NEW AUSTRALIAN OLIGOCENE TO MIOCENE RINGTAIL POSSUMS (PSEUDOCHEIRIDAE) AND REVISION OF THE GENUS MARLU

Abstract:  Four genera of extinct ringtail possums have been reported from Australian Oligocene to Miocene sediments since 1987. The genus Marlu was described from two species, M. kutjamarpensis and M. praecursor (Woodburne, Tedford and Archer), from the Miocene Kutjamarpu Local Fauna (Leaf Locality) and late Oligocene Wadikali Local Fauna respectively, of northern South Australia. New fossil material referable to this genus has been collected from the Leaf Locality and the Oligocene to Miocene Riversleigh World Heritage Area in northwest Queensland. Three new species, Marlu karya sp. nov. from middle Miocene Riversleigh local faunas and Marlu syke sp. nov. and Marlu ampelos sp. nov. from the Leaf Locality as well as early to middle Miocene Riversleigh local faunas are described. A revision and rediagnosis of the genus and published species are made following the re-examination of referred material in conjunction with the new material. Marlu is characterized by simple dentition and synapomorphies with extant pseudocheirids such as a conjoined postmetacristid and preentocristid on m1 and the loss of the entostylid ridge. New material from middle Miocene Riversleigh deposits has been referred to M. kutjamarpensis , extending the known distribution and age of that species. Re-examination of M. praecursor has revealed the presence of a small m1 entostylid ridge, contributing further to differences between M. praecursor and all other Marlu species and raising the possibility that Marlu is paraphyletic. The new material does not contradict a sister group relationship between ' Marlu ' (excluding M. praecursor ) and the Pliocene–Pleistocene genus Pseudokoala. Homology of the pseudocheirid m1 protostylid identified in species of Paljara , Pildra and Marlu with that observed in extant species is reconsidered. The 'protostylid' of the extinct genera is herein described as the buccal stylid.     

Mammalian lineages and the biostratigraphy and biochronology of Cenozoic faunas from the Riversleigh World Heritage Area,Australia

Lineage zones within nine contemporaneous mammalian lineages represented in the Cenozoic fossil vertebrate record from the Riversleigh World Heritage Area in northern Australia are used to resolve a series of faunal intervals from the Late Oligocene to Late Miocene. The results agree with previous interpretations of Cenozoic biostratigraphy, provide evidence for the presence of time horizons subdividing some of the four previously recognized Riversleigh biochrons A–D and also provide evidence supporting the allocation of a greater number of Riversleigh faunas to intervals within the Riversleigh biochronology than other methods in previous work. This approach demonstrates the biostratigraphical potential of evolutionary lineages.     

A NEW HIPPOSIDERID GENUS (MICROCHIROPTERA) FROM AN EARLY MIOCENE BAT COMMUNITY IN AUSTRALIA

Abstract:  A new genus and species of hipposiderid bat is described from an early Miocene cave deposit (Bitesantennary Site) in the Riversleigh World Heritage fossil property, northern Australia. Eight hipposiderid genera are now recorded from Riversleigh's Miocene sediments: Hipposideros , Brachipposideros, Rhinonycteris , Riversleigha, Xenorhinos , Miophyllorhina , Archerops and Brevipalatus gen. nov. The new taxon appears to be most closely related to Australian endemic Rhinonycteris and Brachipposideros species, but its autapomorphically very short palate distinguishes it from other members of this relatively plesiomorphic group. It is one of eight hipposiderid species recovered from the Bitesantennary Site deposit, and one of 11 recorded from Riversleigh's early Miocene sediments. Compared with modern bat faunas, the early Miocene Riversleigh bat community differs strikingly in its high hipposiderid diversity but may differ less in its overall trophic structure.     

The Calcaneum—On the Heels of Marsupial Locomotion

The potential for making functional interpretations from a single postcranial element for marsupials was investigated through morphometric analysis of the calcanea of 61 extant species from Australia and New Guinea. Extant species were grouped into locomotor categories and a canonical variates analysis was carried out on measurements of their calcanea. A relationship between measurements of the calcanea and the locomotor behavior of species was found, allowing for prediction of locomotor behavior from calcaneum morphometrics. This was applied to fossil marsupial taxa, from early–late Miocene/?Pliocene deposits at Riversleigh, in an attempt to determine their locomotor behavior. Hopping (saltatorial) taxa are distinguished from quadruped terrestrial taxa and taxa capable of climbing (arboreal and scansorial) by their relatively longer tuber calcis and wider calcaneal head, by their dorso-ventrally thicker calcaneal head, and by their calcaneocuboid facet being less steeply angled antero-posteriorly. Taxa capable of climbing are distinguished from quadruped terrestrial taxa by their shorter tuber calcis relative to the calcaneal head and by their smaller calcaneo-astragalar facet. The locomotor categories distinguished in this study (arboreal/scansorial, quadruped terrestrial, and saltatorial) highlight differences between species in their use of available substrates and thus are informative with regards to the structural components of their habitat. The results of this analysis can be used, in combination with other data, to make inferences about the habitats of paleocommunities at Riversleigh through the Miocene. The calcaneum is a dense and very robust element and, therefore, has a good chance of being preserved. This method provides a quick and easy way of inferring locomotion and has a wide potential for application to many fossil deposits because it requires only a single element.     

Australia's first fossil marsupial mole (Notoryctemorphia) resolves controversies about their evolution and palaeoenvironmental origins

Fossils of a marsupial mole (Marsupialia, Notoryctemorphia, Notoryctidae) are described from early Miocene deposits in the Riversleigh World Heritage Area, northwestern Queensland, Australia. These represent the first unequivocal fossil record of the order Notoryctemorphia, the two living species of which are among the world's most specialized and bizarre mammals, but which are also convergent on certain fossorial placental mammals (most notably chrysochlorid golden moles). The fossil remains are genuinely 'transitional', documenting an intermediate stage in the acquisition of a number of specializations and showing that one of these-the dental morphology known as zalambdodonty-was acquired via a different evolutionary pathway than in placentals. They, thus, document a clear case of evolutionary convergence (rather than parallelism) between only distantly related and geographically isolated mammalian lineages-marsupial moles on the island continent of Australia and placental moles on most other, at least intermittently connected continents. In contrast to earlier presumptions about a relationship between the highly specialized body form of the blind, earless, burrowing marsupial moles and desert habitats, it is now clear that archaic burrowing marsupial moles were adapted to and probably originated in wet forest palaeoenvironments, preadapting them to movement through drier soils in the xeric environments of Australia that developed during the Neogene.     

A NEW SPECIES OF THE WOMBAT WARENDJA FROM LATE MIOCENE DEPOSITS AT RIVERSLEIGH, NORTH-WEST QUEENSLAND, AUSTRALIA

Abstract:  A new species of the plesiomorphic wombat Warendja is described. Warendja encorensis sp. nov., the second species to be attributed to this genus, is known from a single fossil locality (Encore Site) at Riversleigh, north-west Queensland. Specimens of W. encorensis are limited to isolated teeth, two mandibular fragments and a maxillary fragment. Two molars preserve the unworn crown morphology. Encore Site has been estimated to be late Miocene in age, making these the only known specimens of Warendja to be clearly older than Pleistocene. Warendja encorensis is also the first described species of wombat from Miocene deposits to which hypselodont teeth have been attributed. It differs from W. wakefieldi in its larger size, distribution of enamel, and in details of the incisor and premolar morphology.     

A New Species of the Basal “Kangaroo” Balbaroo and a Re-Evaluation of Stem Macropodiform Interrelationships

Exceptionally well-preserved skulls and postcranial elements of a new species of the plesiomorphic stem macropodiform Balbaroo have been recovered from middle Miocene freshwater limestone deposits in the Riversleigh World Heritage Area of northwestern Queensland, Australia. This constitutes the richest intraspecific sample for any currently known basal “kangaroo”, and, along with additional material referred to Balbaroo fangaroo, provides new insights into structural variability within the most prolific archaic macropodiform clade – Balbaridae. Qualitative and metric evaluations of taxonomic boundaries demonstrate that the previously distinct species Nambaroo bullockensis is a junior synonym of B. camfieldensis. Furthermore, coupled Maximum Parsimony and Bayesian phylogenetic analyses reveal that our new Balbaroo remains represent the most derived member of the Balbaroo lineage, and are closely related to the middle Miocene B. camfieldensis, which like most named balbarid species is identifiable only from isolated jaws. The postcranial elements of Balbaroo concur with earlier finds of the stratigraphically oldest balbarid skeleton, Nambaroo gillespieae, and suggest that quadrupedal progression was a primary gait mode as opposed to bipedal saltation. All Balbaroo spp. have low-crowned bilophodont molars, which are typical for browsing herbivores inhabiting the densely forested environments envisaged for middle Miocene northeastern Australia.     

Mid-Tertiary elapid snakes (Squamata, Colubroidea) from Riversleigh, northern Australia: early steps in a continent-wide adaptive radiation

Vertebral and cranial remains of elapid snakes have been collected from fossil assemblages at Riversleigh, north-west Queensland, Australia; most are Miocene but one may be late Oligocene and another as young as Pliocene. The oldest specimen (probably the oldest elapid yet known anywhere) is a vertebra that can be referred provisionally to the extant taxon Laticauda (Hydrophiinae, sensu Slowinski and Keogh, 2000), implying that the basal divergences among Australasian hydrophiine lineages had occurred by the early Miocene, in contrast to most previous estimates for the age of this geographically isolated adaptive radiation. Associated vertebrae and jaw elements from a Late Miocene deposit are described as Incongruelaps iteratus nov. gen. et sp., which has a unique combination of unusual derived characters otherwise found separately in several extant hydrophiine taxa that are only distantly related. Associated vertebrae from other sites, and two parietals from a possibly Pliocene deposit, suggest the presence of several other taxa distinct from extant forms, but the amount of material (and knowledge of variation in extant taxa) is currently insufficient to diagnose these forms. The Tertiary elapids of Riversleigh thus appear to be relatively diverse taxonomically, but low in abundance and, with one exception, not referable to extant taxa below the level of Hydrophiinae. This implies that the present diversity of hydrophiine elapids (31 recognized terrestrial genera, and approximately 16 marine) represents the result of substantial extinction as well as the “cone of increasing diversity” that could be inferred from phylogenetic studies on extant forms.     

Australian Oligo-Miocene Mystacinids (Microchiroptera): upper dentition, new taxa and divergence of New Zealand species

The upper dentition of two Australian early Miocene mystacinids, Icaropsparadox and I. aenae, from Riversleigh, Queensland, are described for the first time. Also recognised is a late Oligocene mystacinid from Lake Palankarinna, South Australia. The new fossils help refine understanding about the evolutionary history of mystacinids in Australia, including their temporal and geographical range, possible dietary and roosting habits, and likely separation time of New Zealand mystacinids.     

3D Morphometric Analysis Reveals Similar Ecomorphs for Early Kangaroos (Macropodidae) and Fanged Kangaroos (Balbaridae) from the Riversleigh World Heritage Area,Australia

Understanding feeding ecology of extinct kangaroos is fundamental to understanding the evolution of kangaroos and the Australia paleoenvironment during the Oligo-Miocene. Comparisons with extant species have suggested that the macropodiforms of the Oligo/Miocene (kangaroos and allies) from the Riversleigh World Heritage Area, northern Australia, were predominantly folivorous browsers or fungivores, unlike the majority of extant species. To further test this hypothesis, we investigate the relationship between variation in cranial and mandibular shape of extant and extinct macropodiforms and ecological factors such as diet, locomotion, and body mass using 3D geometric morphometric analysis of 42 living species and eight extinct species from two radiations (the extinct clade of Balbaridae and some early representatives of the extant Macropodidae. Dietary class (fungivore, browser, grazer, and mixed feeder) correlated strongly with variation in cranial shape (20–25% of variance explained). There was also significant association between cranial shape, and both locomotor mode and body mass. In a principal component analysis of shape variation for crania (including the shape of the molar row), Riversleigh macropodiforms cluster with extant folivorous browsers on principal components (PC) 1 and 3, providing support for previous interpretations of these species as browsing kangaroos. However, as a group and regardless of phylogenetic association, the shape centroid of extinct species differs significantly from that of extant species. Riversleigh macropodiforms cluster with regular hoppers or arboreal tree kangaroos, but this may be a result of the correlation between diet and locomotor mode in kangaroos. Their similarity to extant browsers supports previous interpretations of rainforest and woodland environments at Riversleigh during the early and middle Miocene, respectively. Procrustes ANOVA Analysis of the full shape dataset and diet also shows that diet accounts for a significant portion of variation; however, when phylogeny is taken into account these results become nonsignificant. In analyses of dentary shape, some balbarid species cluster with extant mixed feeders, although this may reflect phylogenetic differences rather than ecological signal.

     

A new cockatoo (Psittaciformes: Cacatuidae) from the Tertiary of Riversleigh, northwestern Queensland, and an evaluation of rostral characters in the systematics of parrots

The first Tertiary cockatoo or any psittaciform from Australia is recorded from an incomplete rostrum from the Riversleigh deposits of northwestern Queensland, Australia. It is placed in the extant genus Cacatua but cannot be diagnosed at the specific level. This cockatoo was probably similar in size to the modern C. roseicapilla , with a short, unspecialized crest and probably white plumage. The rostrum is useful in studies of the systematics of parrots. Rostral characters support some traditional subdivisions of the order, while suggesting that others warrant re-examination.     

Herds Overhead: Nimbadon lavarackorum (Diprotodontidae), Heavyweight Marsupial Herbivores in the Miocene Forests of Australia

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.     

Oldest known Varanus (Squamata: Varanidae) from the Upper Eocene and Lower Oligocene of Egypt: support for an African origin of the genus

Abstract: A large collection of lizard vertebrae from northern Africa represents the oldest unambiguous occurrence of the genus Varanus. The fossils come from late Eocene and early Oligocene freshwater deposits of the Fayum, Egypt, an area noted for many significant primate finds. The recovery and identification of this material indicate that the genus Varanus arose in Africa, before dispersing to Australia and Asia. This dispersal occurred prior to the early to mid‐Miocene, by which time fossil Varanus are known from Australia and Eurasia. Although the dispersal route remains unknown, the lizard material reported here supports the hypothesis that a corridor existed allowing freshwater and terrestrial organisms to cross from Africa to Asia.     

Bird and Mammal Footprints From the Tertiary of Navarre (Western Pyrenees)

A rich variety of vertebrate footprints is known from a number of Upper Eocene to Lower Miocene localities of Navarre (western Pyrenees). The sediments were deposited in a wide range of depositional environments, from marginal marine to diversified terrestrial. Abundant bird tracks have been found in the coastal deposits of the Upper Eocene Liedena Sandstone of the Yesa and Itzagaondoa areas. Ciconiiformes-like (Leptoptilostipus pyrenaicus) and Charadriiformes-like (Charadriipeda ichnospp.) footprints have been recognized. Mammal ichnites have been discovered in the Oligocene and Lower Miocene deposits of Navarre. Equoid perissodactyl ichnites similar to those of Plagiolophustipus occur in the Oligocene fluviatile rocks of the Mués Sandstone of Olexoa and the Rocaforte Sandstone near Oibar and Sada. Trackways of entelodontids (Entelodontipus) are known in fluviatile-palustrine beds of the Oligocene Mués Sandstone of Olkotz. Additionally, bird (Charadriiformes-like) tracks are known in fluviatile-palustrine floodplain deposits of the Lower Miocene Ujué Formation of Los Arcos. In the same area, the Desoio and Los Arcos outcrops have also yielded perissodactyl trackways of possible Equoidea. Trackways of rhinocerotids (?) and artiodactyls (possibly Pecoripeda) are described from the Lower Miocene (Ramblian) palustrine limestones marginal to the Lerín Formation of Kaparroso and from alluvial fan deposits of the Uncastillo-Perdón Formation of Altzorritz, respectively.     

Lepadiform and scalpelliform barnacles from the Oligocene and Miocene of the Paratethys sea

Abstract: A review on the Oligocene and Miocene stalked barnacles of the Paratethys Sea is presented. The fauna comprises two lepadiform and eight scalpelliform species. Only two species are known from Oligocene deposits, four are documented from Lower Miocene formations, and another four species occur in the Middle Miocene. Only one Oligocene and one Miocene species occur in two different Paratethyan basins in roughly coeval environments, and only a single Middle Miocene species is also known from the adjacent Mediterranean Sea. This low degree of biogeographical similarity is probably a result from the still low number of publications dealing with this group. Stalked barnacles are often indicating bathyal or at least deep sublittoral environments. Especially, the abundance of the benthic deep‐water lepadiform Poecilasma in evaporitic deposits of the Middle Miocene Badenian Salinity Crisis may serve as new evidence for a deeper marine depositional environment. Scalpellum paratethyianum sp. nov. and Lepas sattmanni sp. nov. are described as new species; Arcoscalpellum renevieri (Mayer and Gümbel in Gümbel, 1861) is introduced as new combination.     

铲齿象一新种在甘肃省党河地区下中新统的发现

记述了甘肃省党河地区的铲齿象化石一新种———党河铲齿象 (Platybelodondangheen sis)。其主要特征是 :下颌联合和下门齿构成宽短的铲形 ;下门齿薄 ,由单层稀少的齿柱组成 ;具较大的p3 ,p4前窄后宽 ,前齿带和跟座较小 ,m1构造简单 ,齿冠较低 ,第一齿脊无前斜脊 ,前齿带和后跟座较小。它是欧亚大陆目前所知时代最早 (早中新世 )、构造最原始的一种铲齿象。     

On the adaptive pattern of "Ramapithecus".

A reconstruction of the adaptive pattern of the Miocene hominoid "Ramapithecus," supported by comparative data and morphological comparisons, is presented. It is based upon the completely known dentition, inferior portion of the face, and on the paleoecology and geographic distribution of "Ramapithecus" bearing deposits. The known adaptive and morphological patterns of the "Ramapithecus" species are almost indistinguishable from those of the contemporary species of Sivapithecus, which has taxonomic priority. The "Ramapithecus" species are therefore referred to Sivapithecus, a dryopithecine genus which apparently radiated throughout Eurasia and Africa during the Middle Miocene.     

Virtual Reconstruction and Prey Size Preference in the Mid Cenozoic Thylacinid,Nimbacinus dicksoni (Thylacinidae,Marsupialia)

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.     

Phylogeography of Cape Verde Island skinks (Mabuya)

The Cape Verde Islands are of volcanic origin with most having appeared between the early Miocene and mid-Pleistocene. They contain six known species of Mabuya skinks. Phylogeographical relationships within and among the relatively widespread taxa M. stangeri, M. spinalis and M. delalandii were inferred, based on approximately 1 kbp of the cytochrome b gene (mitochondrial DNA). Reciprocal monophyly of M. spinalis and M. stangeri was established, which may have arisen from an early Pliocene/late Miocene cladogenetic event. Considerable between-island sequence divergence was detected among M. spinalis, which appears to have colonized the older islands (Sal and Boavista) first. Much lower sequence divergence was found in M. delalandii, indicating a more recent range expansion. Here, evidence points to colonization of the younger islands of Brava and Fogo soon after appearance. There are similarities between some of the described patterns and those seen in lizards from the Canary Islands.