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Abstract The relatively high level of morphological diversity in Australasian marsupials compared to that observed among American marsupials remains poorly understood. We undertake a comprehensive macroevolutionary analysis of ontogenetic allometry of American and Australasian marsupials to examine whether the contrasting levels of morphological diversity in these groups are reflected in their patterns of allometric evolution. We collate ontogenetic series for 62 species and 18 families of marsupials (n = 2091 specimens), spanning across extant marsupial diversity. Our results demonstrate significant lability of ontogenetic allometric trajectories among American and Australasian marsupials, yet a phylogenetically structured pattern of allometric evolution is preserved. Here we show that species diverging more than 65 million years ago converge in their patterns of ontogenetic allometry under animalivorous and herbivorous diets, and that Australasian marsupials do not show significantly greater variation in patterns of ontogenetic allometry than their American counterparts, despite displaying greater magnitudes of extant ecomorphological diversity.

Abstract Background Self-powered flight is one of the most energy-intensive types of locomotion found in vertebrates. It is also associated with a range of extreme morpho-physiological adaptations that evolved independently in three different vertebrate groups. Considering that development acts as a bridge between the genotype and phenotype on which selection acts, studying the ossification of the postcranium can potentially illuminate our understanding of bat flight evolution. However, the ontogenetic basis of vertebrate flight remains largely understudied. Advances in quantitative analysis of sequence heterochrony and morphogenetic growth have created novel approaches to study the developmental basis of diversification and the evolvability of skeletal morphogenesis. Assessing the presence of ontogenetic disparity, integration and modularity from an evolutionary approach allows assessing whether flight may have resulted in evolutionary differences in the magnitude and mode of development in bats. Results We quantitatively compared the prenatal ossification of the postcranium (24 bones) between bats (14 species), non-volant mammals (11 species) and birds (14 species), combining for the first time prenatal sequence heterochrony and developmental growth data. Sequence heterochrony was found across groups, showing that bat postcranial development shares patterns found in other flying vertebrates but also those in non-volant mammals. In bats, modularity was found as an axial-appendicular partition, resembling a mammalian pattern of developmental modularity and suggesting flight did not repattern prenatal postcranial covariance in bats. Conclusions Combining prenatal data from 14 bat species, this study represents the most comprehensive quantitative analysis of chiropteran ossification to date. Heterochrony between the wing and leg in bats could reflect functional needs of the newborn, rather than ecological aspects of the adult. Bats share similarities with birds in the development of structures involved in flight (i.e. handwing and sternum), suggesting that flight altriciality and early ossification of pedal phalanges and sternum are common across flying vertebrates. These results indicate that the developmental modularity found in bats facilitates intramodular phenotypic diversification of the skeleton. Integration and disparity increased across developmental time in bats. We also found a delay in the ossification of highly adaptable and evolvable regions (e.g. handwing and sternum) that are directly associated with flight performance.

Fluctuating asymmetry (random fluctuations between the left and right sides of the body) has been interpreted as an index to quantify both the developmental instabilities and homeostatic capabilities of organisms, linking the phenotypic and genotypic aspects of morphogenesis. However, studying the ontogenesis of fluctuating asymmetry has been limited to mostly model organisms in postnatal stages, missing prenatal trajectories of asymmetry that could better elucidate decoupled developmental pathways controlling symmetric bone elongation and thickening. In this study, we quantified the presence and magnitude of asymmetry during the prenatal development of bats, focusing on the humerus, a highly specialized bone adapted in bats to perform under multiple functional demands. We deconstructed levels of asymmetry by measuring the longitudinal and cross-sectional asymmetry of the humerus using a combination of linear measurements and geometric morphometrics. We tested the presence of different types of asymmetry and calculated the magnitude of size-controlled fluctuating asymmetry to assess developmental instability. Statistical support for the presence of fluctuating asymmetry was found for both longitudinal and cross-sectional asymmetry, explaining on average 16% of asymmetric variation. Significant directional asymmetry accounted for less than 6.6% of asymmetric variation. Both measures of fluctuating asymmetry remained relatively stable throughout ontogeny, but cross-sectional asymmetry was significantly different across developmental stages. Finally, we did not find a correspondence between developmental patterns of longitudinal and cross-sectional asymmetry, indicating that processes promoting symmetrical bone elongation and thickening work independently. We suggest various functional pressures linked to newborn bats’ ecology associated with longitudinal (altricial flight capabilities) and cross-sectional (precocial clinging ability) developmental asymmetry differentially. We hypothesize that stable magnitudes of fluctuating asymmetry across development could indicate the presence of developmental mechanisms buffering developmental instability.

Sheath-tailed bats (Family Emballonuridae) from the early Pleistocene Rackham’s Roost Site cave deposit in the Riversleigh World Heritage Area, north-western Queensland are the oldest recorded occurrence for the family in Australia. The fossil remains consist of maxillary and dentary fragments, as well as isolated teeth, but until now their precise identity has not been assessed. Our study indicates that at least three taxa are represented, and these are distinguished from other Australian emballonurids based on morphometric analysis of craniodental features. Most of the Rackham’s Roost Site emballonurid remains are referrable to the modern species Taphozous georgianus Thomas, 1915, but the extant species T. troughtoni Tate, 1952 also appears to be present, as well as a very large, as-yet undetermined species of Saccolaimus Temminck, 1838. We identify craniodental features that clearly distinguish T. georgianus from the externally very similar T. troughtoni. Results suggest that the distributions of T. georgianus and T. troughtoni may have overlapped in north-western Queensland since at least the early Pleistocene.

Morphological shifts observed in the fossil record of a lineage potentially indicate concomitant shifts in ecology of that lineage. Mekosuchine crocodiles of Cenozoic Australia display departures from the typical eusuchian body-plan both in the cranium and postcranium. Previous qualitative studies have suggested that these crocodiles had a more terrestrial habitus than extant crocodylians, yet the capacity of mekosuchine locomotion remains to be tested. Limb bone shape, such as diaphyseal cross-section and curvature, has been related to habitual use and locomotory function across a wide variety of taxa. Available specimens of mekosuchine limbs, primarily humeri, are distinctly columnar compared with those of extant crocodylians. Here we apply a quantitative approach to biomechanics in mekosuchine taxa using both geomorphic morphometric and finite element methods to measure bone shape and estimate locomotory stresses in a comparative context. Our results show mekosuchines appear to diverge from extant semi-aquatic saltwater and freshwater crocodiles in cross-sectional geometry of the diaphysis and generate different structural stresses between models that simulate sprawling and high-walk gaits. The extant crocodylians display generally rounded cross-sectional diaphyseal outlines, which may provide preliminary indication of resistance to torsional loads that predominate during sprawling gait, whereas mekosuchine humeri appear to vary between a series of elliptical outlines. Mekosuchine structural stresses are comparatively lower than those of the extant crocodylians and reduce under high-walk gait in some instances. This appears to be a function of bending moments induced by differing configurations of diaphyseal curvature. Additionally, the neutral axis of structural stresses is differently oriented in mekosuchines. This suggests a shift in the focus of biomechanical optimisation, from torsional to axial loadings. Our results lend quantitative support to the terrestrial habitus hypothesis in so far as they suggest that mekosuchine humeri occupied a different morphospace than that associated with the semi-aquatic habit. The exact adaptational trajectory of mekosuchines, however, remains to be fully quantified. Novel forms appear to emerge among mekosuchines during the late Cenozoic. Their adaptational function is considered here; possible applications include navigation of uneven terrain and burrowing.

Quail-thrushes (Passeriformes: Cinclosomatidae: Cinclosoma) are ground-dwelling corvoid songbirds endemic to Australia and New Guinea. Until now, the only known quail-thrush fossils have been from late Quaternary cave deposits in Australia. A new species of quail-thrush, Cinclosoma elachum sp. nov., is described from the early to middle Miocene deposits in the Riversleigh World Heritage Area, Queensland, Australia. A second, larger quail-thrush is identified from the middle Miocene of Riversleigh. The new fossils considerably extend the geographic and temporal ranges of cinclosomatids, and indicate the presence of two species of quail-thrushes in the Miocene of northern Australia, located more than 300 km from the nearest extant member of Cinclosoma. These fossils provide a minimum age of ~18 million years for Cinclosomatidae. They cannot be confidently assigned to the crown group of the genus Cinclosoma, but can be used to calibrate the split between this genus and Ptilorrhoa in molecular dating studies of the Corvides radiation and Passeriformes overall. This material also adds to the growing diversity of songbirds identified from the pre-Pleistocene record of Australia, and reemphasises the global significance of the Riversleigh deposits in developing understanding about the early evolutionary history of passerines.

Instances of dwarfism in the fossil record are of interest to palaeontologists because they often provide insight into aspects of palaeoecology. Fossil species of Australian-Pacific mekosuchine genus Mekosuchus have been described as dwarf, primarily terrestrial crocodiles, in contrast with the nearly ubiquitous semi-aquatic habitus of extant crocodilians (Willis 1997). This hypothesis has been difficult to test because of limited knowledge of the cranial and postcranial skeleton of extinct taxa and the continuous nature of crocodilian growth. New crocodilian vertebral material from Riversleigh, northwestern Queensland, tentatively referred to Mekosuchus whitehunterensis Willis, 1997, displays morphological maturity indicative of adult snout-vent length little over a half-meter, proportionally smaller than extant dwarf taxa. Further, this material displays morphology that indicates a relatively large epaxial neck musculature for its body-size. These attributes suggest this dwarf mekosuchine employed unusual feeding behaviours. The ability to perform normal death-roll, de-fleshing behaviours would be limited in a mekosuchine of such small size. Given the powerful neck muscles and other anatomical features, it is more likely that this mekosuchine killed and/or dismembered its prey using a relatively forceful lifting and shaking of the head.

Australian Oligo–Miocene mekosuchines (Crocodylia; Crocodyloidea) display wide diversity in cranial shape and inferred hunting strategies. Terrestrial habitus has been inferred for these distinctive predators. A direct morphological signal for locomotion can be expected in the postcrania, particularly the pelvic and pectoral girdles. Here we describe fossil materials of the girdles, which chart their morphological variation in the subfamily from Eocene through to Middle Miocene. Over this period, both girdles undergo significant morphological changes. Notably, an enclosed, ventrally orientated acetabulum in the ilium is developed in one lineage. This recapitulates the erect parasagittal configuration of the pelvic limb seen in many Mesozoic crocodylomorph lineages, suggesting consistent use of erect high-walking in these mekosuchines. Other pelves from the same Oligo–Miocene deposits display morphology closer to modern crocodilians, suggesting a partitioning of locomotory strategy among sympatric mekosuchines. Plesiomorphic and derived pelvic girdles are distinguishable by parsimony analysis, and the earliest examples of the mekosuchine pelvis more closely resemble gavialids and alligatorids while latter forms converge on crown group crocodylids in the morphology of the iliac crest. This suggests that a revaluation of the base relationship of Mekosuchinae within Eusuchia is necessary.

Fourteen of the best sampled Oligo-Miocene local faunas from the Riversleigh World Heritage Area, north-western Queensland, Australia are analysed using classification and ordination techniques to identify potential mammalian palaeocommunities and palaeocommunity types. Abundance data for these faunas are used, for the first time, in conjunction with presence/absence data. An early Miocene Faunal Zone B and two middle Miocene Faunal Zone C palaeocommunities are recognised, as well as one palaeocommunity type. Change in palaeocommunity structure, between the early Miocene and middle Miocene, may be the result of significant climate change during the Miocene Carbon Isotope Excursion. The complexes of local faunas identified will allow researchers to use novel palaeocommunities in future analyses of Riversleigh’s fossil faunas. The utility of some palaeoecological multivariate indices and techniques is examined. The Dice index is found to outperform other binary similarity/distance coefficients, while the UPGMA algorithm is more useful than neighbour joining. Evidence is equivocal for the usefulness of presence/absence data compared to abundance.

The New Zealand endemic bat family Mystacinidae comprises just two Recent species referred to a single genus, Mystacina. The family was once more diverse and widespread, with an additional six extinct taxa recorded from Australia and New Zealand. Here, a new mystacinid is described from the early Miocene (19-16 Ma) St Bathans Fauna of Central Otago, South Island, New Zealand. It is the first pre-Pleistocene record of the modern genus and it extends the evolutionary history of Mystacina back at least 16 million years. Extant Mystacina species occupy old-growth rainforest and are semi-terrestrial with an exceptionally broad omnivorous diet. The majority of the plants inhabited, pollinated, dispersed or eaten by modern Mystacina were well-established in southern New Zealand in the early Miocene, based on the fossil record from sites at or near where the bat fossils are found. Similarly, many of the arthropod prey of living Mystacina are recorded as fossils in the same area. Although none of the Miocene plant and arthropod species is extant, most are closely related to modern taxa, demonstrating potentially long-standing ecological associations with Mystacina.

Thylacinidae is an extinct family of Australian and New Guinean marsupial carnivores, comprizing 12 known species, the oldest of which are late Oligocene (∼24 Ma) in age. Except for the recently extinct thylacine (Thylacinus cynocephalus), most are known from fragmentary craniodental material only, limiting the scope of biomechanical and ecological studies. However, a particularly well-preserved skull of the fossil species Nimbacinus dicksoni, has been recovered from middle Miocene (∼16-11.6 Ma) deposits in the Riversleigh World Heritage Area, northwestern Queensland. Here, we ask whether N. dicksoni was more similar to its recently extinct relative or to several large living marsupials in a key aspect of feeding ecology, i.e., was N. dicksoni a relatively small or large prey specialist. To address this question we have digitally reconstructed its skull and applied three-dimensional Finite Element Analysis to compare its mechanical performance with that of three extant marsupial carnivores and T. cynocephalus. Under loadings adjusted for differences in size that simulated forces generated by both jaw closing musculature and struggling prey, we found that stress distributions and magnitudes in the skull of N. dicksoni were more similar to those of the living spotted-tailed quoll (Dasyurus maculatus) than to its recently extinct relative. Considering the Finite Element Analysis results and dental morphology, we predict that N. dicksoni likely occupied a broadly similar ecological niche to that of D. maculatus, and was likely capable of hunting vertebrate prey that may have exceeded its own body mass.

The marsupial family Diprotodontidae (Diprotodontia, Vombatiformes) is a group of extinct large-bodied (60-2500 kg) wombat-like herbivores that were common and geographically widespread in Cenozoic fossil deposits of Australia and New Guinea. Typically they are regarded to be gregarious, terrestrial quadrupeds and have been likened in body form among placental groups to sheep, rhinoceros and hippopotami. Arguably, one of the best represented species is the zygomaturine diprotodontid Nimbadon lavarackorum which is known from exceptionally well-preserved cranial and postcranial material from the middle Miocene cave deposit AL90, in the Riversleigh World Heritage Area, northwestern Queensland. Here we describe and functionally analyse the appendicular skeleton of Nimbadon lavarackorum and reveal a far more unique lifestyle for this plesiomorphic and smallest of diprotodontids. Striking similarities are evident between the skeleton of Nimbadon and that of the extant arboreal koala Phascolarctos cinereus, including the powerfully built forelimbs, highly mobile shoulder and elbow joints, proportionately large manus and pes (both with a semi-opposable digit I) and exceedingly large, recurved and laterally compressed claws. Combined with the unique (among australidelphians) proportionately shortened hindlimbs of Nimbadon, these features suggest adept climbing ability, probable suspensory behaviour, and an arboreal lifestyle. At approximately 70 kg, Nimbadon is the largest herbivorous mammal to have occupied the forest canopies of Australia - an ecological niche that is no longer occupied in any Australian ecosystem and one that further expands the already significant niche diversity displayed by marsupials during the Cenozoic.

BackgroundWe describe new cranial and post-cranial marsupial fossils from the early Eocene Tingamarra Local Fauna in Australia and refer them to Djarthia murgonensis, which was previously known only from fragmentary dental remains.Methodology/principal findingsThe new material indicates that Djarthia is a member of Australidelphia, a pan-Gondwanan clade comprising all extant Australian marsupials together with the South American microbiotheres. Djarthia is therefore the oldest known crown-group marsupial anywhere in the world that is represented by dental, cranial and post-cranial remains, and the oldest known Australian marsupial by 30 million years. It is also the most plesiomorphic known australidelphian, and phylogenetic analyses place it outside all other Australian marsupials.Conclusions/significanceAs the most plesiomorphic and oldest unequivocal australidelphian, Djarthia may approximate the ancestral morphotype of the Australian marsupial radiation and suggests that the South American microbiotheres may be the result of back-dispersal from eastern Gondwana, which is the reverse of prevailing hypotheses.