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Didelphidae is the largest New World radiation of marsupials, and is mostly represented by arboreal, small- to medium-sized taxa that inhabit tropical and/or subtropical forests. The group originated and remained isolated in South America for millions of years, until the formation of the Isthmus of Panama. In this study, we present the first reconstruction of the biogeographic history of Didelphidae including all major clades, based on parametric models and stratified analyses over time. We also compiled all the pre-Quaternary fossil records of the group, and contrasted these data to our biogeographic inferences, as well as to major environmental events that occurred in the South American Cenozoic. Our results indicate the relevance of Amazonia in the early diversification of Didelphidae, including the divergence of the major clades traditionally ranked as subfamilies and tribes. Cladogeneses in other areas started in the late Miocene, an interval of intense shifts, especially in the northern portion of Andes and Amazon Basin. Occupation of other areas continued through the Pliocene, but few were only colonized in Quaternary times. The comparison between the biogeographic inference and the fossil records highlights some further steps towards better understanding the spatiotemporal evolution of the clade. Finally, our results stress that the early history of didelphids is obscured by the lack of Paleogene fossils, which are still to be unearthed from low-latitude deposits of South America.

Amniote vertebrates share a suite of extra-embryonic membranes that distinguish them from anamniotes. Other than that, however, their reproductive characteristics could not be more different. They differ in basic ectothermic vs endothermic physiology, in that two clades evolved powered flight, and one clade evolved a protective shell. In terms of reproductive strategies, some produce eggs and others give birth to live young, at various degrees of development. Crucially, endotherms provide lengthy parental care, including thermal and food provisioning—whereas ectotherms seldom do. These differences could be expected to manifest themselves in major differences between clades in quantitative reproductive traits. We review the reproductive characteristics, and the distributions of brood sizes, breeding frequencies, offspring sizes and their derivatives (yearly fecundity and biomass production rates) of the four major amniote clades (mammals, birds, turtles and squamates), and several major subclades (birds: Palaeognathae, Galloanserae, Neoaves; mammals: Metatheria and Eutheria). While there are differences between these clades in some of these traits, they generally show similar ranges, distribution shapes and central tendencies across birds, placental mammals and squamates. Marsupials and turtles, however, differ in having smaller offspring, a strategy which subsequently influences other traits.

Polydolopidae is a family of Palaeogene marsupials recorded from outcrops in southern South America and the Antarctic Peninsula. They are mostly represented by skull fragments or maxillary, dentary and molar remains. A taxonomic and systematic revision is carried out with the inclusion of a phylogenetic analysis encompassing almost every polydolopid species and five marsupial species found to be related to them in previous analyses (Bonapartherium hinakusijum, Epidolops ameghinoi, Microbiotherium tehuelchum, Prepidolops didelphoides and Roberthoffstetteria nationalgeographica). The Polydolopidae was recovered as a monophyletic group, even thoughno resolution about its sister-group can be found. The following genera are recovered: Antarctodolops, Amphidolops, Archaeodolops, Eudolops, Hypodolops gen. nov., Kramadolops, Pliodolops, Pseudolops and two unidentified genera (Gen. et sp. indet 1 and 2). One genus and three new species are recognized. The family appeared at the beginning of the Palaeocene and disappeared during the Early Oligocene. The extinction of the group could be related to climaticdeterioration in the Early Oligocene, when temperature and the humidity dropped, provoking desertification in the region where Polydolopids evolved. Fil: Chornogubsky Clerici, Laura. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina

Two enigmatic, poorly preserved specimens of mammalian upper molars have been recovered from the Eric the Red West locality in Victoria, Australia. They are two of the three known specimens of mamm...

With two recognized species, the extinct (late Eocene) Groeberia is periodically at the centre of systematic disputes. On the basis of its best-preserved specimen, we describe in detail the dental ...

Background Different types of proteins diverge at vastly different rates. Moreover, the same type of protein has been observed to evolve with different rates in different phylogenetic lineages. In the present study we measured the rates of protein evolution in Eutheria (placental mammals) and Metatheria (marsupials) on a genome-wide basis and we propose that the gene position in the genome landscape has an important influence on the rate of protein divergence. Results We analyzed a protein-encoding gene set (n = 15,727) common to 16 mammals (12 Eutheria and 4 Metatheria). Using sliding windows that averaged regional effects of protein divergence we constructed landscapes in which strong and lineage-specific regional effects were seen on the molecular clock rate of protein divergence. Within each lineage, the relatively high rates were preferentially found in subtelomeric chromosomal regions. Such regions were observed to contain important and well-studied loci for fetal growth, uterine function and the generation of diversity in the adaptive repertoire of immunoglobulins. Conclusions A genome landscape approach visualizes lineage-specific regional differences between Eutherian and Metatherian rates of protein evolution. This phenomenon of chromosomal position is a new element that explains at least part of the lineage-specific effects and differences between proteins on the molecular clock rates.

We have made quantitative volumetric analyses of cerebral cortical (pallial) structures in the brains of three species of monotreme (Ornithorhynchus anatinus, Tachyglossus aculeatus, Zaglossus bruijni) and compared the findings with similar measurements in a range of therian mammals (6 marsupials and 50 placentals). We have found that although the iso- and periallocortical grey matter volume of the monotremes is about what would be expected for their brain size, the proportion of iso- and periallocortical white matter in monotremes is substantially lower than that in the forebrains of therians. This suggests that the forebrains of the three monotremes have fewer association, commissural and/or projection connections than those of similarly sized forebrains of therian mammals. We also found that the iso- and periallocortex of the platypus is relatively smooth-surfaced compared to similarly sized brains of therian mammals, with a distinct caudal shift in the positioning of cortical white matter in the forebrain, consistent with expansion of the posterior thalamic radiation. Central laminated olfactory structures (anterior olfactory nucleus and piriform cortex) are large in the tachyglossid monotremes (Tachyglossus aculeatus and Zaglossus bruijni) and large in xenarthran placental mammals, suggesting convergence of the forebrain structure of monotreme formivores with that of similarly specialized therians like the xenarthrans Myrmecophaga tridactyla and Dasypus novemcinctus. Fil: Ashwell, Ken. University of New South Wales; Australia Fil: Gurovich, Yamila. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Centro de Investigación Esquel de Montaña y Estepa Patagóica. Universidad Nacional de la Patagonia "San Juan Bosco". Facultad de Ciencias Naturales - Sede Esquel. Centro de Investigación Esquel de Montaña y Estepa Patagónica; Argentina. University of South Wales; Australia

Kangaroos and wallabies of the Macropus complex include the largest extant marsupials and hopping mammals. They have traditionally been divided among the genus Macropus (with three subgenera: Macropus, Osphranter and Notamacropus) and the monotypic swamp wallaby, Wallabia bicolor. Recent retrotransposon and genome-scale phylogenetic analyses clarify the placement of Wallabia as sister to Notamacropus, with Osphranter and Macropus branching successively deeper. In view of the traditional Macropus concept being paraphyletic, we undertake to resolve the species-level phylogeny and genus-level taxonomy of the Macropus complex. For the first time, we include nuclear and mitochondrial DNA covering all extant species, and the first DNA sequences from the extinct Toolache wallaby (Notamacropus greyi), which we find groups with the black-gloved wallaby (Notamacropus irma). Morphological variation was examined using geometric morphometric methods on three-dimensional surface-scanned skulls. Wallabia skull shape fell close to Notamacropus (or Thylogale when controlling for allometry). We recommend the subgenera Macropus, Osphranter and Notamacropus be elevated to genera, alongside Wallabia, based on comparisons with other established macropodine genera for cranial disparity, ecology and molecular divergence. Our time tree estimates that all four ‘Macropus’ genera diverged close to the Miocene–Pliocene boundary (~6–5 Mya), then diversified coincident with Pliocene expansion of grasslands in Australia.

The northernmost record of Metatheria, Unnuakomys hutchisoni gen. et sp. nov., is described from the Pediomys Point locality in Upper Cretaceous (lower Maastrichtian) strata of the Prince Creek For...

Sparassodont metatherians were the dominant terrestrial mammalian predators during South America's long Cenozoic isolation. This group's early fossil record is very poor, however, particularly for the late Eocene and early Oligocene. Here, we describe a new sparassodont, Chlorocyon phantasma, gen. et sp. nov., based on a specimen from Los Helados, a new locality within the Abanico Formation of the Andean Main Range of central Chile. New 40Ar/39Ar dates at Los Helados bracketing the fossil-bearing level constrain the age of this specimen to 37–36 Ma (late Eocene), indicating that this new taxon likely pertains to the Mustersan South American Land Mammal “Age.” Chlorocyon is the first Paleogene sparassodont reported from Chile and the first sparassodont described from the Abanico Formation. Distinctive features, including a p3 with an anterior edge that is more curved than the posterior edge and the lack of a hypoconulid on m4, suggest that Chlorocyon is a borhyaenoid closely related to Pharsophorus or Plesiofelis, although much smaller. Chlorocyon represents a welcome addition to the sparse record of late Eocene sparassodonts and indicates that the diversity of non-proborhyaenid borhyaenoids prior to the late Oligocene was greater than previously thought.

International audience; Andinodelphys cochabambensis Marshall & Muizon, 1988 is one of the best preserved metatherian species from the early Palaeocene fauna of Tiupampa (Bolivia). It is represented by five almost complete skulls, three of them being securely associated to sub-complete to partial skeleton. Four skulls could be extracted from a block including several intermingled skeletons. The present paper provides a thorough description of the dental, cranial, and dentary anatomy of A. cochabambensis. The cranial anatomy of A. cochabambensis is similar to that of Pucadelphys andinus. The skull of Andinodelphys however differs from that of Pucadelphys in its larger size and proportionally longer rostrum. Other differences include the presence, in Andinodelphys, of large anteriorly protruding I1s, small palatal vacuities, a transverse canal, and a small hypotympanic sinus. Andinodelphys has the same dental formula as Pucadelphys (I 5/4, C 1/1, P 3/3, M4/4), the plesiomorphic condition for metatherians. Furthermore, both genera share the lack a tympanic process of the alisphenoid, a deep groove for the internal carotid artery at the anterior apex of the promontorium, a small prootic canal perforating the lateral edge of the petrosal and opening laterally in the deep sulcus for the prootic sinus, and a vestigial anterior lamina of the petrosal. Dentally Andinodelphys closely resembles Pucadelphys, the two genera differing in the larger size of the former and in the inconstant presence in the former of a twinned stylar cusp C. Although 25% smaller, the cheek teeth of Andinodelphys closely resemble those of Itaboraidelphys camposi from the early Eocene of Itaboraí (Brazil). As far as dental morphology is concerned, both genera are likely to have diverged from a direct common ancestor, probably Andinodelphys-like, with Itaboraidelphys displaying more derived dental structures. Two isolated petrosal from Itaboraí (Type 2 petrosals) are morphologically close to those of Andinodelphys but distinctly larger. In this paper, a previous interpretation including the teeth of Itaboraidelphys and these petrosals in the same taxon is followed. A phylogenetic analysis retrieved Itaboraidelphys as a sister taxon of the clade Pucadelphys + Andinodelphys, thus lending support to inclusion of the former in the Pucadelphyidae. Three sets of parsimony analyses were performed. A first set of analyses (with all characters) retrieved a strict consensus tree with a clade as follows: (pucadelphyids, (deltatheroidans (stagodontids, Gurlin Tsav skull-GTS), sparassodonts)). An implied weighting analysis with the same data matrix placed the stagodontids in an early diverging position but retained a clade (pucadelphyids, (deltatheroidans, (GTS, sparassodonts))), the deltatheroidans, being therefore inserted in the pucadelphydans. This result implies an independent arrival of pucadelphyids and sparassodonts to South America, which consequently must have been present in North America in the Late Cretaceous. Possible North American sparassodonts could be the poorly known genera Atokatheridium and Olklatheridium (currently referred to deltatheroidans) and the pucadelphyids may have been present in the Late Cretaceous of North America with the genus Aenigmadelphys. However, this hypothesis is less parsimonious (with regard to palaeobiogeography) than a single southward migration of an ancestral Pucadelphyda (Pucadelphyidae + Sparassodonta). Because the result of this first set of analyses may have been induced by heavily homoplastic dental characters related to hypercarnivory, a second set of analyses was performed excluding all the dental characters. The strict consensus is poorly resolved but retains monophyletic Marsupialia and Sparassodonta. An implied weighting analysis retrieved a monophyletic Pucadelphyda but split the deltatheroidans, the polyphyly of which is regarded as a possible artefact related to the lack of dental characters. The GTS is sister taxon to Pucadelphyda. Because the polyphyly of deltatheroidans contradicts all previous hypotheses, a third set of analyses has been performed excluding only those molar characters that supported the close relationships of the hypercarnivorous clades (deltatheroids, stagodontids, and sparassodonts). The strict consensus tree retrieved monophyletic deltatheroidans, Marsupialia and sparassodonts. An implied weighting analysis resulted in deltatheroidans forming a paraphyletic stem assemblage of Metatheria and monophyletic Pucadelphyda. The GTS was no longer related to sparassodonts but was the sister taxon of a clade including the North American taxa of the data matrix, Asiatherium, and Marsupialia. This topology, which is favoured here, supports (as well as that of the second set of analyses) a single pucadelphydan southward migration, probably in the Late Cretaceous, with a Tiupampian radiation of South American carnivorous metatherians.

International audience; The phylogenetic relationships of "opossum-like" metatherians, Herpetotheriidae and Peradectidae, continue to be debated, and yet they are key taxa in the evolutionary history of the crown-group Marsupialia. With state-of-the-art technologies such as X-ray computed tomography (CT scanning), it is now possible to access the inner anatomy of the skull and particularly the ear region, which is more and more frequently used in phylogenetic analyses. Here we describe for the first time the petrosal and inner ear anatomy of the type species Peratherium elegans from the early Oligocene of Ronzon, together with Amphiperatherium elegans from the Eocene of the Montmartre Gypsum. A parsimony analysis of cranial (including petrosal), dental and postcranial characters in 25 metatherians (extant and fossil taxa including the four herpetotheriids studied herein) does not support inclusion of herpetotheriids or peradectids within Didelphimorphia. The results indicate the monophyly of Peratherium cuvieri, P. elegans and A. minutum, but the other herpetotheriid in the analysis, Herpetotherium cf. fugax, is more closely related to the crowngroup clade Marsupialia. Peradectids and herpetotheriids, traditionally considered as "opossum-like", are here found to be stem metatherians and stem marsupials, respectively. The hypothesis in favour of an ancient origin of opossums (Didelphidae) is therefore questioned.

The Pediomyiodea has been largely regarded as a Late Cretaceous North American metatherian lineage; however, some studies have suggested that a few South American taxa could be related to this family. Herein, Austropediomys marshalli gen. et sp. nov. from the Itaborai Basin, Brazil (lower Eocene – Itaboraian SALMA), is described. Austropediomys gen. nov. represents the first report of a pediomyoid metatherian in the Paleogene of South America backed by a phylogenetic analysis. Several derived features in the upper molars support its belonging to the Pediomyoidea: presence of accessory ‘conular-like’ structures (here named conuloids) lingual to the conules; supernumerary stylar cusps; asymmetric stylar shelf, with parastylar shelf reduced to a narrow rim in M1; straight centrocrista; short and obliquely oriented prepacrista, which contacts StA in M2–3. The presence of ‘conular-like’ structures lingual to the paraconule and metaconule is identified in Austropediomys gen. nov. and in Late Cretaceous pediomyoids as well, and results in the phylogenetic analysis as a synapomorphy of the Pediomyoidea. Austropediomys gen. nov. is autapomorphyc in the presence of enlarged conuloids on protocristae. The record of a pediomyoid in the Paleogene of South America increases the temporal and biogeographical range of this lineage. Also, our results support the hypothesis that several non-related metatherian lineages dispersed from North to South America via the Caribbean Plate, most probably during the latest Cretaceous, in an island-hopping or sweepstakes model.

The Sparassodonta (Mammalia: Metatheria) were the principal group of carnivorous mammals in Cenozoic South America and an important component of this continent’s terrestrial predator guild for nearly 60 million years. However, knowledge of the evolutionary history of this group is biased towards species larger than 1.5 kg from extra-tropical latitudes. Here, we describe a new, small sparassodont from the late middle Miocene (Serravallian) of Quebrada Honda, Bolivia, a locality in the middle latitudes (southern tropics) of South America (~ 22° S). This species, Australogale leptognathus, gen. et sp. nov., is represented by a partial, well-preserved dentary and can be distinguished from other sparassodonts by a combination of features including a p3 that is much smaller than p2, absence of a posterobasal heel on p2, a conical entoconid, absence of an entocristid on m1, and an entocristid on m2 that is lingual to the trigonid. This new species is among the smallest Neogene sparassodonts, with an estimated body mass of ~840 g. Australogale leptognathus expands the known taxonomic and morphological diversity of late middle Miocene sparassodonts and provides evidence that the group may have been as diverse during the late middle Miocene as during the late Oligocene and early Miocene.

Molecular estimates of the divergence of placental and marsupial mammals and their broader clades (Eutheria and Metatheria, respectively) fall primarily in the Jurassic period. Supporting these estimates, Juramaia-the oldest purported eutherian-is from the early Late Jurassic (160 million years ago) of northeastern China. Sinodelphys-the oldest purported metatherian-is from the same geographic area but is 35 million years younger, from the Jehol biota. Here we report a new Jehol eutherian, Ambolestes zhoui, with a nearly complete skeleton that preserves anatomical details that are unknown from contemporaneous mammals, including the ectotympanic and hyoid apparatus. This new fossil demonstrates that Sinodelphys is a eutherian, and that postcranial differences between Sinodelphys and the Jehol eutherian Eomaia-previously thought to indicate separate invasions of a scansorial niche by eutherians and metatherians-are instead variations among the early members of the placental lineage. The oldest known metatherians are now not from eastern Asia but are 110 million years old from western North America, which produces a 50-million-year ghost lineage for Metatheria.

The origin of the South American Sparassodonta has always been debated. Current data indicate a North American origin for this group, but no definitive fossil evidence was recovered so far to prove this hypothesis. Herein, Varalphadon janetae sp. nov. is described from the upper Cenomanian and lower Coniacian of southern Utah, which represents the oldest known Sparassodonta. This taxon can be considered to represent Varalphadon as it presents a markedly mesiodistally compressed protocone and M3; small StA, flattened StD, slightly labial contact of the cristid obliqua, labially salient protoconid and lacks StC. It differs from other species of the genus in the smaller size, proportionally narrower protoconal area, paracone slightly more labial than metacone in M3, more labiolingually compressed StD, keeled metaconid and narrower hypoconid and talonid basin. Varalphadon shares with other sparassodonts the presence of vestigial cuspules between StB and StD, flattened and poorly developed StD, absent StC, straight centrocrista, crested conules and an m1 markedly smaller and strongly labiolingually compressed. The phylogenetic analysis recovered Varalphadon as a basal Sparassodonta, corroborating previous studies that considered this taxon as more basal than remaining “Alphadontidae”, which is represented by Protalphadon, Eoalphadon, Alphadon, Albertatherium and Nortedelphys. “Varalphadon” crebreforme is recovered as a Hatcheriformes. The results demonstrate that Metatheria originates in North America during the late Early Cretaceous, with Gondwanan lineages evolving in the Northern Hemisphere during the Late Cretaceous prior to their dispersal to South American millions of years later (late Maastrichtian?). The analysis recovered Metatheria as a Gondwanan invasive lineage.

The Quebrada de Los Colorados Formation (Los Cardones National Park, Salta Province, north-western Argentina), is an Eocene–Oligocene unit well represented in the Calchaquí Valley area. Here we describe a new metatherian association recorded from the base of this formation, inferred as middle Eocene. Represented taxa are: Sparassodonta, family indet.: Patene simpsoni; Polydolopimorphia, Bonapartheriiformes, Argyrolagoidea: family, genus, and species indet.; and Bonapartherioidea, Prepidolopidae: Punadolops alonsoi and Coloradolops cardonensis gen. et sp. nov. Patene simpsoni is also found in the Tonco Valley, near Los Cardones, and at São José de Itaboraí, Rio de Janeiro, Brazil. Punadolops alonsoi was originally described from the Geste Formation in Salta and Catamarca provinces. Hitherto, no other Argyrolagoidea have been found in north-western Argentina in the Palaeogene. Coloradolops cardonensis is unique to the locality and levels here described. The preservation of part of the skull constitutes the best prepidolopid remains known. North-western Argentina is a key region for the study of the evolution of South American mammals, since it has a singular geographical location between the Neotropics and the southernmost part of South America. Due to biogeographical constraints, it may lead to a better comprehension of the continental distribution of metatherian lineages during Cenozoic times. Fil: Chornogubsky Clerici, Laura. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; Argentina. Universidad Nacional de Luján. Departamento de Ciencias Básicas; Argentina Fil: Zimicz, Ana Natalia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Salta. Instituto de Bio y Geociencias del NOA. Universidad Nacional de Salta. Facultad de Ciencias Naturales. Museo de Ciencias Naturales. Instituto de Bio y Geociencias del NOA; Argentina Fil: Goin, Francisco Javier. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Paleontología Vertebrados; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Fernicola, Juan Carlos. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; Argentina Fil: Payrola Bosio, Patricio Augusto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Salta. Instituto de Bio y Geociencias del NOA. Universidad Nacional de Salta. Facultad de Ciencias Naturales. Museo de Ciencias Naturales. Instituto de Bio y Geociencias del NOA; Argentina Fil: Cardenas, Magali. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigación en Paleobiología y Geología; Argentina

One major innovation of mammals is the tribosphenic molar, characterized by the evolution of a neomorphic upper cusp (=protocone) and a lower basin (=talonid) that occlude and provide shearing and crushing functions. This type of molar is an evolutionarily flexible structure that enabled mammals to achieve complex dental adaptations. Among carnivorous mammals, hypercarnivory is a common trend that evolved several times among therians (marsupials, placentals, and stem relatives). Hypercarnivory involves an important simplification of the carnassial molar pattern from the ancestral tribosphenic molar pattern, with the modification of the triangular tooth crown, and the loss of several cusps and cuspids typical of the tribosphenic molar. These losses confer to the molars of the hypercarnivorous mammals a plesiomorphic/paedomorphic morphology that resembles more the earliest mammaliaforms than the earliest therians. Here, we demonstrate that the modification of the molar morphology is fully explained by a patterning cascade mode of cusp development. Contrary to what was previously proposed, our study concludes that the metaconid (mesiolingual cusp of lower molars, associated with a puncturing function) does not influence cusp development of the talonid (distal crushing heel of lower molars). Moreover, it provides a new example of how heterochronic changes were crucial to the evolution of mammal dentition. To overcome the difficulty of applying behavioral or ecological definitions of diets to fossil animals, we characterize hypercarnivorous dentitions on the basis of the molar morphology and more particularly on the loss or retention of crushing structures, each dentition resulting from adaptations to a distinct ecomorphotype. Despite repeated and convergent evolution of hypercarnivorous forms, hypercarnivory appears as a highly constrained specialization (i.e., "dead end") that is unlikely to evolve back to omnivorous dentition, especially when the crushing structures are lost.

The Paleogene vertebrate-bearing sites of Brazil are restricted to few formations such as Maria Farinha (Late Cretaceous-early Paleocene), Itaborai (early Eocene), Entre-Corregos (Eocene-Oligocene), and Tremembe (late Oligocene-early Miocene). A new Paleogene fauna is reported from an outcrop of the Guabirotuba Formation located at Curitiba, Parana State, southern Brazil. The Guabirotuba Formation is a stratigraphic unit of the Curitiba Basin, which belongs to the Cenozoic continental rift system of southeastern Brazil, a predominantly half-graben and horst system. The sedimentary facies indicate a fluvial distributary depositional system associated with floodplain, generated under humid climate conditions alternating with drier periods. This new Paleogene fauna, here named the Guabirotuba Fauna, has invertebrate and vertebrate taxa, including mammals (Cingulata, Notoungulata, Astrapotheria, and Metatheria). The fossil remains of seven armored xenarthrans are identified, including the description of a new species and genus named Proeocoleophorus carlinii. The Guabirotuba ungulates are assigned to Interatheriidae, Oldfieldthomasiidae, and Astrapotheria. The metatherian mammals are represented by a sparassodont, a palaeothentoid, and an argyrolagoid. This new mammalian fauna preliminarily suggests a late middle Eocene age (Barrancan South American Land Mammal Age) for the Guabirotuba Formation.

Mammals and birds provide food for their young after birth, but such provisioning is almost absent in other vertebrates. A recent theory suggested that, in addition to thermoregulation, the large discrepancy in size between adult and young ectothermic vertebrates precludes them from providing for their young, whereas the relatively large offspring of endotherms are easier to provision. I show here that reptile neonates and hatchlings are as large as those of mammals and birds. Differences in size between adults and young thus cannot explain the lack of parental provisioning in reptiles. I suggest that the large size at birth is the ancestral condition in amniotes as a whole and that provisioning has thus evolved after endothermy.

Argyrolagus constitutes, both for its craniodental and postcranial anatomy, one of the most notably specialized South American Neogene metatherians. Differentiating it from any other South American mammal, bipedal jumping has been proposed for Argyrolagus, even though this hypothesis was not supported by morphofunctional studies. Here, we describe the postcranium of A. scaglai (from the Pliocene of Argentina), perform a functional analysis, and interpret it against a varied background of locomotor adaptations of extant mammals. The configuration of joints, the degree of development and location of muscular insertions were mainly analyzed, and functional indices were evaluated. This study indicates that Argyrolagus had stabilized glenohumeral and humeroulnar joints, a great development of the arm retractors, flexors-extensors of the digits, pronator, and supinator muscles, low restrictive humeroradial joint, powerful extensor muscles of the hip, knee, and ankle, good development of the iliac muscle, and restrictive hind limb joints. Joint configurations are interpreted to be optimal to resist the impacts during jumping, avoiding dislocation, compatible with digging activity. A compromise between the capacities to dig and manipulate objects is inferred. It is concluded that Argyrolagus had bipedal jumping locomotion as well as good capacity to dig, constituting an astonishing case of convergence with the small bipedal rodents and small Australian macropodids. We suggest that bipedal jumping in Miocene and Pliocene argyrolagids should not be necessarily related to a particular arid environment. Finally, we evaluate the importance of postcranial features to understand the phylogenetic relationships of Argyrolagidae in a comprehensive phylogeny of Metatheria., Laboratorio de Sistemática y Biología Evolutiva

Marsupials and their fossil relatives, which collectively comprise Metatheria, have been of scientific interest for centuries, with many aspects of their evolution and systematics subject to intense research and debate. Here, we review progress over the last 25 years, which has included the description of many new species (modern and fossil), and major improvements in understanding of their phylogenetic relationships, as well as the overall evolutionary history and biogeography of Marsupialia (crown-clade) and Metatheria (total-clade). Significant advances have included the deployment of increasingly sophisticated molecular, morphological, and total evidence analyses, which have resolved most previously disputed relationships among and within the modern marsupial orders. A broad systematic consensus is now emerging, although several major areas of contention remain, particularly among fossil metatherians. New modern species continue to be described at an impressive rate, with almost 50 named in the last 25 years, and many more await discovery. There has also been an explosion in the discovery and description of fossil marsupials and non-marsupial metatherians (~270 species), principally from Australasia and the Americas but also from Antarctica, Europe, and Asia. Most are represented by dental specimens only, but some consist of complete and well-preserved material, which has led to major improvements in our understanding of the evolution of cranial and postcranial morphology. Improvements in the fossil record and advances in methods for inferring divergence times have helped clarify when and where key events occurred in metatherian evolution, and the patterns of subclade diversification. We also have improved understanding of biogeographical relationships among metatherians on different landmasses. Despite enormous progress, numerous key uncertainties remain due to major gaps in the fossil record (e.g., Antarctica, Late Cretaceous, and early Paleogene of Australia) and a comparative lack of studies that directly combine molecular and fossil data. Future advances will largely depend on improvements in the fossil record and studies that better integrate neontological and paleontological evidence.

International audience; The global palaeobiogeography of early marsupialiform mammals is still poorly understood due to a meagre fossil record outside western North America. Here, two isolated teeth of a marsupialiform mammal from the lowermost Upper Cretaceous (lower Cenomanian) of southwestern France are described and referred to Stagodontidae, a dentally specialized group whose unambiguous members are so far known exclusively from the Upper Cretaceous of North America. The occurrence of stagodontids in Europe represents additional evidence of faunal exchanges between the two landmasses during the latest Albian–early Cenomanian interval, around 100 million years ago, and shows that early (mid-Cretaceous) marsupialiforms were more widely distributed than previously thought

Paucituberculatan marsupials, particularly members of the family Palaeothentidae, were important components of South American mammal communities during much of the Cenozoic. However, after the late early Miocene, palaeothentid remains are rare in the fossil record, and the group is last recorded at late middle Miocene sites in Colombia, Bolivia and possibly Argentina. Here, we describe new specimens of palaeothentids from one of these late middle Miocene sites, Quebrada Honda, Bolivia, which include: (1) the first described lower dentitions of Acdestis maddeni, which exhibit distinctive features such as a greatly elongated paracristid and a single-rooted m4; (2) the first described late middle Miocene palaeothentines, representing two new species of Palaeothentes, P. serratus sp. nov. and P. relictus sp. nov., distinguished from other species of Palaeothentes by the presence of an anterobasal cingulid and reduced anterior trigonid crest, among other features; and (3) remains of a third new species, Chimer...

Recently discovered specimens of the marsupialHerpetotherium merriami(Stock and Furlong, 1922) from the John Day Formation, Oregon, are described. The species was previously known only from a single (type) specimen. These additional specimens have allowed for a revised diagnosis of the species based on the presence of an additional stylar cusp on the upper molars, as well as the relative size of the stylar cusps. This new material also allows for an examination of the variability within the species and establishes a time range for the species, which extends through most of the Arikareean (late Oligocene: Ar1–Ar3: late Rupelian-Aquitanian).Herpetotherium merriamiis distinct within the genus because it is among the larger of the species, but one of the latest occurring, whereas the general pattern of the genus is a reduction in size through time.

A state-of-the-art review of the Cenozoic fossil record from Western Amazonia is provided, based on literature and new data (regarding Paleogene native ungulates). It allows summarizing the evolution and dynamics of middle Eocene–Holocene mammalian guilds, at the level of species, families, and orders. Major gaps in the Western Amazonian mammal record occur in the pre-Lutetian and early Miocene intervals, and in the Pliocene epoch. Twenty-three orders, 89 families, and 320 species are recognized in the fossil record, widely dominated by eutherians from the middle Eocene onward. Probable Allotheria (Gondwanatheria) occur only in the earliest interval, whereas Metatheria and Eutheria are conspicuous components of any assemblage. Taxonomic diversity was probably fairly constant at the ordinal level (~12–14 orders in each time slice considered) and much more variable in terms of family and species richness: if most intervals are characterized by 40–50 co-occurring species and 19–31 co-occurring families, the early Miocene period illustrates a depauperate fauna (21 species, 17 families), strongly contrasting with the late Miocene climactic guild (82 species, 38 families). Recent mammalian taxonomic diversity from Western Amazonia (12 orders, 37 families, and 286 species) is at odds with all past intervals, as it encompasses only three orders of South American origin (Didelphimorphia, Cingulata, and Pilosa) but four North American immigrant orders (Artiodactyla, Perissodactyla, Carnivora, and Lagomorpha). In terms of taxonomic diversity, recent mammalian guilds are fully dominated by small-sized taxa (Chiroptera, Rodentia, and Primates). This overview also confirms the scarcity of large mammalian flesh-eaters in ancient Neotropical mammalian assemblages. The pattern and the timing of mammalian dispersals from northern landmasses into Western Amazonia are not elucidated yet. Fil: Antoine, Pierre Olivier. Université de Montpellier; Francia Fil: Salas Gismondi, Rodolfo. Université de Montpellier; Francia. Universidad Nacional Mayor de San Marcos; Perú Fil: Pujos, François Roger Francis. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mendoza. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales. Provincia de Mendoza. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales. Universidad Nacional de Cuyo. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales; Argentina Fil: Ganerød, Morgan. Geological Survey of Norway; Noruega Fil: Marivaux, Laurent. Université de Montpellier; Francia

The musculoskeletal system of marsupial mammals has numerous unusual features beyond the pouch and epipubic bones. One example is the widespread absence or reduction (to a fibrous “patelloid”) of the patella (“kneecap”) sesamoid bone, but prior studies with coarse sampling indicated complex patterns of evolution of this absence or reduction. Here, we conducted an in-depth investigation into the form of the patella of extant marsupial species and used the assembled dataset to reconstruct the likely pattern of evolution of the marsupial patella. Critical assessment of the available literature was followed by examination and imaging of museum specimens, as well as CT scanning and histological examination of dissected wet specimens. Our results, from sampling about 19% of extant marsupial species-level diversity, include new images and descriptions of the fibrocartilaginous patelloid in Thylacinus cynocephalus (the thylacine or “marsupial wolf”) and other marsupials as well as the ossified patella in Notoryctes ‘marsupial moles’, Caenolestes shrew opossums, bandicoots and bilbies. We found novel evidence of an ossified patella in one specimen of Macropus rufogriseus (Bennett’s wallaby), with hints of similar variation in other species. It remains uncertain whether such ossifications are ontogenetic variation, unusual individual variation, pathological or otherwise, but future studies must continue to be conscious of variation in metatherian patellar sesamoid morphology. Our evolutionary reconstructions using our assembled data vary, too, depending on the reconstruction algorithm used. A maximum likelihood algorithm favours ancestral fibrocartilaginous “patelloid” for crown clade Marsupialia and independent origins of ossified patellae in extinct sparassodonts, peramelids, notoryctids and caenolestids. A maximum parsimony algorithm favours ancestral ossified patella for the clade [Marsupialia + sparassodonts] and subsequent reductions into fibrocartilage in didelphids, dasyuromorphs and diprotodonts; but this result changed to agree more with the maximum likelihood results if the character state reconstructions were ordered. Thus, there is substantial homoplasy in marsupial patellae regardless of the evolutionary algorithm adopted. We contend that the most plausible inference, however, is that metatherians independently ossified their patellae at least three times in their evolution. Furthermore, the variability of the patellar state we observed, even within single species (e.g. M. rufogriseus), is fascinating and warrants further investigation, especially as it hints at developmental plasticity that might have been harnessed in marsupial evolution to drive the complex patterns inferred here.

Thylacosmiline sparassodonts (previously recognized as thylacosmilids) are among the most iconic groups of endemic South American Cenozoic mammals due to their distinctive morphology and convergent resemblance to saber-toothed placental carnivores. However, the early evolution of this group and its relationship to other sparassodonts remains poorly understood, primarily because only highly specialized Neogene taxa such as Thylacosmilus, Anachlysictis, and Patagosmilus are well known. Here, we describe a new Paleogene sparassodont, Eomakhaira molossus, from the Cachapoal locality of central Chile, the first sparassodont reported from early Oligocene strata of the Abanico Formation. Eomakhaira shares features with both Neogene thylacosmilines and Paleogene “proborhyaenids,” and phylogenetic analyses recover this taxon as sister to the clade of Patagosmilus + Thylacosmilus. This broader clade, in turn, is nested within the group conventionally termed Proborhyaenidae. Our analyses support prior hypotheses of a close relationship between thylacosmilines and traditionally recognized proborhyaenids and provide the strongest evidence to date that thylacosmilines are proborhyaenids (i.e, the latter name as conventionally used refers to a paraphyletic group). To reflect the internestedness of these taxa, we propose use of Riggs' (1933) original name Thylacosmilinae for the less inclusive grouping and Proborhyaenidae for the more inclusive one. Saber teeth arose just once among metatherians (among thylacosmilines), perhaps reflecting a developmental constraint related to nonreplacement of canines in metatherians; hypselodonty may have relaxed this potential constraint in thylacosmilines. The occurrence of Eomakhaira in strata of early Oligocene age from the Chilean Andes demonstrates that the stratigraphic range of thylacosmilines spanned almost 30 million years, far surpassing those of saber-toothed placental lineages.

The Eocene-Oligocene boundary (EOB) marks a period of remodeling in the metatherian faunas of South America. Paucituberculata was one of the groups that successfully diversified as the climate and environment conditions changed, and they became, during the first part of the Neogene, an important component of micromammal assemblages. Among paucituberculatans, the non-pichipilid palaeothentoids (NPP) has been recognized as the clade that diversified most widely in post-EOB times. Here we explore the evolutionary response of the NPP to the climatic-environmental changes around the EOB, by analysing the temporal patterns of disparity, taxonomic diversity and body mass in a phylogenetic context. To asses the magnitude of the NPP radiation comparisons based on these macroevolutionary parameters were done with its sister-group Pichipilidae, and its next closest relative, the Caenolestidae. In all considered parameters, NPP reached values significantly higher than the remaining paucituberculatans clades. From its initial diversification in the middle Eocene, taxonomic diversity increased through time, but it was decoupled from disparity across the EOB, and from the late Oligocene to early Miocene. The Oligocene emerges as the key period in NPP evolution, which is evidenced by a significant and concordant expansion of disparity and taxonomic diversity, suggesting evolution into empty ecospace., Facultad de Ciencias Naturales y Museo

This study investigates whether terrestrial mammalian carnivore guilds of ancient South America, which developed in relative isolation, were similar to those of other continents. We do so through analyses of clade diversification, ecomorphology and guild structure in the Sparassodonta, metatherians that were the predominant mammalian carnivores of pre-Pleistocene South America. Body mass and 16 characters of the dentition are used to quantify morphological diversity (disparity) in sparassodonts and to compare them to extant marsupial and placental carnivores and extinct North American carnivoramorphans. We also compare trophic diversity of the Early Miocene terrestrial carnivore guild of Santa Cruz, Argentina to that of 14 modern and fossil guilds from other continents. We find that sparassodonts had comparatively low ecomorphological disparity throughout their history and that South American carnivore palaeoguilds, as represented by that of Santa Cruz, Argentina, were unlike modern or fossil carnivore guilds of other continents in their lack of mesocarnivores and hypocarnivores. Our results add to a growing body of evidence highlighting non-analogue aspects of extinct South American mammals and illustrate the dramatic effects that historical contingency can have on the evolution of mammalian palaeocommunities.

International audience; he present paper describes a disarticulated skull of Allqokirus australisMarshall & Muizon, 1988, a basal sparassodont (Metatheria, Mammalia) from the early Palaeocene (c. 65 Ma.) of Tiupampa (Bolivia). The specimen includes the rostrum and palate with right premaxilla, both maxillae, left lacrimal, palatines and most upper teeth. The second largest element includes the frontals, the left squamosal, the parietals, the supraoccipital, the basisphenoid, the presphenoid, the alisphenoid, and part of the pterygoids. The nasals, basioccipital and exoccipitals are missing. Other elements are the left petrosal, the right jugal and squamosal, and both dentaries. The elements of the specimen allow for a good reconstruction of the skull, which is thoroughly described and compared to that of other sparassodonts and to the Tiupampa pucadelphyids, Pucadelphys and Andinodelphys. The dental morphology of Allqokirus australis is extremely similar to that of Patene simpsoni from the early Eocene of Itaboraí (Brazil) and presents distinct (although incipient) carnivorous adaptations. Furthermore, some characters of the ear region (e.g. medial process of the squamosal, deep groove for the internal carotid artery at the ventral apex of the petrosal) are also present in most other sparassodonts and in the pucadelphyids from the same locality. A parsimony analysis performed on the basis of a data matrix of 364 characters and 38 taxa placed Allqokirus in a sparassodont clade (the Mayulestidae) that also included Mayulestes and Patene. This family constitutes the sister group of all other sparassodonts. Our analysis also retrieved a large clade composed of the sparassodonts and the pucadelphyids, formally named Pucadelphyda n. superord. This superorder represents the large metatherian carnivorous radiation of the Tertiary of South America, which is first known at Tiupampa, and which started to diversify probably slightly earlier, during the late Cretacous in South America. So far, no representative of Pucadelphyda has been discovered in North America. At Tiupampa, Allqokirus and Mayulestes are the largest metatherians of the fauna and they fill the predaceous mammalian ecological niche. They are the earliest representatives of Sparassodonta, a successful metatherian carnivorous radiation which persisted in South America until the late Pliocene, i.e., during more than 63 Ma.

Sparassodonta is a diverse group of extinct metatherian predators that include forms with diets ranging from omnivores to hypercarnivores, including potential bone-crushers and sabre-Tooth specialised species. Most of the previous dietary studies on the group were based on qualitative approaches or dental morphometric indexes and/or bite force estimations. In this study, we explore the evolution of mandible shape and diet of Sparassodonta in a comparative phylogenetic framework, using geometric morphometric tools and allometric and discriminant analyses. We analysed the mandible shape of 142 extant species of marsupials and placental carnivores, and 15 fossil sparassodont species. We found that the relationship between shape and size of the mandible is strongly structured by phylogeny, where the more derived borhyaenoids tend to possess stronger and larger mandibles. Derived borhyaenoid sparassodonts and basal borhyaenoids were classified as hypercarnivores (with short and robust mandibular body). Hathliacynid were classified as mesocarnivores or as hypercarnivores, but with lower probabilities and less specialised morphologies (with a long and slender mandible). Although dental morphology suggests that most of the species of Sparassodonta would have been hypercarnivores, the robustness of the mandible seems to be informative regarding the prey size and degree of specialisation. The relationship between mandibular size and shape, and talonid/trigonid relative size, is strongly influenced by the phylogenetic legacy, suggesting that ecological factors could have influenced the evolution of the sparassodonts. Fil: Echarri, Sebastián. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales ; Argentina Fil: Ercoli, Marcos Darío. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Universidad Nacional de Jujuy. Instituto de Ecorregiones Andinas. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Salta. Instituto de Ecorregiones Andinas; Argentina. Universidad Nacional de Jujuy. Instituto de Geología Minera; Argentina Fil: Chemisquy, Maria Amelia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Universidad Nacional de La Rioja. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Universidad Nacional de Catamarca. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Secretaría de Industria y Minería. Servicio Geológico Minero Argentino. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Provincia de La Rioja. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja; Argentina Fil: Turazzini, Guillermo Fidel. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Ciencias Geológicas; Argentina Fil: Prevosti, Francisco Juan. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Universidad Nacional de La Rioja. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Universidad Nacional de Catamarca. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de la Rioja. - Secretaria de Industria y Minería. Servicio Geológico Minero Argentino. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Provincia de La Rioja. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja; Argentina

We describe a near-complete, three-dimensionally preserved skeleton of a metatherian (relative of modern marsupials) from the middle Eocene (Lutetian: 44–43 million years ago) Lülük member of the Uzunçarşıdere Formation, central Turkey. With an estimated body mass of 3–4 kg, about the size of a domestic cat (Felis catus) or spotted quoll (Dasyurus maculatus), it is an order of magnitude larger than the largest fossil metatherians previously known from the Cenozoic of the northern hemisphere. This new taxon is characterised by large, broad third premolars that probably represent adaptations for hard object feeding (durophagy), and its craniodental morphology suggests the capacity to generate high bite forces. Qualitative and quantitative functional analyses of its postcranial skeleton indicate that it was probably scansorial and relatively agile, perhaps broadly similar in locomotor mode to the spotted quoll, but with a greater capacity for climbing and grasping. Bayesian phylogenetic analysis of a total evidence dataset comprising 259 morphological characters and 9kb of DNA sequence data from five nuclear protein-coding genes, using both undated and “tip-and-node dating” approaches, place the new taxon outside the marsupial crown-clade, but within the clade Marsupialiformes. It demonstrates that at least one metatherian lineage evolved to occupy the small-medium, meso- or hypo-carnivore niche in the northern hemisphere during the early Cenozoic, at a time when there were numerous eutherians (placentals and their fossil relatives) filling similar niches. However, the known mammal fauna from Uzunçarşıdere Formation appears highly endemic, and geological evidence suggests that this region of Turkey was an island for at least part of the early Cenozoic, and so the new taxon may have evolved in isolation from potential eutherian competitors. Nevertheless, the new taxon reveals previously unsuspected ecomorphological disparity among northern hemisphere metatherians during the first half of the Cenozoic.

Zeusdelphys complicatus is one of the most enigmatic metatherians from the Itaboraí Basin. The type and only known specimen was previously regarded as the upper dentition of Eobrasilia; an M4 of a new taxon; an M3 of a Kollpaniidae (now regarded as a group of “condylarths”); a probable M1 of an incertae sedis taxon; and as an M1 of a Protodidelphidae. Herein, we present a morphological review of the dental structures of Zeusdelphys complicatus, presenting new interpretations and comparing it with other North and South American taxa. We also perform a phylogenetic analysis in order to test the affinities of Zeusdelphys and the validity of most studied characters. The results recovered Zeusdelphys complicatus as more closely related to Hatcheritherium alpha than to any other metatherian. Glasbiidae were recovered as the sister lineage of Protodidelphidae within Didelphimorphia, as true marsupials. Ectocentrocristus was recovered as the sister taxon of Zeusdelphys + Hatcheritherium, as a Hatcheriformes. The analysis recovered this suborder as an independent lineage from Polydolopimorphia, being more closely related to “Alphadontidae”. The affinities with Protodidelphidae are a result of convergent evolution, as Zeusdelphys is more closely related to Hatcheritherium alpha from the Late Cretaceous of North America. The results support a North American origin for Hatcheriformes. The presence of strong sea-level lowstands and islands in the Caribbean Plate during the Late Cretaceous provide valid data to support a faunal interchange between Americas during the latest Late Cretaceous. Based on the results, Zeusdelphys represents a South American early Eocene surviving Hatcheriformes.

Two centuries ago, Georges Cuvier, the father of vertebrate palaeontology, described an almost complete skeleton of a small ‘opossum’ from the Montmartre Gypsum (late Eocene): Peratherium cuvieri. His description of the visible bones remains famous, as it was an unequivocal demonstration of his principle of correlation of parts. With the state-of-the-art technology of X-ray microtomography (computed tomography scanning), it is now possible to extend the investigation of the Cuvier's ‘opossum’, one of the most iconic specimens of the Museum national d'Histoire naturelle (MNHN). Beneath the visible upper cheek teeth the skull is damaged and fragmented, but provides significant new data, with the most complete and informative structures being the paired petrosal bones. Comparisons with the didelphid Marmosa murina (Linnaeus, 1758) and three other fossil herpetotheriids [Amphiperatherium minutum Aymard, 1846, Herpetotherium cf. fugax Cope, 1873, and Peratherium elegans (Aymard, 1846)] suggest that dental features alone are not sufficient to discriminate between the different species, being based on highly variable, polymorphic features of the stylar shelf. The anatomy of the petrosal bone, which has yet to be described for P. elegans or A. minutum, reveals marked differences between P. cuvieri and H. cf. fugax, however, which brings into question the monophyly of Herpetotheriidae. Further investigation of petrosal characters for herpetotheriids would therefore shine new light on the phylogenetic affinities of Herpetotheriidae, in terms of both the relationships within the family and the position of Herpetotheriidae within Metatheria as a whole.

This work is based on new dental, cranial, and postcranial remains of the argyrolagidMicrotragulus bolivianus (Metatheria, Polydolopimorphia, Bonapartheriiformes)and dental pieces of Microtragulus sp. coming from late Pliocene-early Pleistocene levelsof the Uquía Formation exposed at San Roque, Humahuaca (Jujuy Province,Argentina). They were found in association with amphibians, lizards, birds, rodents,and didelphid marsupials, forming an assemblage probably generated by the trophicactivity of owls. Specimens were assigned to M. bolivianus based on the followingcombination of features: M3 subcircular, with a flexus between the mesiolabial lobeand the metacone, paracone and metacone not differentiated in M4, absence of entoflexidin m1-2 and shallow entoflexid in m3, proportionally large talonid in m4, with a distinguishabledistal flexid. Microtragulus bolivianus, initially recorded in Pliocenesediments exposed at the Bolivian Altiplano, was represented only by a mandibularfragment with m3-m4. Based on a much more complete sample we present a detaileddental and postcranial description of the species. Since 1904 the family name Argyrolagidaeand the generic names Microtragulus and Argyrolagus have been subject ofseveral nomenclatural changes. A revision of these modifications, mainly those occurringin the last 40 years, is also presented. Furthermore, we analyze some mandibulartraits of argyrolagids such as the maxillary canal (retrodental canal), a very odd featurepresent in all the members of the family, which could be related to the passage of aconnection between the inferior alveolar and inferior orbital veins, as in some extantmammals. Este trabajo se basa en nuevos restos dentales, craneales y postcraneales del argirolágido Microtragulus bolivianus (Metatheria, Polydolopimorphia, Bonapartheriiformes) y piezas dentales de Microtragulus sp. provenientes de niveles de edad pliocena tardía-pleistocena temprana de la Formación Uquía, aflorantes en San Roque, Humahuaca (provincia de Jujuy, Argentina). Fueron encontados en asociación con anfibios, lagartijas, aves, roedores y marsupiales didélfidos, constituyendo una asociación probablemente generada por la actividad trófica de lechuzas. Los especímenes fueron asignados a M. bolivianus en base a la siguiente combinación de características: M3 subcircular, con un flexo entre el lóbulo mesiolabial y el metacono, paracono y metacono no diferenciados en M4, ausencia de entofléxido en m1-2 y un entofléxido somero en m3, un talónido proporcionalmente grande en m4, con un fléxido distal distinguible. Microtragulus bolivianus, inicialmente registrado en sedimentos pliocenos del altiplano boliviano, se hallaba representado únicamente por un fragmento mandibular con un incisivo y m3-m4. En base a una muestra mucho más completa presentamos una descripción dental y postcraneal detallada de la especie. Desde 1904 el nombre de la familia Argyrolagidae y los nombres genéricos Microtragulus y Argyrolagus han sido sujetos a varios cambios nomenclaturales. También se presenta una revisión de estas modificaciones, principalmente aquellas de los últimos 40 años. A su vez, analizamos algunas características de los argirolágidos tales como el canal maxilar (canal retrodental), un atributo muy particular presente en todos los miembros de la familia, el cual podría estar relacionado con el pasaje de una conexión entre las venas alveolar inferior y orbital inferior, como en algunos mamíferos actuales. Ce travail est basé sur des nouveaux restes dentaires, crâniens, et postcrâniens de l'argyrolagidé Microtragulus bolivianus (Metatheria, Polydolopimorphia, Bonapartheriiformes) et des restes dentaires de Microtragulus sp. provenant des niveaux du Pliocène récent-Pléistocène ancien de la formation d'Uquía qui affleurent à San Roque, Humahuaca (province de Jujuy, Argentine). Ils ont été trouvés en association avec des amphibiens, des lézards, des oiseaux, des rongeurs, et des marsupiaux didelphidés, formant un assemblage probablement accumulé par l'activité trophique de chouettes. Les spécimens ont été attribués à M. bolivianus d'après la combinaison de caractères suivante : une M3 subcirculaire avec un flexus entre le lobe mésiolabial et le métacône, le paracône et le métacône non différenciés sur la M4, une absence d'entoflexide sur la m1 et la m2, et un entoflexide peu profond sur la m3, un talonide relativement développé sur la m4 avec un flexide distal perceptible. Microtragulus bolivianus, espèce à l'origine connue dans les sédiments pliocènes affleurant dans l'Altiplano bolivien, n'était jusqu'à présent représenté que par un seul fragment de mandibule avec m3-m4. D'après un échantillon bien plus complet, nous présentons une description détaillée des restes dentaires et postcrâniens de cette espèce. Depuis 1904, le nom de famille Argyrolagidae et les noms de genre Microtragulus et Argyrolagus ont été le sujet de nombreux changements de nomenclature. Une révision de ces modifications, principalement celles des 40 dernières années, est également présentée. De plus, nous analysons certains traits mandibulaires des argyrolagidés, tels que le canal maxillaire (canal rétrodentaire), un caractère très curieux qui est présent chez tous les membres de cette famille et qui pourrait être lié au passage d'une connexion entre la veine ophtalmique inférieure et la veine alvéolaire inférieure, comme chez certains mammifères actuels. Fil: Babot, María Judith. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico - Tucumán. Unidad Ejecutora Lillo; Argentina Fil: Garcia Lopez, Daniel Alfredo. Universidad Nacional de Tucumán. Facultad de Ciencias Naturales e Instituto Miguel Lillo; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Tucumán. Instituto Superior de Correlación Geológica. Universidad Nacional de Tucumán. Facultad de Ciencias Naturales e Instituto Miguel Lillo. Departamento de Geología. Cátedra Geología Estructural. Instituto Superior de Correlación Geológica; Argentina

Didelphid marsupials — opossums — are solitary creatures, a trait often considered primitive for marsupials. This view could change with the discovery of a mass-death fossil group of Pucadelphys andinus, a primitive mouse-sized relative of marsupials from the early Palaeocene (around 64 million years ago) of Bolivia. The remains of 35 individuals are jumbled together in a small area, apparently buried in a single event. This find is so remarkable that we now have more specimens of this ancient mammal than we do of some marsupials living today. The vast majority of Mesozoic and early Cenozoic metatherian mammals (extinct relatives of modern marsupials) are known only from partial jaws or isolated teeth, which give insight into their probable diets and phylogenetic relationships but little else. The few skulls known are generally crushed, incomplete or both1,2,3,4, and associated postcranial material is extremely rare. Here we report the discovery of an exceptionally large number of almost undistorted, nearly complete skulls and skeletons of a stem-metatherian, Pucadelphys andinus, in the early Palaeocene epoch5 of Tiupampa in Bolivia6,7,8. These give an unprecedented glimpse into early metatherian morphology, evolutionary relationships and, especially, ecology. The remains of 35 individuals have been collected, with 22 of these represented by nearly complete skulls and associated postcrania. These individuals were probably buried in a single catastrophic event, and so almost certainly belong to the same population9. The preservation of multiple adult, sub-adult and juvenile individuals in close proximity (

European terrestrial vertebrate sites of the Upper Paleocene-Lower Eocene deposits are predominantly known from the central and eastern parts of the Paris Basin. However, several outcrops covering this interval are scattered along the Upper Normandy coast, in the western part of the Paris Basin. Here we report the discovery of a new terrestrial vertebrate site in the Mortemer Formation, at the top of the cliffs of Sotteville-sur-Mer in Upper Normandy, France. The vertebrate level is situated about 1.5. m above the onset of the Paleocene-Eocene Carbon Isotope Excursion (CIE) based on dispersed organic carbon and is therefore Earliest Eocene in age. The vertebrate fauna is composed of fish, amphibians, lizards and mammals, including the earliest peradectid marsupials and paromomyid plesiadapiform of Europe. A diverse and rich charophyte flora is well represented throughout the lower part of the outcrop and allows the conclusion that the CIE falls in the Peckichara disermas biozone. La plupart des sites à vertébrés terrestres du Paléocène supérieur-Eocène inférieur d'Europe sont connus du Centre et de l'Est du Bassin de Paris. Cependant, plusieurs affleurements de cet intervalle de temps sont éparpillés le long de la côte en Haute-Normandie dans l'Ouest du Bassin de Paris. Dans cet article, nous faisons état de la découverte d'un nouveau site à vertébrés terrestres dans la Formation de Mortemer, au sommet des falaises de Sotteville-sur-Mer en Haute-Normandie, France. Le niveau à vertébrés est situé environ 1,5. m au-dessus du point initial de l'Excursion Isotopique du Carbone de la limite Paléocène-Eocène (CIE), mise en évidence à partir du carbone organique dispersé. Il est donc daté de l'Eocène basal. La faune de vertébrés comprend des poissons, des amphibiens, des lézards et des mammifères dont les plus anciens marsupiaux peradectidés et le plus ancien plésiadapiforme paromomyidé d'Europe. Une flore riche et variée de charophytes, bien distribuée le long de la partie inférieure de l'affleurement permet de préciser que la CIE est située dans la biozone à Peckichara disermas.

New metatherian petrosal bones from the mid to Late Palaeocene of Itaborai, belonging to three morphotypes (VI, VII, and VII), are formally described and compared to fossil and extant taxa known by their auditory region. An attempt at assigning petrosal types to tooth-based taxa from Itaborai was made by combining parsimony and morphometric methods. The first large scale phylogenetic analysis of the Itaboraian metatherians, involving basicranial and dental characters in a larger number of taxa, is provided here and is at the basis of a systematic revision of the metatherians from Itaborai. The combination of morphometric and cladistic analyses helps in understanding the affinities between the petrosals and the tooth-based taxa. The metatherians from Itaborai were taxonomically diverse, belonging to each of the most important radiations in marsupial evolutionary history (Didelphimorphia, Paucituberculata, Eometatheria). The inclusion of Palaeocene taxa in the crown group Marsupialia and above all in the Eometatheria radiation points to an early emergence of these clades in South America and corroborates the main molecular hypotheses. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159, 746–784.

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New petrosal bones, assigned to Pucadelphys and Andinodelphys, from the Lower Paleocene of Tiupampa, Bolivia, are described. These remains provide new information on the anatomy of the ear region of these taxa. The re-examination of characters from the petrosal and basicranium shed light on the phylogenetic relationships of the three Tiupampan genera known from complete cranial remains (i.e. Mayulestes, Pucadelphys and Andinodelphys). The combination of dental, general cranial and basicranial characters led to two alternative hypotheses. The first is that borhyaenoids (including Mayulestes) are nested within Notometatheria. Pucadelphyds (i.e. Pucadelphys and Andinodelphys) are the sister group of a clade comprising MHNC 8369 (one isolated petrosal from Tiupampa) and Marsupialia. The second favours the paraphyly of ‘borhyaenoids’ (i.e. the exclusion of Mayulestes from borhyaenoids) and the polyphyly of ‘Notometatheria’. In this case, Mayulestes and borhyaenids represent the stem group of a clade including Asiatic, American and Australian metatherians. This analysis of combined datasets (dental, general cranial and basicranial) highlighted contradictory information in the dental and cranial characters, serving to emphasize that in a large anatomical complex like an entire skull mosaic evolution of the characters is likely.

With less than 3 g of estimated body mass, the early Eocene Minusculodelphis minimus Paula Couto (Mammalia, Metatheria, Jaskhadelphyidae) is one of the smallest mammals, living or extinct. It has alternatively been regarded as a didelphid or a derorhynchid "ameridelphian," or even as an eometatherian marsupial. Here, we describe a new species of Minusculodelphis coming from the same locality (Itaborai Quarry, Brazil) and age (Itaboraian age) of the type species of the genus. It differs from M. minimus in its larger size and several dental characters. The new species offers data on the upper dentition and femur, which are unknown in the type species. Compared to other Paleogene metatherians, Minusculodelphis shows closer relationships with Jaskhadelphys, from the early Paleocene of Tiupampa, Bolivia, as well as with Kiruwamaq, from the late Eocene-early Oligocene of Peru. A cladistic analysis places all three genera within the family Jaskhadelphyidae (Metatheria, Order indet.), which includes small to tiny, insectivorous-like metatherians. We argue that insectivory (soft insects) is the best-supported diet for both species of Minusculodelphis, and that the most probable microhabitat for them was the understorey or leaf litter of tropical, rain forested environments.

I describe Archaeonothos henkgodthelpi gen. et. sp. nov., a small (estimated body mass ~40-80g) tribosphenic metatherian from the early Eocene Tingamarra Fauna of southeastern Queensland, Australia. This taxon, known only from a single isolated upper molar (M2 or M3) is characterised by a very distinctive combination of dental features that, collectively, probably represent faunivorous adaptations. These include: a straight, elevated centrocrista; a metacone considerably taller than the paracone; a wide stylar shelf (~50% of the total labiolingual width of the tooth); reduced stylar cusps; a long postmetacrista; a small and anteroposteriorly narrow protocone; an unbasined trigon; and the absence of conules. Some of these features are seen in dasyuromorphians, but detailed comparisons reveal key differences between A. henkgodthelpi and all known members of this clade. A. henkgodthelpi also predates recent molecular estimates for the divergence of crown-group Dasyuromorphia. Similar dental features are seen in a number of other metatherians, including the South American sparassodonts, Wirunodon chanku from the ?Middle-Late Eocene Santa Rosa local fauna of Peru, and Kasserinotherium tunisiense from the Early Eocene Chambi fauna of Tunisia, although whether A. henkgodthelpi is closely related to any of these taxa is unclear based on available evidence. I therefore refer A. henkgodthelpi to Metatheria incertae sedis. Potential relatives of A. henkgodthelpi are unknown from any other Australian fossil deposit.

Although sexual size dimorphism is an important component of the biology of many living New World metatherians, little is known about sexual dimorphism in extinct members of this group. Here, we describe morphometric variation in two traditionally recognized species of the paucituberculatan Acdestis from the early Miocene Santa Cruz Formation of Patagonia, A. owenii and A. spegazzinii (=A. lemairei). The results of this morphometric study show that specimens of Acdestis from the Santa Cruz Formation form a single morphological continuum, with no features that clearly distinguish the two recognized morphotypes besides overall body size and the depth of the mandibular ramus, both features which are sexually variable in extant marsupials. This suggests that the two species of Acdestis from the Santa Cruz Formation are better regarded as gender morphs of a single dimorphic species rather than two highly similar monomorphic species. The degree of sexual dimorphism inferred here for Acdestis is compar...

The long-term changes of the mammalian diversity of the Quercy and Limagne area (Massif Central, France) is quantitatively investigated through a ∼16-Myr time span running from the late Middle Eocene (∼38 Myr) to the Early Miocene (∼22 Myr). Within the specific context of ecosystem response to extreme climatic events, the goal of such a deep-time approach to local community and regional metacommunity evolution is to give time-scaled reference series with which extant changes can be objectively compared. The results indicate that local and regional ecological integration levels show very distinct evolutionary dynamics, involving at least partially independent sets of controlling parameters; those driving the metacommunity still remain largely to be better understood. Clearly, the analyzed data set constitutes a rather unique deep-time key into several major current macroecological debates.

A new species of Sparassodonta (Mammalia, Metatheria), Lycopsis padillai, sp. Nov., is described on the basis of a partial left maxilla with M1-M4 and fragments of lacrimal and jugal. The material comes from the early to early middle Miocene Castilletes Formation, La Guajira Peninsula, Colombia. This specimen represents the northernmost record of a fossil metatherian in South America and integrates a highly diverse vertebrate association, recently discovered in the north of Colombia. The La Guajira specimen is referred to the genus Lycopsis, as supported by the results of our phylogenetic analysis. This analysis also demonstrates that species of Lycopsis (L. torresi, L. longirostrus, L. viverensis, and L. padillai) constitute a monophyletic group and are placed as the basal taxon of Borhyaenoidea. Lycopsis padillai is a large-sized sparassodont with a body mass of about 22 kg. The presence of Lycopsis from La Guajira extends the geographical distribution of the genus to the entire South America, representing the sparassodont with the widest latitudinal distribution. Fil: Suarez Gomez, Sandra Catalina. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Paleontología Vertebrados; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Forasiepi, Analia Marta. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mendoza. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales. Provincia de Mendoza. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales. Universidad Nacional de Cuyo. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales; Argentina Fil: Goin, Francisco Javier. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Paleontología Vertebrados; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Jaramillo, Carlos. Smithsonian Tropical Research Institute; Panamá

Four Paleocene petrosal bones from the Itaborai fissure fillings of Brazil belong to two metatherian morphotypes. They are compared with ear regions of other fossil and recent marsupials. The results of a parsimony analysis using thirty-eight characters from the petrosal and ear region are partly congruent with recent studies and confirm that characters from the ear region are highly informative for phylogenetic reconstruction. Metatheria are diagnosed by five petrosal synapomorphies: stapedial artery absent in adults; reduced, intramural prootic canal; extrabullar internal carotid artery; inferior petrosal sinus between petrosal, basisphenoid, and basioccipital; and cava supracochleare and epiptericum completely separated. The stem group of South American and Australian metatherians is composed of the Paleocene taxa Pucadelphys, one of the oldest known metatherians from South America, and Petrosal Type II from Itaborai. Among the South American-Australian metatherians two clades are defined here...

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

The sequence of differentiation of major elements of the skeletal, muscular and nervous systems of the head is examined in developmental series of five eutherian (placental) and four metatherian (marsupial) mammals. The analysis identifies the elements that are conserved across the Theria, those that are unique to the Metatheria and to the Eutheria, and those that are variable. It is shown that although there are slight shifts in the sequence of development within the somatic tissues of the head, the primary difference between marsupial and placental mammals involves the timing and rate of differentiation of structures of the central nervous system (CNS) relative to a specific subset of structures of the cranial skeleton and musculature. In eutherians, CNS morphogenesis is well underway before the somatic tissues of the head begin differentiation. In metatherians, CNS development is delayed considerably and certain elements of the skeletal and muscular systems are advanced. It is concluded that the developmental differences between marsupial and placental mammals are best explained by the interaction of several processes including neurogenesis as a potential rate-limiting step, the developmental requirements of somatic elements, and the extremely short period of organogenesis of marsupial mammals. Several other issues, including the way that these data may be applied to determine the primitive therian developmental condition, and the use of comparative developmental data to address basic questions on morphogenetic processes, are discussed.