Craniodental anatomy of late Oligocene archaeohyracids (Notoungulata,Mammalia) from Bolivia and Argentina and new phylogenetic hypotheses

The rich Deseadan fauna from the locality of Salla Luribay (Bolivia) documents the last occurrence of archaeohyracids, a poorly known group of small typotherian notoungulates. Exceptionally, archaeohyracid remains are extremely abundant in the Salla deposits and are assigned to a single new species Archaeohyrax suniensis sp. nov . The anatomy of the new taxon is presented and the ontogeny of the cheek teeth is described. Archaeohyrax patagonicus Ameghino, 1897 from the Deseadan of Patagonia is also redescribed and compared with the new Bolivian species. Additionally, juvenile teeth of Sallatherium altiplanense (Hegetotheriidae) are described because they provide crucial phylogenetic information for understanding the phylogeny of archaeohyracids. A cladistic analysis performed on archaeohyracids and hegetotheriids supports for the first time the existence of a clade of late archaeohyracids (post‐Mustersan), which is the sister taxon of all hegetotheriids. It also argues for the origin of mesotheriids within archaeohyracids and for the existence of a hegetotheriine clade. These conclusions fit well with temporal data known for each taxa. Nevertheless, the present analysis also underlines the fact that the lack of data concerning the cranial anatomy of many archaeohyracids (Eohyrax, Pseudhyrax, Archaeotypotherium, Protarchaeohyrax) weakens the phylogenetic signal. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 458–509.     

New rodents (Mammalia) from the late Oligocene of Cabeza Blanca (Chubut) and the first rodent radiation in Patagonia

Caviomorph rodents, the New World Hystricognathi, are one of the most characteristic groups of South American mammals. Although they have been in the continent at least since the middle Eocene, those of the Deseadan Land mammal Age (early–late Oligocene) are the best source to understanding their early history due of their good record, large geographic distribution and good temporal calibration of many of the local faunas. Here, we describe the following new taxa from the classical locality of Cabeza Blanca: Octodontoidea Acaremyidae Galileomys baiosn. sp., Octodontoidea incertae sedis Ethelomys loomisin. gen.,n. comb., Acarechimys leucotheaen. sp., Protacaremys? adilosn. sp., Chinchilloidea incertae sedis Loncolicu tretosn. gen., n. sp., Incamys menniorumn. sp., Caviomorpha incertae sedis Llitun notucan. gen., n. sp., Leucokephalos zeffiaen. gen., n. sp. and Cephalomyidae Cephalomys ceciaen. sp. The DP4 of an ‘eocardiid’, Asteromys punctus? is described for the first time. These new taxa allow us to reinterpret the relationships of some of the previously known Deseadan species and genera. They show a great early diversification at least for extra Andean Patagonia, involving at least three of the main caviomorph lineages: octodontoids, chinchilloids and cavioids.     

Skull anatomy of the Oligocene toothed mysticete Aetioceus weltoni (Mammalia; Cetacea): implications for mysticete evolution and functional anatomy

Toothed mysticetes of the family Aetiocetidae from Oligocene rocks of the North Pacific play a key role in interpretations of cetacean evolution because they are transitional in grade between dorudontine archaeocetes and edentulous mysticetes. The holotype skull of Aetiocetus weltoni from the late Oligocene (28–24 Ma) of Oregon, USA, has been further prepared, revealing additional morphological features of the basicranium, rostrum and dentary that have important implications for mysticete evolution and functional anatomy. The palate of Aetiocetus weltoni preserves diminutive lateral palatal foramina and associated delicate sulci which appear to be homologous with the prominent palatal foramina and sulci that occur along the lateral portion of the palate in extant mysticetes. In modern baleen whales these foramina allow passage of branches of the superior alveolar artery, which supplies blood to the epithelia of the developing baleen racks. As homologous structures, the lateral palatal foramina of A. weltoni suggest that baleen was present in this Oligocene toothed mysticete. Cladistic analysis of 46 cranial and dental characters supports monophyly of the Aetiocetidae, with toothed mysticetes Janjucetus and Mammalodon positioned as successive sister taxa. Morawanacetus is the earliest diverging aetiocetid with Chonecetus as sister taxon to Aetiocetus species. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 308–352.     

Killer sperm whale: a new basal physeteroid (Mammalia, Cetacea) from the Late Miocene of Italy

Zygophyseter varolai , a new genus and species of Physeteroidea (Cetacea, Odontoceti), is based on an almost complete skeleton from the Late Miocene (Tortonian) in southern Italy. The extreme elongation of the zygomatic process of the squamosal and the circular supracranial basin (probably for housing the spermaceti organ) delimited by a peculiar anterior projection of the supraorbital process of the right maxilla are the most distinctive features of this bizarre sperm whale. Large body size, large teeth present in both lower and upper jaw, and anteroposteriorly elongated temporal fossa and zygomatic process of the squamosal indicate that this cetacean (for which we suggest the English common name killer sperm whale) was an active predator adapted to feeding on large prey, similarly to the extant killer whale ( Orcinus orca ). A phylogenetic analysis reveals that Zygophyseter belongs to a Middle–Late Miocene clade of basal physeteroids, together with Naganocetus (new genus for the type of ' Scaldicetus ' shigensis ). Moreover, the phylogenetic analysis shows evidence of a wide physeteroid radiation during the Miocene and that the extant Physeter and Kogia belong to two distinct families that form a clade representing the crown-group Physeteroidea.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 103–131.     

New data on the Cainotheriidae (Mammalia, Artiodactyla) from the early Oligocene of south-western France

The Cainotheriidae are small artiodactyls that suddenly appeared in the late Eocene of western Europe. A revision of early Oligocene cainotheriid lineages is proposed on the basis of newly dated material from the Quercy Phosphorites (south-western France). A significant diversification of the group occurred at the end of the Eocene. Few species seem to have persisted through the Eocene/Oligocene boundary, but the Cainotheriidae subsequently diversified rapidly during the early Oligocene.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 145−166.     

New ceratomorph perissodactyls (Mammalia) from the Middle and Late Eocene of Mongolia: their implications for phylogeny and dating

Eight species rank taxa arc described from Eocene sites in the Gobi Desert, Mongolia. Irdinolophus gen. now is erected for Desmalotherium mongoliense Osborn and for Irdinolophus? tuiensis sp. nov. The genus is placed at the stem of the family Depcrctellidae. Intraspecific variation in lekolophus magnus is documented for the first time; Pachylophus is synonymized with leleoloplms; and Deperetella cf.birmanica is recorded far north of its previously known range in Burma. The first known lower teeth of Colodon inceptus support referral ofDesmalotherium Jissum to Colodon. The stratigraphic range of the hyracodontid Ardynia is extended back from the late Eocene to the early middle Eocene. A range of primitive tapiroids and rhinocerotoids arc analysed cladistically to resolve the problematic affinities of the extinct families Dc-peretellidae and Rhodopagidae. The Depcrctellidae are shown to belong to a monophyletic superfamily Tapiroidca, whilst the Rhodopagidae belongs to the Rhinocerotoidea, but is distinct from the Hyracodontidae. Veragromovia, previously synonymized with the tapiroid Helaleles, is resurrected as a valid genus within the Rhodopagidae. Tsagan Khutcl and other sites in the Kholbolchi formation may be older than Irdinmanhan faunas in Asia, equaling with those from the Arshanto Formation of China of probable early Lutetian or latest Ypresian age. The Irdinmanhan Mergen fauna may be slightly older than that from Irdin Manha, because of the primitive aspect of its Lophialetes.     

A giant rhinocerotoid (Mammalia, Perissodactyla) from the Late Oligocene of north-central Anatolia (Turkey)

A giant rhinocerotoid is described for the first time south of the Black Sea, in Turkey. The single specimen, a fragmentary radius referred to Paraceratherium sp., originates from conglomerates nearby at Gözükizilli, in the Çankiri–Çorum Tertiary basin. These layers correspond to the Lower member of the Kizilirmak Formation. The same locality (Gözükizilli-2) yields also the small rhinocerotid Protaceratherium sp., cf. P. albigense (Roman, 1912). Three other mammal localities (Gözükizilli-1, in the Lower Member of the formation, with several rodent species; Tepe 641 and Kizilirmak, in the Upper Member, with a diversified micro- and macro-mammal fauna) allow us to refer the Kizilirmak Formation as a whole to the Late Oligocene. All the observed taxa have strong Asian and/or European affinities, which precludes any geographical insulation for this part of Anatolia during the Late Oligocene.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152 , 581–592.     

A new Oligocene Ctenodactylinae (Rodentia: Mammalia) from Ulantatal (nei Mongol): new insight on the phylogenetic origins of the modern Ctenodactylidae

The four living genera of Ctenodactylidae (Rodentia) are the only survivors of a flourishing Tertiary group. In this paper, we describe a new Oligocene species from Ulantatal (Chinese Mongolia) that highlights the origins of crown ctenodactylids. This species, Helanshania deserta gen. et sp. nov. , is lophodont and displays semi‐hypsodont teeth, a dental pattern that is somewhat transitional between that of primitive Oligocene ctenodactylids and the later hypsodont genera. We perform here a cladistic assessment of the dental evidence for species produced by the successive radiations of the group. In order to get new data, to decipher homologies for the dental pattern of modern ctenodactylids, and to specify their dental replacement, we describe additional dental material of Ctenodactylus, Massoutiera, and Felovia. The phylogenetic analysis (using PAUP) considered 45 characters (mainly dental) and 31 species. The performed heuristic searches yielded 596 equally most parsimonious trees. Protataromys and Karakoromys are stem ctenodactylids and appear as the earliest offshoots of the Ctenodactylidae clade, which represents a well‐supported family rank. Within this family, the Tataromyinae appear paraphyletic, whereas the Ctenodactylinae sensu lato are a clade including the new taxon Helanshania. As such, a revision of the Tataromyinae is envisaged and a new subfamily is erected (Yindirtemyinae). Amongst the Ctenodactylinae, a tribe Ctenodactylini encompassing the crown ctenodactylines is proposed. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 531–550.     

Re-assessment of the Oligocene genera Prosotherium and Propachyrucos (Hegetotheriidae,Notoungulata)

Several systematic revisions of the Hegetotheriidae have been published in the last twenty years, but some important taxonomic work remains limited to unpublished studies. In one such thesis, the late Oligocene genera Prosotherium and Propachyrucos were revised, so that Prosotherium garzoni and Propachyrucos smithwoodwardi were considered the only valid species for each genus. Later authors loosely accepted these informal revisions, but recent phylogenetic coding has yet to accurately reflect these revisions, with character states often at odds with the known material for these genera. This study re-examines most of the fossil material assigned to Propachyrucos and Prosotherium, including both published and unpublished fossils, and contributes to a better understanding of the systematics, taxonomic diversity and early morphological evolution of the pachyrukhines. We conclude that Prosotherium garzoni and Propachyrucos smithwoodwardi are indeed the only valid species for each genus, confirming the influential but unpublished revisions that have become widely accepted. The known material for these species is clarified to aid future phylogenetic studies. At the same time, Propachyrucos aequilatus and Pr. robustus are herein considered nomina dubia. Additionally, Miocene species originally ascribed to Propachyrucos, along with unpublished material loosely assigned to the same genus, are here synonymized with other genera. Finally, several fossils previously recognized as Prosotherium sp. are here recognized as P. garzoni.     

Protein variation and its systematic implications for the South African Leporidae (Mammalia:Lagomorpha)

Analysis of 9 genetic loci by horizontal starch gel electrophoresis was performed on 203 specimens of South African representatives of the leporid genera Lepus and Pronolagus . Three loci were found to be polymorphic within samples of the various species of both genera: transferrin (Tf-I), esterase-1 (Es-I) and esterase-2 (Es-II). Only one system, carbonic anhydrase (CA-I), proved useful in species identification, this being restricted to the genus Lepus . Based on this system all Lepus specimens analysed would be assigned to one of two groups corresponding to the species L. capensis and L. saxatils .     

Juvenile cranial anatomy of Nimravidae (Mammalia, Carnivora): biological and phylogenetic implications

The anatomy of the spectacular sabretoothed carnivorans has been studied many times. Their behavioural and biomechanical implications are well known in adults, but not in juvenile individuals, which are poorly represented in the fossil record. This work reports on a newly discovered Hoplophoneini (Nimravidae Nimravinae) of 5–6 months of age, which represents the earliest and one of the most complete juvenile crania discovered. We discuss and compare the juvenile anatomy of Palaeogene nimravids relative to that of other mammals, especially other carnivorans including the Miocene nimravids (i.e. Barbourofelinae). We also discuss the ontogenetic development of this individual and other sabretoothed predators, which provides important data for reconstructing the biology of these extinct predators. Our study of the juvenile anatomy of the Hoplophoneini reveals an original association of basicranial features: the ectotympanic rotates away from the cranium earlier than in other mammals, which may be an additional autapomorphy distinguishing the Palaeogene Nimravidae from other Carnivora; within Palaeogene Nimravidae, at a similar ontogenetic stage, there is a major difference in development and maximum ventral extension of ossification of the caudal entotympanic; in contrast to the Miocene nimravid Barbourofelis, there is no strong relationship between the eruption of the upper canines and the development of other sabretoothed features (especially the mandibular flange) in the individual described. These results confirm the conclusions of recent studies which suggested that Palaeogene Nimravidae (Nimravinae) and Miocene Nimravidae (Barbourofelinae) are not closely related. Muscular reconstruction and biomechanical models also confirmed that juvenile individuals were probably not able to kill or hunt any prey. They may have fed on carcasses, eating muscles by using the incisors, and associated with a pulling or twisting action of neck. © 2003 The Linnean Society of London. Zoological Journal of the Linnean Society, 2003, 138 , 477–493.     

An X-radiographic and SEM study of the osseous inner ear of multituberculates and monotremes (Mammalia): implications for mammalian phylogeny and evolution of hearing

Multituberculate petrosals with well-preserved, three-dimensional internal anatomy from the Late Cretaceous/early Paleocene Bug Creek Anthills, Montana, U.S.A., are described from X-radiographic and SEM images, as well as from conventional visual observations, and are compared with the anatomy of the osseous inner ear in monotremes and in primitive non-therian and therian mammals. Results of this study indicate that: (1) the cochlea of at least some multituberculates retained a lagena, previously known only in monotremes among mammals; (2) an enlarged vestibule evolved in several lineages of multituberculates independently, and hence is not a synapomorphy of the order; (3) the cochlear canal lacks osseous laminae in support of the short, wide basilar membrane, which was probably inefficient in responding to high-frequency airborne vibrations; and (4) consequently, bone-conducted hearing in some multituberculate species may have been important in interpretation of their surroundings. Comparisons with the inner ear of monotremes and primitive therians indicate that curvature of the cochlea and cribriform plates for passage of vestibulocochlear nerve branches through the petrosal are unlikely homologues between monotremes and therians. From non-therian to therian mammals, there is a distinct morphological gap in the inner ear transition, characterized by acquisition of a number of neomorphs in the therian inner ear; an intermediate stage has yet to be discovered.     

A new Hyperodapedon (Archosauromorpha,Rhynchosauria) from the Upper Triassic of India: implications for rhynchosaur phylogeny

A new species of the rhynchosaur genus Hyperodapedon, namely H. tikiensis, is described from well‐preserved skeletal elements that were collected from the Upper Triassic Tiki Formation of India. Hyperodapedon tikiensis is diagnosed on the basis of several cranial and postcranial features including longer than wide basipterygoid process, crest‐shaped maxillary cross section lateral to the main longitudinal groove, deeply excavated neural arches of mid‐dorsal vertebrae, long scapular blade, a pronounced deltopectoral crest, proximal humeral end much broader than distal end, iliac length greater than iliac height, equal pre‐ and postacetabular iliac lengths and circular femoral cross section. Two distinct morphotypes of the maxillary tooth plates can be discerned, which are attributed to ontogenetic variations. A maximum‐parsimony analysis was carried out to show that the order Rhynchosauria is characterized by nine cranial and one postcranial character states. The analysis reveals that Otischalkia elderae is invalid and the basal forms, Howesia and Mesosuchus, are closely related. The Mid‐Triassic genus Ammorhynchus is more derived and forms a sister group to the Late Triassic subfamily Hyperodapedontinae. Isalorhynchus and Teyumbaita are basal to the pandemic genus Hyperodapedon. Twenty‐four characters that are not homoplasious document major patterns of skeletal evolution in rhynchosaurs. From laterally oriented scapula and slender propodials, the postcranial skeleton evolved into a more robust form as is evident from nearly vertical scapula and increase in the robustness of the propodials. Shortening of the femur is noted in the derived Late Triassic forms as exemplified in Hyperodapedon gordoni, Hyperodapedon huxleyi and H. tikiensis.     

Comparative Endocranial Anatomy,Encephalization, and Phylogeny of Notoungulata (Placentalia,Mammalia)

Journal of Mammalian Evolution - Cranial endocasts are one of the most direct tools available to obtain information about the endocranial cavity of fossil mammals, but few anatomical comparisons...     

The phylogeny and taxonomy of Hippopotamidae (Mammalia: Artiodactyla): a review based on morphology and cladistic analysis

The phylogeny and taxonomy of the whole family Hippopotamidae is in need of reconsideration, the present confusion obstructing palaeoecology and palaeobiogeography studies of these Neogene mammals. The revision of the Hippopotamidae initiated here deals with the last 8 Myr of African and Asian species. The first thorough cladistic analysis of the family is presented here. The outcome of this analysis, including 37 morphological characters coded for 15 extant and fossil taxa, as well as non-coded features of mandibular morphology, was used to reconstruct broad outlines of hippo phylogeny. Distinct lineages within the paraphyletic genus Hexaprotodon are recognized and characterized. In order to harmonize taxonomy and phylogeny, two new genera are created. The genus name Choeropsis is re-validated for the extant Liberian hippo. The nomen Hexaprotodon is restricted to the fossil lineage mostly known in Asia, but also including at least one African species. The genus Hippopotamus is confirmed. These changes represent substantial advances for understanding the evolutionary history of the Hippopotamidae, and provide a new framework for future studies.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 1–26.     

Earliest ontogeny of Middle Ordovician rhynchonelliform brachiopods (Clitambonitoidea and Polytoechioidea): implications for brachiopod phylogeny

New data on the earliest ontogeny of Mid-Ordovician Baltoscandian clitambonitoid ( Apomatella , Neumania and Oslogonites ) and polytoechioid ( Antigonambonites and Raunites ) brachiopods reveal significant differences in the life history of the taxa belonging to these two superfamilies. The Polytoechioidea and probably other members of the Billingsellida had planktotrophic larvae, in which the dorsal and ventral mantle lobes formed separately and without reversion. The 'pedicle sheath' in Antigonambonites is secreted by a section of modified ventral mantle and thus this 'pedicle' is not homologous within the pedicle of rhynchonellate brachiopods. It is likely that polytoechioids and other members of the strophomenate clade had the same type of ontogeny and mode of attachment. In contrast, the ontogeny and mode of attachment of clitambonitoids are similar to that of recent rhynchonellates: their mantle lobes and both valves formed simultaneously, and the pedicle most likely formed from the larval pedicle lobe. Evidence for the lecithotrophic nature of clitambonitoid larva is discussed. This confirms that the Clitambonitoidea, unlike the Polytoechioidea, represents an ingroup within the Rhynchonellata.     

A non‐aquatic otter (Mammalia,Carnivora, Mustelidae) from the Late Miocene (Vallesian,MN 10) of La Roma 2 (Alfambra,Teruel, Spain): systematics and functional anatomy

A new genus and species of otter‐like mustelid, Teruelictis riparius, is created on the basis of a partial skeleton from the Late Miocene (Vallesian age, MN 10) locality of La Roma 2 (Teruel, Spain), including several postcranial elements, the skull, and the mandible. The combination of a typically lutrine dentition, similar to that of other fossil otters such as Paralutra jaegeri, with a very slender postcranial skeleton, including a long back and gracile long bones and metacarpals, thus lacking any aquatic adaptations, was previously unknown in the fossil record. This mosaic of features strongly suggests the possibility that the aquatic lifestyle of otters could have appeared after the initial development of the distinctive dental morphology of this specialized group of mustelids. © 2013 The Linnean Society of London