Evolutionary History of Lagomorphs in Response to Global Environmental Change

Although species within Lagomorpha are derived from a common ancestor, the distribution range and body size of its two extant groups, ochotonids and leporids, are quite differentiated. It is unclear what has driven their disparate evolutionary history. In this study, we compile and update all fossil records of Lagomorpha for the first time, to trace the evolutionary processes and infer their evolutionary history using mitochondrial genes, body length and distribution of extant species. We also compare the forage selection of extant species, which offers an insight into their future prospects. The earliest lagomorphs originated in Asia and later diversified in different continents. Within ochotonids, more than 20 genera occupied the period from the early Miocene to middle Miocene, whereas most of them became extinct during the transition from the Miocene to Pliocene. The peak diversity of the leporids occurred during the Miocene to Pliocene transition, while their diversity dramatically decreased in the late Quaternary. Mantel tests identified a positive correlation between body length and phylogenetic distance of lagomorphs. The body length of extant ochotonids shows a normal distribution, while the body length of extant leporids displays a non-normal pattern. We also find that the forage selection of extant pikas features a strong preference for C₃ plants, while for the diet of leporids, more than 16% of plant species are identified as C₄ (31% species are from Poaceae). The ability of several leporid species to consume C₄ plants is likely to result in their size increase and range expansion, most notably in Lepus. Expansion of C₄ plants in the late Miocene, the so-called ‘nature’s green revolution’, induced by global environmental change, is suggested to be one of the major ‘ecological opportunities’, which probably drove large-scale extinction and range contraction of ochotonids, but inversely promoted diversification and range expansion of leporids.     

Molecular phylogeny and biogeographic history of the European Maja spider crabs (Decapoda, Majidae)

We have assessed for the first time the phylogenetic relationships and biogeographic history of the crabs of the genus Maja that inhabit European coasts: M. brachydactyla, M. crispata, M. goltziana and M. squinado. Using mitochondrial markers, we have recovered a well-resolved phylogenetic tree that supports a single origin for the European species, most likely from an Indo-West Pacific ancestor during the Early Miocene. In this phylogeny, M. goltziana appears as the basal European species, with a sister lineage bifurcating into an Eastern Atlantic (M. brachydactyla) and a Mediterranean (M. crispata and M. squinado) clade. We propose the Tethyan Seaway as the initial colonization route, although an entrance through South Africa cannot be discounted. The Eastern Atlantic/Mediterranean split seems to predate the Messinian salinity crisis, which, in turn, could have promoted the recent divergence within the Mediterranean. In addition, Pleistocene glaciations could explain the current diversity in the Eastern Atlantic Ocean, where a unique mitochondrial lineage is found. According to this, the genetic profile of South African crabs appears to belong to M. brachydactyla, questioning the validity of the putative species M. capensis.     

Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae)

Most of the 29 living species of Potamididae show a close association with mangroves. The trees provide the snails with shelter, protection from predators, a solid substrate and sometimes food. Using sequences from three genes (nuclear 18S rRNA and 28S rRNA, mitochondrial COI) we derive a molecular phylogeny and recognize six living genera (Terebralia, Telescopium, Tympanotonos, Cerithidea, Cerithideopsis, Cerithideopsilla). The oldest modern genera (Terebralia, Cerithideopsis) appeared in the Tethyan realm in the Middle Eocene, shortly after the origin of mangrove trees. Whereas most potamidid genera are now restricted to either the Indo-West Pacific (IWP) or to the eastern Pacific plus Atlantic (EPA), sister clades of Cerithideopsis survive in both realms. Based on a reinterpretation of the fossil record (particularly of the monotypic Tympanotonos and extinct Potamides), and parsimonious reconstruction of ancestral habitats, we suggest that the living potamidids are an adaptive radiation that has always been closely associated with mangroves. The specialized tree-climbing groups Cerithidea and Cerithideopsis were independently derived from mud-dwelling ancestors. Cerithideopsilla cingulata (a species complex in the IWP) and 'Potamides' conicus (in the Mediterranean and Indian Ocean) form a single clade within the genus Cerithideopsilla. This refutes the hypothesis that 'P.'conicus is the sole relict of the Tethyan Potamides that has occurred in the Mediterranean region since the Palaeocene. Instead, the phylogeny and fossil record suggest that an ancestor of Cerithideopsilla conica with planktotrophic larvae dispersed from the IWP to the Mediterranean in the Middle Miocene, that its direct development evolved in the Mediterranean during the Pliocene, and that it reinvaded the Indian Ocean during the Plio-Pleistocene.     

Molecular phylogenies and historical biogeography of a circumtropical group of gastropods (Genus: Nerita): implications for regional diversity patterns in the marine tropics

To determine how historical processes, namely speciation, extinction, and dispersal, have contributed to regional species diversity patterns across the marine tropics, we examined the biogeographical history of a circumtropical genus of intertidal gastropods. A species-level phylogeny of Nerita, representing approximately 87% of extant species, was developed from 1608bp of mitochondrial (COI and 16S) and nuclear (ATPSalpha) markers. Phylogenetic relationships generally corresponded to prior classifications; however, comprehensive sampling revealed a number of previously undetected ESUs. Using the resulting tree as a framework, we combined geographical distributions and fossil evidence to reconstruct ancestral ranges, produce a time-calibrated chronogram, and estimate diversification rates. Analyses revealed two monophyletic eastern Pacific+Atlantic (EPA) clades, each of which likely split from an Indo-West Pacific (IWP) sister clade prior to an early Miocene Tethys Seaway closure. More recent diversification throughout the IWP appears to have been driven by both vicariance and dispersal events; EPA diversity has been further shaped by speciation across the Central American Seaway prior to its closure and dispersal across the Atlantic. Despite the latter, inter-regional dispersal has been rare, and likely contributes little to regional diversity patterns. Similarly, infrequent transitions into temperate regions combined with reduced diversification rates may explain low diversity in West and South Pacific clades. Since origination, Nerita diversification appears remarkably constant, with the exception of a lag in the late Eocene-early Oligocene and elevated rates in the late Oligocene-early Miocene. However, a comparison among regions suggested that IWP clades have experienced, on average, higher rates of speciation. Fossil evidence indicates that the EPA likely witnessed greater extinction relative to the IWP. We propose that regional differences in species diversity in Nerita have been largely shaped by differential rates of speciation and extinction.     

FILLING A GAP: THE FIRST OCCURRENCES OF EPIPHAXUM (CNIDARIA: HELIOPORACEA: LITHOTELESTIDAE) IN THE EOCENE, OLIGOCENE AND MIOCENE

Abstract:  A new species of the genus Epiphaxum (family Lithotelestidae) is described and illustrated in detail, and compared to other species. Epiphaxum arbuscula sp. nov. has been collected from Upper Eocene (Priabonian), Upper Oligocene (Chattian) and Lower Miocene (Upper Burdigalian) deposits of the Aquitaine Basin, south-west France. Epiphaxum is a poorly documented genus but its fossil record extends back to the Late Cretaceous; it was previously known only from the Paleocene (Danian). Epiphaxum arbuscula differs from all others species of the genus in the form of its colony. In contrast to the creeping colonies of previously known species, it has branched colonies. It is very common at one Upper Oligocene outcrop from which an assemblage with submarine cave remains has been described. A close relationship between the three extant species (two from the Caribbean Sea and one from the Indo-West Pacific region) and the Paleogene species is also noted. These constitute a group that has not undergone any important morphological changes for the last 65 million years.     

Late Miocene Survival of a Hyper-Longirostrine Dolphin and the Neogene to Recent Evolution of Rostrum Proportions Among Odontocetes

Hyper-longirostry, the character of having extremely elongated rostra, emerged in the early and middle Miocene among several different clades of echolocating toothed whales (odontocetes) followed by a rapid decline near the end of the middle Miocene, and postdated by a much lower number of occurrences in the late Miocene and Pliocene and a complete absence among extant odontocetes. New finds of unreworked fossils of Xiphiacetus cristatus (Eurhinodelphinidae) in the middle Tortonian Diest Formation in Belgium (9.54–8.8 Ma) allow for the documentation of the survival of a hyper-longirostrine dolphin into the early late Miocene. An extensive dataset of the rostral index (calculated as the ratio between rostral length and condylobasal length) of Neogene and extant odontocetes is compiled and presented here, which facilitates discussion of evolutionary trends of rostrum proportions during a time period spanning 23 million years. Of interest, the iterative survival into the late Miocene of a single different species of hyper-longirostrine dolphins in a number of paleogeographic regions (North Sea Basin, Atlantic Coastal Plain, and probably the southeastern Pacific) is noted, whereas hyper-longirostrine morphologies only seem to re-appear by the late Messinian in the Northeastern Pacific. A correlation between this pattern and a decrease in habitat size for coastal to estuarine dolphins linked to a major sea level drop is tentatively proposed; such a process may also have played a role in the ecological shift in several dolphin families to freshwater habitats.

     

Cryptic speciation and paraphyly in the cosmopolitan bryozoan Electra pilosa--impact of the Tethys closing on species evolution

Cosmopolitan nature of the marine bryozoan Electra pilosa was studied to clarify geographic structure and to outline evolution and phylogeography of the species. Several local populations from the Northeast Atlantic (North Sea and Baltic Sea), Arctic (Barents Sea and White Sea) and Indo-West Pacific (New Zealand) were compared. In addition, we examined the closely related species E. posidoniae from the Mediterranean Sea. Phylogenetic analysis based on both 16S and 18S rDNA indicate that the Indo-West Pacific E. pilosa is a sister species to the Atlantic-Mediterranean clade, with the latter including the species E. posidoniae and the Atlantic population of E. pilosa. The topology of the phylogenetic tree leads us to conclude that E. pilosa is a paraphyletic species group relative to E. posidoniae, and a molecular dating of its divergence is consistent to geologic events associated with the closure of the Tethys Sea.     

DNA evidence for a Paleocene origin of the Alcidae (Aves: Charadriiformes) in the Pacific and multiple dispersals across northern oceans

The Alcidae is a group of marine, wing-propelled diving birds known as auks that are distributed along the coasts of the northern oceans. It has been suggested that auks originated in the Pacific coastal shores as early as the Miocene, and dispersed to the Atlantic either through the Arctic coasts of Eurasia and North America (northern dispersal route), or through upwelling zones in the coastal areas of California to Florida (southern dispersal route), before the closure of the Isthmus of Panama in the Pliocene. These hypotheses have not been tested formally because proposed phylogenies failed to recover fully bifurcating, well-supported phylogenetic relationships among and within genera. We therefore constructed a large data set of mitochondrial and nuclear DNA sequences for 21 of the 23 species of extant auks. We also included sequences from two other extant and one extinct species retrieved from GenBank. Our analyses recovered a well-supported phylogenetic hypothesis among and within genera. Aethia is the only genus for which we could not obtain strong support for species relationships, probably due to incomplete lineage sorting. By applying a Bayesian method of molecular dating that allows for rate variation across lineages and genes, we showed that auks became an independent lineage in the Early Paleocene and radiated gradually from the Early Eocene to the Quaternary. Reconstruction of ancestral areas strongly suggests that auks originated in the Pacific during the Paleocene. The southern dispersal route seems to have favored the subsequent colonization of the northern Atlantic Ocean during the Eocene and Oligocene. The northern route across the Arctic Ocean was probably only used more recently after the opening of the Norwegian Sea in the Middle Miocene and the opening of the Bering Strait in the Late Miocene. We postulate that the ancestors of auks lived in a warmer world than that currently occupied by auks, and became gradually adapted to feeding in cool marine currents with high biomass productivity. Hence, warmer tropical waters are now a barrier for the dispersal of auks into the Southern Hemisphere, as it is for penguins in the opposite direction.     

Antitropical distribution and evolution in the Indo-West Pacific Ocean

Antitropical distributions of continental shelf, Indo-West Pacific species are probably not due to transgression of the tropics during the glacial periods, isothermic submergence, island integration, rising Neogene temperatures, or the Mesozoic dispersal of fragments from a Pacific continental mass. Characteristics of common antitropical patterns, plus information from systematic works on a variety on a variety of animal and plant groups, indicate that the long discarded "relict theory" of Theel (1885) appears to best fit the evidence, for it provides a mechanism whereby antitropical distribution may be brought about. The relict theory is compatible with the concept that the East Indies part of the Indo-West Pacific has been functioning as a center of evolutionary origin. It suggests that antitropical and associated disjunct patterns are produced as an older species, that has spread out to occupy a broad range, loses ground and gradually becomes supplanted by a younger species that had subsequently evolved in the East Indies. As this process goes on, the older species becomes restricted to a few isolated localities on the fringe of its original range. These isolates are often found to the north and south of the equatorial region but may include relict populations at the western edge of the Indian Ocean.     

Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina

A phylogenetic approach to the origin and maintenance of species diversity ideally requires the sampling of all species within a clade, confirmation that they are evolutionarily distinct entities, and knowledge of their geographical distributions. In the marine tropics such studies have mostly been of fish and reef-associated organisms, usually with high dispersal. In contrast, snails of the genus Echinolittorina (Littorinidae) are restricted to rocky shores, have a four-week pelagic development (and recorded dispersal up to 1400 km), and show different evolutionary patterns. We present a complete molecular phylogeny of Echinolittorina, derived from Bayesian analysis of sequences from nuclear 28S rRNA and mitochondrial 12S rRNA and COI genes (nodal support indicated by posterior probabilities, maximum likelihood, and neighbor-joining bootstrap). This consists of 59 evolutionarily significant units (ESUs), including all 50 known taxonomic species. The 26 ESUs found in the Indo-West Pacific region form a single clade, whereas the eastern Pacific and Atlantic species are basal. The earliest fossil occurred in the Tethys during the middle Eocene and we suggest that the Indo-West Pacific clade has been isolated since closure of the Tethyan seaway in the early Miocene. The geographical distributions of all species (based on more than 3700 locality records) appear to be circumscribed by barriers of low temperature, unsuitable sedimentary habitat, stretches of open water exceeding about 1400 km, and differences in oceanographic conditions on the continuum between oceanic and continental. The geographical ranges of sister species show little or no overlap, indicating that the speciation mode is predominantly allopatric. Furthermore, range expansion following speciation appears to have been limited, because a high degree of allopatry is maintained through three to five branching points of the phylogeny. This may be explained by infrequent long-distance colonization, habitat specialization on the oceanic/continental gradient, and perhaps by interspecific competition. In the eastern Pacific plus Atlantic we identify five cases of divergence on either side of the Isthmus of Panama, but our estimates of their ages pre-date the emergence of the Isthmus. There are three examples of sister relationships between species in the western Atlantic and eastern Atlantic, all resulting from dispersal to the east. Within the Indo-West Pacific, we find no geographical pattern of speciation events; narrowly endemic species of recent origin are present in both peripheral and central parts of the region. Evidence from estimated divergence times of sister species, and from a plot of the number of lineages over time, suggest that there has been no acceleration of diversification during the glacio-eustatic cycles of the Plio-Pleistocene. In comparison with reefal organisms, species of Echinolittorina on rocky shores may be less susceptible to extinction or isolation during sea-level fluctuations. The species richness of Echinolittorina in the classical biogeographic provinces conforms to the common pattern of highest diversity (11 species) in the central "East Indies Triangle" of the Indo-West Pacific, with a subsidiary focus in the eastern Pacific and western Atlantic, and lowest diversity in the eastern Atlantic. The diversity focus in the East Indies Triangle is produced by a mosaic of restricted allopatric species and overlap of a few widespread ones, and is the result of habitat specialization rather than historical vicariance. This study emphasizes the plurality of biogeographic histories and speciation patterns in the marine tropics.     

Miocene biochronology and paleoceanography of the North Pacific

Biostratigraphic correlation based on microfossil datum levels, directly or indirectly tied to the paleomagnetic time scale, provides a high resolution time control for the Miocene in the equatorial and middle latitude North Pacific. Faunal changes and abundance fluctuations of planktic foraminiferal species combined with the oxygen Pacific. Faunal changes and abundance fluctuations of planktic foraminiferal species combined with the oxygen isotope record of foraminifers, reveal the paleoclimatic and paleoceanographic history. The planktic foraminiferal assemblage change in the early Miocene, extinction of Oligocene fauna and rise of a highly diverse Neogene fauna, appears to be related to increased water mass stratification in the world oceans presumably resulting from the establishment of circum-Antarctic circulation. An increase in the siliceous productivity in the eastern equatorial Pacific region between 20 and 18 Ma suggests that the vertical and horizontal circulation was intensified at that time. Climates cooled rapidly during the middle Miocene between 14 and 13 Ma suggesting the growth of a major east Antarctic ice sheet. Paleoclimatic conditions remained generally cool, although oscillating, during the late Miocene. In the late early to middle Miocene faunal provincialism developed between low and middle latitudes, and by late Miocene time a distinct provincialism similar to the present was established.     

TRACING BACK THE ORIGIN OF THE INDO-PACIFIC MOLLUSC FAUNA: BASAL TRIDACNINAE FROM THE OLIGOCENE AND MIOCENE OF THE SULTANATE OF OMAN

Abstract:  Two new tridacnine species are described from the Chattian and Aquitanian of the Arabian Peninsula. For these, the new names Omanidacna eos gen. et sp. nov. and Tridacna evae sp. nov. are erected. Omanidacna is interpreted as an Oligocene ancestor of Hippopus , being the oldest record of this tridacnine lineage. The Aquitanian Tridacna evae is the first occurrence of the genus Tridacna . These Arabian taxa imply that the modern tridacnine lineages are rooted in the Palaeogene and early Neogene of the East African-Arabian Province, although their Eocene ancestors, such as Byssocardium , are Western Tethyan taxa. During the Neogene they successfully settled the Indo-Polynesian Province and became typical elements of the entire Indo-West Pacific Region. The tridacnines are thus an example of a successive transformation and gradual eastward dispersal of an originally Tethyan element contributing to late Neogene diversity in the Indo-West Pacific.     

Late Miocene—early Pliocene planktonic foraminiferal biostratigraphy and paleoceanography of low-latitude marine sequences

Plate reorganization and development of polar glaciation are closely associated with the changing climatic conditions of the Cenozoic. The planktonic foraminiferal fauna in three low-latitude DSDP sites (224, Ninety East Ridge; 317B, Manihiki Plateau; 366A, Sierra Leone Rise) has been examined to determine how these regions responded to the late Miocene climatic cooling that has been previously observed in high-latitude regions. It has been proposed that an expansion of Antarctic glaciation and a resultant eustatic regression are associated with this cooling, with the latter being at least partially responsible for the Messinian “salinity crisis” in the Mediterranean.Only one of the sites, 366A, shows any significant faunal change during the late Miocene and early Pliocene (approximately 8-3 Ma). During the late Miocene, there is a decrease in the abundance of species considered to be tropical-subtropical, suggesting the incursion of a cool water mass into the Sierra Leone Rise region during this time. The lack of any major faunal changes at Sites 214 and 317B indicates that the water masses of the low-latitude regions of the Indo-Pacific were relatively unaffected by this cooling in the late Miocene.The three sites also show no evidence for a change in the level of the CCD between the late Miocene and early Pliocene; however, this is most likely due to their equatorial position and shallow water depth. Higher-latitude sites from both the Atlantic and the Pacific reveal a definite shoaling of the CCD during the late Miocene.     

Systematics and evolutionary significance of the small Abrocomidae from the early Miocene of southern South America

Octodontoidea is the most species-rich clade among hystricomorph rodents, and has a fossil record going back to at least the late Oligocene. Affinities of fossils previous to the late Miocene differentiation of the extant families Abrocomidae, Echimyidae and Octodontidae are controversial, essentially because these fossils may share few apomorphies with modern species. In fact, pre-late Miocene representatives of Abrocomidae had not been recognised until very recently. Here we revise the early Miocene genus Acarechimys, originally assigned to Echimyidae, and alternatively to stem Octodontoidea or to Octodontidae. A systematic and parsimony-based phylogenetic analysis of the species traditionally included in Acarechimys showed that this genus is part of stem Abrocomidae. These results are primarily supported by morphology of the mandible and lower molars. Acarechimys is here restricted to three species, A. minutus, A. pulchellus and Acarechimys pascuali sp. nov., while another species, A. constans, is here transferred to a new abrocomid genus. The remaining species were nested within Octodontidae. According to these results, Abrocomidae might have been as diverse as its sister clade Octodontidae-Echimyidae during the late Oligocene–early Miocene. Extinction of this diversity would have resulted in marked loss of evolutionary history, with extant abrocomids being currently restricted to late-diverged euhypsodont representatives.     

An island in the stream: Australia's place in the cosmopolitan world of Indo-West Pacific freshwater shrimp (Decapoda: Atyidae: Caridina)

Mitochondrial DNA sequences were used to investigate phylogenetic and biogeographic relationships among Australian freshwater shrimp from the genus Caridina H. Milne Edwards, 1837 (Atyidae) and congeners from potential source populations throughout the Indo-West Pacific region. Numerous Australian taxa have close evolutionary relationships with non-Australian taxa from locations throughout the region, indicating a diverse origin of the Australian freshwater fauna. This implies many colonisations to or from Australia over a long period, and thus highlights the surprising adeptness of freshwater shrimp in dispersal across ocean barriers and the unity of much of the region's freshwater biota. Interestingly, a study on Australia's other main genus of atyid shrimp, Paratya Miers, 1882, inferred only a single colonisation. A number of potential species radiations within Australia were also identified. This agrees with patterns detected for a large number of Australian freshwater taxa, and so implies a vicariant explanation due to the development of colder, dryer climates during the late Miocene/early Pliocene.     

Biogeographic responses to geodynamics: A key study all around the Oligo–Miocene Tethyan Seaway

Extensive terrestrial exchanges were initiated by the closure of the Tethyan Seaway during the Early Miocene. Proboscideans are among the most prominent African immigrants, which arrived in Eurasia about 19 Ma ago via the “Gomphotherium Landbridge”. Several distinct waves of continental migrations, however, document that the formation of this landbridge was a multiphase process. Until the closure, a marine faunal exchange was enabled via the Mesopotamian Trough and the Zagros Basin, as reflected by contributions of Indonesian corals in the Iranian basins and by the occurrence of “western” gastropods in Pakistan and India. Nevertheless, the emergence of the landbridge was preceded in the marine biosphere by first biogeographic divergences on both sides of the seaway already during Oligocene times (e.g. within the tridacnines and strombids). Around the closure event, the breakdown of biogeographic relations was near-complete and the Proto-Mediterranean faunas bear little in common with those of the Indo-West Pacific Region (IWPR). Some of the discussed examples suggest that the Western Tethys Region (WTR) had acted as centre of origin and diversity during Oligocene and Early Miocene times. After the closure of the seaway, this centre had shifted to the southeast, heralding the enormous biodiversity of the modern IWPR. Some originally WTR elements managed to follow this shift and formed the Miocene stock for the modern IWPR faunas. In contrast, the marine fauna in the Mediterranean cul-de-sac suffered strong impoverishment due to the Miocene cooling, the Messinian Salinity Crisis and the late Pliocene and Pleistocene glacials – a fact which might explain the receptivity of the Mediterranean Sea for Lessepsian migrants.

This synthesis tries to document the practical problem of recognising biogeographic patterns despite the heterochronous developments in different systematic groups, which, in addition, are often obscured by a stratigraphically incomplete and geographically patchy fossil record.     

The European Tertiary Neritiliidae (Mollusca, Gastropoda, Neritopsina): indicators of tropical submarine cave environments and freshwater faunas

The oldest freshwater neritiliid, Neritilia bisinuata , is described from the Middle Eocene of the Loire Basin. Another European species, N. neritinoides , ranging from the Lower Oligocene to Lower Miocene (Upper Burdigalian) is recognized; its habitat appears to have been freshwater, but very close to the sea. Two new marine neritiliid species from the Aquitaine Basin are described: Bourdieria favia sp. nov. from the Upper Oligocene and Pisulinella aucoini  sp. nov. from the Lower Miocene. A third undescribed species from the Lower Miocene is referred to the same family and related to Pisulinella . The Oligocene species has a strong spiral sculpture, a character completely absent in previously known neritiliid species. The genus Agapilia , founded on juvenile N. neritinoides and adult Vitta picta , appears to be a junior synonym of the genus Vitta. The associated occurrence of shells of the families Neritiliidae, Neritopsidae and Pickworthiidae (well-known inhabitants of Indo-West Pacific submarine caves) at Peyrère suggest the first occurrence of a characteristic assemblage of dark submarine caves during the Oligocene. Both factorial analysis and relative abundance show that at Peyrère these families are associated with other cryptic fossils (various gastropods, bivalves, Brachiopoda, corals, Annelida). However, there are indications of other submarine cave assemblages in various Cenozoic deposits from the Palaeocene to the middle Miocene.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 447–467.     

A new Middle Miocene tarsier from Thailand and the reconstruction of its orbital morphology using a geometric-morphometric method

Tarsius is an extant genus of primates endemic to the islands of Southeast Asia that is characterized by enormously enlarged orbits reflecting its nocturnal activity pattern. Tarsiers play a pivotal role in reconstructing primate phylogeny, because they appear to comprise, along with Anthropoidea, one of only two extant haplorhine clades. Their fossils are extremely rare. Here, we describe a new species of Tarsius from the Middle Miocene of Thailand. We reconstructed aspects of its orbital morphology using a geometric-morphometric method. The result shows that the new species of Tarsius had a very large orbit (falling within the range of variation of modern Tarsius) with a high degree of frontation and a low degree of convergence. Its relatively divergent lower premolar roots suggest a longer mesial tooth row and therefore a longer muzzle than in extant species. The new species documents a previous unknown Miocene group of Tarsius, indicating greater taxonomic diversity and morphological complexity during tarsier evolution. The current restriction of tarsiers to offshore islands in Southeast Asia appears to be a relatively recent phenomenon.     

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.