The palaeoecology of the Vombatidae: did giant wombats burrow?

Debate over the origins of burrowing in the Vombatidae has continued since the discovery of the remains of the largest of all wombats, Phascolonus gigas, in the nineteenth century. In this paper, we argue that the largest of the ancestors of extant wombats did not burrow due to physical and physiological limitations of burrows. Further, we suggest that the burrowing characteristics of the extant wombats were derived from an ancestor of similar body size (20–40 kg) that is presently not represented in the fossil record.     

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.     

New Discoveries of “Opposum-Like” Marsupials from Antarctica (Seymour Island, Medial Eocene)

Five new species of marsupials are described from the middle Eocene La Meseta Formation of Seymour Island, Antarctic Peninsula. Three are derorhynchid didelphimorphians; one species is a prepidolopid polydolopimorphian, and the last is a microbiotheriid australidelphian. Additionally, fragmentary specimens representing an indetermined derorhynchid and a possible marsupial are also described. The prepidolopid and one of the derorhynchids are sufficiently derived as to preclude any close relationship to other members of that family, but the remaining taxa show the closest affinity with species otherwise known only from Itaboraian and older faunas in Patagonia. This differs from the affinity to early Eocene (Casamayoran) taxa shown by the polydolopid marsupials and placental mammals previously known from the La Meseta Formation. The newly described marsupials indicate that the relict La Meseta Fauna is composed of forms that must have dispersed to Antarctica no later than about early late Paleocene, whereas the previously known taxa apparently arrived in the early Eocene. Ecologically, the La Meseta Fauna is composed mostly of small-sized marsupials of likely insectivorous to frugivorous habits and larger-sized placental herbivores. Whereas the ratite bird of the La Meseta Fauna was probably also herbivorous, the phorusrhachoid and falconid birds comprised a large and smaller carnivorous to possibly scavenging component, respectively. Compared to contemporary faunas of Patagonia, the medium- to large-sized marsupial carnivores are lacking in the Antarctic Peninsula. Nevertheless, the La Meseta Fauna is Patagonian in origin and affinity. In conjunction with new faunas of Itaboraian age (early late Paleocene) in Patagonia, the evidence available indicates that from at least Itaboraian time onward the land mammal fauna of Patagonia and northern South America, as well, is a self-contained unit, developing the diversity characteristic of the Paleogene in that continent, including the australidelphian (but South American) microbiotheres. This, in combination with the apparent separation of Australia from Antarctica at ca. 64 Ma, reinforces interpretations that the precursors of the Australian marsupial fauna most likely dispersed from South America to Australia in the late Cretaceous–early Paleocene.     

Patterns of evolutionary transformation in the petrosal bone and some basicranial features in marsupial mammals, with special reference to didelphids

Twelve petrosal and four nonpetrosal characters were coded for representatives of all 15 extant genera of Didelphidae and for 16 additional genera of marsupials representing all extant orders. Three basal metatherians were used as outgroup comparison. Histological sections of a subset of the data were examined. An intermediate position of the hiatus Fallopii supports the monophyly of Didelphidae. Several basicranial regions support different clades within the Didelphidae that recent molecular work has identified, including a sister group relationship of Caluromys and Caluromysiops , the monophyly of large opossums, a Lestodelphys-Thylamys clade, and a Lestodelphys-Thylamys-Gracilinanus-Marmosops clade. Glironia lacks petrosal and jaw synapomorphies of Caluromys and Caluromysiops. The transverse canal, a synapomorphy of the crown-group Marsupialia, opens as a single foramen anterior to the carotid foramen in most marsupials or as numerous foramina in the pterygoid fossa in diprotodontians. It is either intramural (most marsupials) or simply endocranial (most diprotodontians excluding koalas and wombats). Loss of a deep sulcus in the anterior pole of the promontorium for the internal carotid artery and a rostral tympanic process of the petrosal also characterize the groundplan of the crown group Marsupialia. Pouch-young wombats show a groove in the anterior pole of the petrosal for the internal carotid artery. The absence of a prootic canal foramen in the tympanic side of the petrosal of adults supports the monophyly of Australidelphia. Some pouch-young marsupials possess a prootic canal that is later lost in ontogeny. A rather flat promontorium and a crest running medio-distally in the middle of the promontorium characterize Macropodidae.     

Embryo implantation and proteinase activities in a marsupial (Macropus eugenii)

Summary Embryo implantation remains superficial (epithelio-chorial type) in most marsupials including the Macropodidae, but does involve formation of specialized contact zones of the trophoblast with the uterine epithelium. Since in eutherian mammals proteinases appear to play a central role in implantation-initiation mechanisms, a systematic histochemical investigation of proteinase patterns as related to implantation was performed in the tammar wallaby, Macropus eugenii (Macropodidae).Tammar uteri with embryos were collected at diapause and at days 7, 17, 18, 19, 20, 21 and 26 of the 27-day gestational period. Proteinase patterns were studied using a sensitive histochemical gelatin-substrate-film test previously optimized for the detection of trophoblast-dependent proteinase (blastolemmase) in the rabbit. Proteinase patterns were correlated with light-microscopical morphology of the processes of shedding of the extracellular embryo coverings (shell membrane) and attachment of the trophoblast to the uterine epithelium.At acid pH values an intracellular proteinase is detected in yolk sac endoderm and trophoblast as well as in endometrial glands and certain stromal cells. This enzyme is proposed to be a cathepsin indicating high catabolic activity connected particularly with protein transport from the endometrium into the yolk sac. Peak activity is found in the avascular (bilaminar) yolk sac at the phase when contact with the endometrium is being established.A particularly interesting proteinase active at alkaline pH values is detected in the trophoblast-endoderm complex. This enzyme appears to be extruded into the interface between trophoblast and uterine epithelium where it shows maximal activity for only approximately one day, around day (18-)19, exclusively in the bilaminar (avascular) yolk sac. The activity is correlated with the process of shedding of the extracellular embryo coverings (shell membrane) and of subsequent attachment of the trophoblast to the uterine epithelium, in the bilaminar but not the trilaminar (vascular) yolk-sac region. This is the first report on an extracellular (alkaline) proteinase activity possibly serving a specific function in embryo implantation in a marsupial.Abreviations BYS bilaminar (avascular) yolk sac membrane = bilaminar omphalopleure - dp.c. days post coitum - d RPY days after removal of pouch young - TYS trilaminar (vascular) yolk sac membrane = trilaminar omphalopleurePreliminary reports on portions of these investigations were presented at the 14th Annual Meeting of the Society for the Study of Reproduction 1981 (Biol Reprod 24 Suppl 1, p 78 A, 1981) and at the 3. Arbeitstagung der Anatomischen Gesellschaft 1982 (Anat Anz 153, 268, 1983)     

Dentition and Jaw of Kokopellia juddi, a Primitive Marsupial or Near-Marsupial from the Medial Cretaceous of Utah

We add to the knowledge of the dentition and lower jaw of the primitive marsupial or near marsupial, Kokopellia juddi, based on newly collected materials from the medial Cretaceous (Albian–Cenomanian) of central Utah. The dental formula, i4 c1 p3 m4, is primitive for (or with respect to) Marsupialia, as are a number of features of the dentary and dentition: presence of a labial mandibular foramen, ?an inflected angle, ?and a trace of the meckelian groove; lack of “staggering” of the lower incisor series; lack of “twinning” between entoconid and hypoconulid on lower molars; incompletely lingual position of lower molar paraconid; upper molar protocone relatively small and mesiodistally narrow; and conules placed about halfway between the protocone and the paracone–metacone. Other than the stylocone, cusps are lacking from the stylar shelf; we argue that this represents the primitive marsupial condition based on the economy of character change and the stratigraphic record of marsupials in the Cretaceous of North America. Recent discoveries of early marsupials, eutherians, and therians of metatherian–eutherian grade provide data indicating that some derived features of the dentary and dentition (e.g., loss of coronoid, meckelian groove, and labial mandibular foramen; acquisition of strong, “winged” conules, double rank postvallum/prevallid shearing, and stylar cusp D) probably arose independently, in some cases more than once, among the major groups of tribosphenic mammals. In turn, this suggests that a common ancestor for marsupials and placentals was more primitive than has generally been appreciated.     

Relatedness structure detected by microsatellite analysis and attempted pedigree reconstruction in an endangered marsupial, the northern hairy-nosed wombat Lasiorhinus krefftii

The northern hairy-nosed (NHN) wombat is perhaps Australia's most endangered mammal. Being fossorial and nocturnal as well as rare, NHN wombats are difficult to observe in the wild. Hence little is known of their social biology, such as their mating and dispersal systems. A hypothesis has been advanced that adult females of the species disperse post-breeding, leaving their young to inhabit the natal burrow. Female-biased dispersal is expected to result in higher relatedness amongst males in a burrow cluster than amongst females in a burrow cluster. The usefulness of a panel of microsatellite markers in estimating the relatedness structure, and in reconstructing pedigrees for, the sole known population of NHN wombats was assessed. Microsatellite genotypes at eight or nine loci were obtained from 58 of the 85 known individuals, and used to estimate pairwise individual relatedness using Queller & Goodnight's (1989) RELATEDNESS 4.2. Our analysis gave the unexpected result that both males and females were significantly more closely related to their same-sex burrow cluster mates than random, while opposite-sex animals sharing burrows were only slightly (nonsignificantly) more related than random. This raises the possibility of dispersal patterns which lead to association of same-sex relatives. The observed relatedness structure is not expected to make likely a high incidence of inbred matings, as close relatives of the opposite sex are not significantly associated in space. Parentage analysis was attempted using genetic exclusion and LOD likelihood ratios, but proved difficult because of low genetic variation, incomplete sampling of potential parents, and paucity of ecological data such as known mother/offspring pairs and ages of individuals.     

Genetic variation of microsatellite loci in a bottlenecked species: the northern hairy-nosed wombat Lasiorhinus krefftii

We investigate the utility of hypervariable microsatellite loci to measure genetic variability remaining in the northern hairy-nosed wombat, one of Australia's rarest mammals. This species suffered a dramatic range and population reduction over the past 120 years and now exists as a single colony of about 70 individuals at Epping Forest National Park, central Queensland. Because our preliminary research on mitochondrial DNA and multilocus DNA fingerprints did not reveal informative variation in this population, we chose to examine variation in microsatellite repeats, a class of loci known to be highly polymorphic in mammals. To assess the suitability of various wombat populations as a reference for comparisons of genetic variability and subdivision we further analysed mitochondrial DNA cytochrome b sequence, using phylogenetic methods. Our results show that appreciable levels of variation still exist in the Epping Forest colony although it has only 41% of the heterozygosity shown in a population of a closely-related species. From museum specimens collected in 1884, we also assessed microsatellite variation in an extinct population of the northern hairy-nosed wombat, from Deniliquin, New South Wales, 2000 km to the south of the extant population. The apparent loss of variation in the Epping Forest colony is consistent with an extremely small effective population size throughout its 120-year decline.