Cranial anatomy of the earliest marsupials and the origin of opossums.

BACKGROUND: The early evolution of living marsupials is poorly understood in part because the early offshoots of this group are known almost exclusively from jaws and teeth. Filling this gap is essential for a better understanding of the phylogenetic relationships among living marsupials, the biogeographic pathways that led to their current distribution as well as the successive evolutionary steps that led to their current diversity, habits and various specializations that distinguish them from placental mammals. METHODOLOGY/PRINCIPAL FINDINGS: Here we report the first skull of a 55 million year old peradectid marsupial from the early Eocene of North America and exceptionally preserved skeletons of an Oligocene herpetotheriid, both representing critical groups to understand early marsupial evolution. A comprehensive phylogenetic cladistic analysis of Marsupialia including the new findings and close relatives of marsupials show that peradectids are the sister group of living opossums and herpetotheriids are the sister group of all living marsupials. CONCLUSIONS/SIGNIFICANCE: The results imply that North America played an important role in early Cenozoic marsupial evolutionary history and may have even been the center of origin of living marsupials and opossums. New data from the herpetotheriid postcranium support the view that the ancestral morphotype of Marsupialia was more terrestrial than opossums are. The resolution of the phylogenetic position of peradectids reveals an older calibration point for molecular estimates of divergence times among living marsupials than those currently used.

Exceptionally preserved North American Paleogene metatherians: adaptations and discovery of a major gap in the opossum fossil record.

A major gap in our knowledge of the evolution of marsupial mammals concerns the Paleogene of the northern continents, a critical time and place to link the early history of metatherians in Asia and North America with the more recent diversification in South America and Australia. We studied new exceptionally well-preserved partial skeletons of the Early Oligocene fossil Herpetotherium from the White River Formation in Wyoming, which allowed us to test the relationships of this taxon and examine its adaptations. Herpetotheriidae, with a fossil record extending from the Cretaceous to the Miocene, has traditionally been allied with opossums (Didelphidae) based on fragmentary material, mainly dentitions. Analysis of the new material reveals that several aspects of the cranial and postcranial anatomy, some of which suggests a terrestrial lifestyle, distinguish Herpetotherium from opossums. We found that Herpetotherium is the sister group to the crown group Marsupialia and is not a stem didelphid. Combination of the new palaeontological data with molecular divergence estimates, suggests the presence of a long undocumented gap in the fossil record of opossums extending some 45Myr from the Early Miocene to the Cretaceous.

A comprehensive morphological analysis of talpid moles (Mammalia) phylogenetic relationships.

Some talpid moles show one of the most specialized suites of morphological characters seen among small mammals. Fossorial and more generalized shrew-looking moles inhabit both North America and Eurasia but these land masses share none of the same genera. One of the central questions of mole evolution has been that of how many times specialized fossorial habits evolved. We investigated the origin of mole characters with a maximum parsimony analysis of 157 characters, mostly craniodental and postcranial, of representatives of all 17 living mole genera and three shrews and one hedgehog as outgroups. The result was one most-parsimonious tree and its most novel aspect was the position of a Japanese shrew mole clade (Urotrichus, Dymecodon), which branched off after Uropsilus and was not closely related to the American shrew mole (Neurotrichus). The desmans (Galemys and Desmana) were the next clade in the tree, followed by Neurotrichus. We confirmed the monophyly of the Eurasian fossorial mole clade Talpini (Euroscaptor, Parascaptor, Mogera, Scaptochirus and Talpa). Condylura, the star-nosed mole from North America, was sister group to a clade consisting of the Talpini plus Scaptonyx and the Scalopini (Scalopus, Scapanus, Parascalops, and Scapanulus). Based on our results and on the assumption that moles originated in Eurasia, it is most parsimonious to infer one migration from Eurasia to North America and two back-migrations to Eurasia. It is ambiguous if Talpini and Scalopini evolved their full fossorial habits independently or not.

First combined cladistic analysis of marsupial mammal interrelationships.

We combine osteological, dental, and soft tissue data with sequences from three nuclear and five mitochondrial genes, sampling all major living clades of marsupials plus several extinct taxa, to perform a simultaneous analysis of marsupial interrelationships. These data were analyzed using direct optimization and sensitivity analysis on a parallel supercomputing cluster, and compared with trees produced with conventional parsimony and likelihood algorithms using a static alignment. A major issue in marsupial phylogeny is the relationships among australidelphians. Optimal analyses using direct optimization and those based on the static alignment support the basal positions of peramelians (bandicoots) and Dromiciops ('monito del monte') within Australidelphia, and in all but one case these analyses support a monophyletic Eometatheria, a group consisting of all australidelphians excluding peramelians. Dromiciops is basal within Eometatheria in analyses that maximize congruence across partitions, including the equally weighted parameter set. The topologies resulting from direct optimization under all parameter sets show some differences, but all show a high degree of resolution. Direct optimization supports high-level clades supported by analyses of partitioned molecular (e.g., Notoryctes as sister group of Dasyuromorphia) and morphological (e.g., Diprotodontia) data.

The anatomy of the world's largest extinct rodent.

Phoberomys is reported to be the largest rodent that ever existed, although it has been known only from isolated teeth and fragmentary postcranial bones. An exceptionally complete skeleton of Phoberomys pattersoni was discovered in a rich locality of fossil vertebrates in the Upper Miocene of Venezuela. Reliable body mass estimates yield approximately 700 kilograms, more than 10 times the mass of the largest living rodent, the capybara. With Phoberomys, Rodentia becomes one of the mammalian orders with the largest size range, second only to diprotodontian marsupials. Several postcranial features support an evolutionary relationship of Phoberomys with pakaranas from the South American rodent radiation. The associated fossil fauna is diverse and suggests that Phoberomys lived in marginal lagoons and wetlands.

A Report on "One Day Symposium on Numerical Cladistics".

A recent symposium on numerical cladistics held at the American Museum of Natural History in New York City, addressed novel methods for searching tree space, applications of randomizations in cladistic analysis, and data management. One of the major concerns in systematics is that of finding the global optimum in tree length. The space to search is complex because it includes many local optima. It is a difficult task to escape local optima without a great loss in efficiency. The ideal is to search among suboptimal topologies and still obtain an answer in a reasonable amount of time. Nixon presented a new family of methods called "parsimony ratchet," which are successful at escaping local optima. Moilanen presented a new program which may have similar advantages. Two presentations, one by Goloboff and Farris and another by Farris, Goloboff, Källersjö, and Oxelman, introduced modifications to parsimony jackknifing that improved its accuracy when compared to normal heuristic searches. Wheeler discussed the advantages of new methods of analyzing DNA and protein sequence data, which eliminate multiple alignment; the most recent one packs nucleotides into strings which constitute the new characters. Siddall discussed different applications of randomization in cladistics and their logical consistency, finding some more acceptable than others. Nixon and Carpenter presented a new program for managing data. This symposium will probably be a landmark judging from the originality and practicality of the points presented.