pseudochirulus mayeri, and Pseudochirulus caroli, with subsequent loss in P. caroli, or evolved independently in P. mayeri and P. forbesi. Divergence times within the New Guinean Pseudochirops clade are generally coincident with the uplift of the central cordillera and other highlands. Diversification within New Guinean Pseudochirulus occurred in the Plio-Pleistocene after the establishment of the Central Range and other highlands

Skeleton of an unusual cat-sized marsupial relative (Metatheria: Marsupialiformes) from the middle Eocene (Lutetian: 44-43 million years ago) of Turkey

We describe a near-complete, three-dimensionally preserved skeleton of a metatherian (relative of modern marsupials) from the middle Eocene (Lutetian: 44–43 million years ago) Lülük member of the Uzunçarşıdere Formation, central Turkey. With an estimated body mass of 3–4 kg, about the size of a domestic cat (Felis catus) or spotted quoll (Dasyurus maculatus), it is an order of magnitude larger than the largest fossil metatherians previously known from the Cenozoic of the northern hemisphere. This new taxon is characterised by large, broad third premolars that probably represent adaptations for hard object feeding (durophagy), and its craniodental morphology suggests the capacity to generate high bite forces. Qualitative and quantitative functional analyses of its postcranial skeleton indicate that it was probably scansorial and relatively agile, perhaps broadly similar in locomotor mode to the spotted quoll, but with a greater capacity for climbing and grasping. Bayesian phylogenetic analysis of a total evidence dataset comprising 259 morphological characters and 9kb of DNA sequence data from five nuclear protein-coding genes, using both undated and “tip-and-node dating” approaches, place the new taxon outside the marsupial crown-clade, but within the clade Marsupialiformes. It demonstrates that at least one metatherian lineage evolved to occupy the small-medium, meso- or hypo-carnivore niche in the northern hemisphere during the early Cenozoic, at a time when there were numerous eutherians (placentals and their fossil relatives) filling similar niches. However, the known mammal fauna from Uzunçarşıdere Formation appears highly endemic, and geological evidence suggests that this region of Turkey was an island for at least part of the early Cenozoic, and so the new taxon may have evolved in isolation from potential eutherian competitors. Nevertheless, the new taxon reveals previously unsuspected ecomorphological disparity among northern hemisphere metatherians during the first half of the Cenozoic

Upper Oligocene–lower-middle Miocene peramelemorphians from the Etadunna, Namba and Wipajiri formations of South Australia

The oldest fossils referable to the marsupial order Peramelemorphia (which includes modern bandicoots and bilbies) stratigraphically date from the upper Oligocene of Australia. Here we describe new ancient peramelemorphian remains from the Etadunna, Namba and Wipajiri formations, which fill gaps in the documented evolutionary history of the clade spanning some 10 million years. The identified taxa include three new species of Bulungu that incorporate the oldest stem peramelemorphian known to date from Faunal Zone A of the Etadunna Formation. We also report a new genus and species of potential thylacomyid, which pushes back the divergence of bilbies by ∼10 million years. Finally, we present a phylogenetic analysis that clarifies the evolutionary relationships of these new taxa with a reassignment of the genus Bulungu to Yaraloidea

Two new species of the genus Gumardee (Marsupialia, Macropodiformes) reveal the repeated evolution of bilophodonty in kangaroos

Two new species of the fossil macropodiform genus Gumardee are described that provide insights into the evolution of early kangaroos. Gumardee has a continuous record from the late Oligocene to the early Miocene and is likely to have become extinct before the middle Miocene. The two new species are the most derived members of the genus. They exhibit some dental size variation, especially in P3/p3, which we interpret as evidence for sexual dimorphism. From the stratigraphically oldest species of Gumardee to the youngest, the dental morphology also shows a progression from bunolophodont to increasingly bilophodont. This trend appears to have occurred in at least four different lineages of macropodoids; however, a fully bilophodont condition was achieved only twice, once in the ancestral lineage of balbarids, and again in ancestral radiation of modern macropodids as implied by the species of Gumardee. Kenny. J. Travouillon[[email protected]], Western Australian Museum, Locked Bag 49, Welshpool DC, WA, 6986 Australia; Kaylene Butler[[email protected]], School of Earth and Environmental Sciences, The University of Queensland, St Lucia, Queensland 4072, Australia; Michael Archer[[email protected]], Earth and Sustainability Science Research Centre, School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia; Suzanne J. Hand[[email protected]], Earth and Sustainability Science Research Centre, School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia

Revision of Oligo-Miocene kangaroos, Ganawamaya and Nambaroo (Marsupialia: Macropodiformes, Balbaridae).

The generic and specific status of fossil kangaroo specimens attributed to Nambaroo and Ganawamaya from the Riversleigh World Heritage Area, Australia, are revised and new material is described. Results indicate that the previously proposed 12 species (eight of Nambaroo and four of Ganawamaya) represent four species from a single genus (Ganawamaya). Previous studies distinguished species of Ganawamaya from Nambaroo in lacking a protostylid on the lower first molar. However, our analyses indicate that the apparent presence/absence of the protostylid is related to the degree of dental wear, being totally worn in older individuals. Unworn specimens from Riversleigh were previously typically attributed to Nambaroo, while worn specimens were most commonly attributed to Ganawamaya. With support from phylogenetic analyses, we refer Riversleigh's N. couperi and N. gillespieae to Ganawamaya. The diagnoses of both genera are revised. We also assign undescribed specimens from the Ngapakaldi Quarry and Leaf Locality, South Australia, to Gan. couperi comb. nov. and Gan. acris, respectively, and describe new material of Gan. acris and Gan. aediculis

The osteology and systematics of the enigmatic Australian Oligo-Miocene Metatherian Yalkaparidon (Yalkaparidontidae; Yalkaparidontia;?Australidelphia; Marsupialia)

We provide the first detailed description of the osteology of the enigmatic Oligo-Miocene Australian metatherian Yalkaparidon. This taxon exhibits a number of unusual craniodental apomorphies but appears to be plesiomorphic within Metatheria in retaining four molars, rather than three as previously reported. We demonstrate that the only known skull of Yalkaparidon almost certainly represents a single individual. We also tentatively refer a number of isolated tarsals to the genus. Maximum parsimony analyses of a 258 character morphological matrix (with information from the tarsals described here either included or excluded) place Yalkaparidon within the superordinal clade Australidelphia, but Bayesian analyses of the same matrix are less well resolved, placing Yalkaparidon within Marsupialia but without unequivocally supporting australidelphian affinities. Bayesian analyses of a total evidence matrix that combines the morphological data with 9 kb of sequence data from five nuclear protein-coding genes (APOB, BRCA1, IRBP, RAG1 and VWF), 78 indels, and 53 retroposon insertion characters are similarly poorly resolved and do not clarify the supraordinal relationships of Yalkaparidon beyond suggesting that it is probably a member of Marsupialia. However, if the tarsal remains are correctly attributed to Yalkaparidon, then membership of Australidelphia seems likely, as these specimens exhibit characteristic australidelphian apomorphies. We conclude that the ordinal status of Yalkaparidon remains justified based on current evidence, and we present a revised diagnosis for Yalkaparidontia. We maintain the two currently recognized species, Y. coheni and Y. jonesi, but present revised specific diagnoses. We suggest a revised phylogenetic definition for Marsupialia, and provide phylogenetic definitions for Eomarsupialia (the clade comprising all extant Australian marsupial orders) and for the clade comprising Dasyuromorphia, Peramelemorphia, and Notoryctemorphia to the exclusion of Diprotodontia; we propose the name Agreodontia for the latter clade