Late Oligocene mesotheriids (Mammalia, Notoungulata) from Salla and Lacayani (Bolivia): implications for basal mesotheriid phylogeny and distribution

The oldest known mesotheriids are referred to the genus Trachytherus , a taxon traditionally included in its own subfamily 'Trachytheriinae'. The ontogeny, morphological variation and systematics of the genus Trachytherus are still poorly understood and the monophyly of this taxon is questionable. For the first time, a large collection of basal mesotheriids (belonging to the genus Trachytherus ) yielded by the Bolivian Salla beds (Deseadan, late Oligocene) and previously referred to T. spegazzinianus was included in a large-scale analysis. This has led to the recognition of a unique new species in the locality of Salla-Luribay, distinct from T. spegazzinianus : Trachytherus alloxus sp. nov . Skulls of this species are described and cheekteeth variation range throughout ontogeny is discussed. These new data are incorporated in a parsimony analysis of basal mesotheriids. The most probable hypothesis of a clade T. spegazzinianus -mesotheriines is discussed. The distinction of the new Salla species from the Patagonian species T. spegazzinianus confirms the faunal differences observed between these Bolivian and Patagonian Deseadan localities. Moreover, the assignment to Trachytherus spegazzinianus of mesotheriid remains from the Bolivian Lacayani deposits complicates the problem of these faunal differences. Further systematic investigations on the Lacayani fauna and on the age of the Patagonian localities are needed.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 153–200.     

A basal sphenodontian (Lepidosauria) from the Jurassic of Patagonia: new insights on the phylogeny and biogeography of Gondwanan rhynchocephalians

Herein we describe a new rhynchocephalian taxon from the Middle Jurassic of Patagonia, Argentina, representing the first Jurassic record of the group in South America. The new taxon, consisting of a complete dentary, is ascribed to Sphenodontia based on the presence of a deep and wide Meckelian groove, long posterior process, well‐developed coronoid process, and acrodont teeth showing dental regionalization including successional, alternate hatchling, and additional series. This allocation is reinforced by a phylogenetic analysis that places the new taxon in a basal position within a clade of sphenodontians that excludes Diphydontosaurus and Planocephalosaurus. Additionally, the new taxon clusters within a Gondwanan clade with the Indian Godavarisaurus from the Jurassic Kota Formation, sharing the presence of recurved and relatively large posterior successional teeth that are ribbed and bear a peculiar anterolingual groove. This sister‐group relationship is intriguing from a palaeobiogeographical viewpoint, as it suggests some degree of endemism during the initial stages of the breakup of Pangaea. We also discuss the ontogenetic stage of the new taxon and provide insights on the evolution of successional dentition in rhynchocephalians. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 342–360.     

Combined phylogenetic analysis of a new North American fossil species confirms widespread Eocene distribution for stem rollers (Aves,Coracii)

We report a nearly complete skeleton of a new species of stem roller (Aves, Coracii) from the early Eocene Green River Formation of North America. The new species is most closely related to two species‐depauperate lineages, Coraciidae (rollers) and Brachypteraciidae (ground rollers), that form a monophyletic crown clade (Coracioidea) with an exclusively Old World extant distribution. Phylogenetic analysis utilizing a matrix of 133 morphological characters and sequence data from three genes (RAG‐1, c‐myc, and ND2) identifies the new species as a stem member of the Coracii more closely related to the crown clade than the only previously identified New World taxon, Primobucco mcgrewi. The phylogenetic placement of the new species and Primobucco mcgrewiindicates a widespread northern hemisphere distribution in the Eocene with subsequent restriction to Africa, Madagascar, Australia, and temperate to tropical parts of Europe and Asia. It provides evidence of further ecological diversity in early stem Coracii and convergence on crown morphologies. The new species contributes to mounting evidence that extant distributions for major avian subclades may be of comparatively recent origin. Further late Palaeogene sampling is needed to elucidate potential drivers for shifting avian distributions and disappearance of Coracii from North America. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 586–611.     

Cranial osteology and phylogenetic position of the theropod dinosaur Proceratosaurus bradleyi (Woodward, 1910) from the Middle Jurassic of England

The cranial osteology of the small theropod dinosaur Proceratosaurus from the Bathonian of Minchinhampton, England, is described in detail, based on new preparation and computed tomography (CT) scan images of the type, and only known, specimen. Proceratosaurus is an unusual theropod with markedly enlarged external nares and a cranial crest starting at the premaxillary–nasal junction. The skull is highly pneumatic, with pneumatized nasals, jugals, and maxillae, as well as a highly pneumatic braincase, featuring basisphenoid, anterior tympanic, basipterygoid, and carotid recesses. The dentition is unusual, with small premaxillary teeth and much larger lateral teeth, with a pronounced size difference of the serrations between the mesial and distal carina. The first dentary tooth is somewhat procumbent and flexed anteriorly. Phylogenetic analysis places Proceratosaurus in the Tyrannosauroidea, in a monophyletic clade Proceratosauridae, together with the Oxfordian Chinese taxon Guanlong. The Bathonian age of Proceratosaurus extends the origin of all clades of basal coelurosaurs back into the Middle Jurassic, and provides evidence for an early, Laurasia‐wide, dispersal of the Tyrannosauroidea during the late Middle to Late Jurassic. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009.     

What is Geosaurus? Redescription of Geosaurus giganteus (Thalattosuchia: Metriorhynchidae) from the Upper Jurassic of Bayern,Germany

The holotype and referred specimens of Geosaurus giganteus, a metriorhynchid crocodile from the Tithonian (Upper Jurassic) of Germany, is redescribed, along with a historical overview of the genus and species. This taxon is unique among metriorhynchids as its serrated, strongly lateromedially compressed dentition is arranged as opposing blades, suggesting it was adapted to efficiently slice through fleshy prey. A new phylogenetic analysis of Crocodylomorpha is presented, which finds G. giganteus to be nested within what is currently considered Dakosaurus, whereas the other species currently assigned to Geosaurus form a clade with Enaliosuchus and the holotype of Cricosaurus. The phyletic relationship of G. giganteus with other metriorhynchids indicates that the current definition of the genus Geosaurus is polyphyletic, and that the inclusion of subsequent longirostrine species to this genus is in error. The re‐analysis presented herein demonstrates Geosaurus to be composed of three species sensu stricto. The appropriate taxonomic amendments to the Metriorhynchidae are also provided. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 551–585.     

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.     

Descriptions and phylogenetic relationships of a new genus and two new species of Oligo‐Miocene cormorants (Aves: Phalacrocoracidae) from Australia

Tertiary cormorant fossils (Aves: Phalacrocoracidae) from Late Oligocene deposits in Australia are described. They derive from the Late Oligocene – Early Miocene (26–24 Mya) Etadunna and Namba Formations in the Lake Eyre and Lake Frome Basins, South Australia, respectively. A new genus, Nambashag gen. nov. , with two new species ( Nambashag billerooensis sp. nov. , 30 specimens; Nambashag microglaucus sp. nov. , 14 specimens), has been established. Phylogenetic analyses based on 113 morphological and two integumentary characters indicated that Nambashag is the sister taxon to the Early Miocene Nectornis miocaenus of Europe and all extant phalacrocoracids. As Nambashag, Nectornis, and extant phalacrocoracids constitute a strongly supported clade sister to Anhinga species, the fossil taxa have been referred to Phalacrocoracidae. Sulids and Fregata were successive sister taxa to the Phalacrocoracoidea, i.e. phalacrocoracids + Anhinga. As phalacrocoracids lived in both Europe and Australia during the Late Oligocene and no older phalacrocoracid taxa are known, the biogeographical origin of cormorants remains unanswered. The phylogenetic relationships of extant taxa were not wholly resolved, but contrary to previous morphological analyses, considerable concordance was found with relationships recovered by recent molecular analyses. Microcarbo is sister to all other extant phalacrocoracids, and all Leucocarbo species form a well‐supported clade. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 277–314.     

The anatomy of the basal ornithischian dinosaur Eocursor parvus from the lower Elliot Formation (Late Triassic) of South Africa

Ornithischia is a morphologically and taxonomically diverse clade of dinosaurs that originated during the Late Triassic and were the dominant large‐bodied herbivores in many Cretaceous ecosystems. The early evolution of ornithischian dinosaurs is poorly understood, as a result in part of a paucity of fossil specimens, particularly during the Triassic. The most complete Triassic ornithischian dinosaur yet discovered is Eocursor parvus from the lower Elliot Formation (Late Triassic: Norian–Rhaetian) of Free State, South Africa, represented by a partial skull and relatively complete postcranial skeleton. Here, the anatomy of Eocursor is described in detail for the first time, and detailed comparisons are provided to other basal ornithischian taxa. Eocursor is a small‐bodied taxon (approximately 1 m in length) that possesses a plesiomorphic dentition consisting of unworn leaf‐shaped crowns, a proportionally large manus with similarities to heterodontosaurids, a pelvis that contains an intriguing mix of plesiomorphic and derived character states, and elongate distal hindlimbs suggesting well‐developed cursorial ability. The ontogenetic status of the holotype material is uncertain. Eocursor may represent the sister taxon to Genasauria, the clade that includes most of ornithischian diversity, although this phylogenetic position is partially dependent upon the uncertain phylogenetic position of the enigmatic and controversial clade Heterodontosauridae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 648–684.     

A new family of lithophoran Proseriata (Platyhelminthes), with the description of seven new species from the Indo‐Pacific and South America,and the proposal of three new genera

In this contribution a new representative of the taxon Meidiama Marcus, 1946, Meidiama uruguayensis sp. nov. , from Uruguay, is described, as are six more new species, for which three new genera are proposed: Dreuxiola philippi gen. nov. sp. nov. , from the French subantarctic archipelago Kerguelen; Yorknia aprostatica gen. nov. sp. nov. ; Serrula byronensis gen. nov. sp. nov. ; Serrula maxillaria sp. nov. ; Serrula concharum sp. nov. ; and Serrula acuta sp. nov. , from eastern Australia and Tasmania. Arguments are presented to propose a new taxon to contain these new species, rather than include them in the Archimonocelididae Meixner, 1938 (of which Meidiama has been considered a member so far), as well as to remove the Calviriinae Martens & Curini‐Galletti from the Archimonocelididae to become a separate taxon Calviriidae. Possible autapomorphies for the three families are discussed. It is also concluded that, with the present state of our knowledge, no sound indications can be given about close relationships. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 759–773.     

Neurocranial osteology and systematic relationships of Varanus (Megalania) prisca Owen, 1859 (Squamata: Varanidae)

The neurocranial osteology of the giant monitor lizard Varanus (Megalania) prisca Owen, 1859 is described in detail for the first time. Optimization of neurocranial characters onto phylogenetic topologies for varanoids, including Lanthanotus, Heloderma, and Varanus species nests V. prisca within an Indo‐Australian clade of Varanus on the basis of characters of the otic capsule. A sister‐taxon relationship between V. prisca and Varanus komodoensis Ouwens, 1912 is proposed based on apomorphies of the crista prootica, fenestra vestibuli, occipital recess, and supraoccipital. These results support a monophyletic clade of giant monitors among Indo‐Australian species, and unambiguously synonymize Megalania with Varanus at both generic and subgeneric levels. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 445–457.     

A new genus of red‐toothed shrew (Mammalia,Soricidae) from the Early Pleistocene of Gran Dolina (Atapuerca,Burgos, Spain), and a phylogenetic approach to the Eurasiatic Soricinae

This paper describes a new genus and species of shrew from the Pleistocene (levels 4, 5, and 6) of the Gran Dolina site, located in the Sierra de Atapuerca (Burgos, Spain). Dolinasorex glyphodon gen. et sp. nov. (Mammalia, Soricidae) is a type of large‐sized soricine, endemic in character, which inhabited the Sierra de Atapuerca in the last part of the Early Pleistocene (c. 780–900 kya). Morphometric and phylogenetic analyses indicate that the new species is more closely related to Asiatic forms than to the species Beremendia fissidens with its primarily European distribution. The main palaeoecological and palaeobiogeographical implications of the new taxon in the area around Atapuerca and in the Iberian Peninsula in general are discussed. The paper also presents a tentative phylogeny for the Soricinae subfamily in Eurasia, including for the first time exclusively fossil species, producing results that may prove to be of great interest for more complete studies in the future. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 904–925.     

A spectacular new species of Nepenthes L. (Nepenthaceae) pitcher plant from central Palawan,Philippines

A new species of Nepenthes L., N. attenboroughii (Nepenthaceae), from Palawan Island in the Philippines, is described and illustrated. It is restricted to rocky, ultramafic soils that comprise the summit region of Mount Victoria, Municipality of Narra, where it occurs in isolation from other members of the genus. On the basis of the morphological features, this new taxon appears to be related to both N. mira Jebb & Cheek of Palawan and N. rajah Hook.f. of Borneo. Its substantial size places it among the largest of known pitcher plants. The diagnostic morphological characters are discussed and an updated key is provided for a revised complex of Nepenthes species from the Palawan and North Borneo phytogeographical region. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 195–202.     

The skull and the palaeoecological significance of Labidosaurus hamatus, a captorhinid reptile from the Lower Permian of Texas

The cranial skeleton of the large captorhinid reptile Labidosaurus hamatus , known only from the Lower Permian of Texas, is described on the basis of new, undescribed specimens. Labidosaurus is distinguished from other captorhinids by the more extreme sloping of the ventral (alveolar) margin of the premaxilla, a low dorsum sellae of the parabasisphenoid, a reduced prootic, a narrow stapes, and a relatively small foramen intermandibularis medius. Despite the presence of a single row of teeth in each jaw, the skull of Labidosaurus resembles most closely those of moradisaurines, the large multiple-tooth-rowed captorhinids of the latest Early and Middle Permian. A phylogenetic analysis confirms that the single-tooth-rowed L. hamatus is related most closely to moradisaurines within Captorhinidae, a relationship that supports the hypothesis of a diphyletic origin for multiple rows of marginal teeth in captorhinids (in the genus Captorhinus and in the clade Moradisaurinae). In view of the close relationship between L. hamatus and moradisaurines, which are regarded to have been herbivorous, L. hamatus is a critical taxon for studies of the evolution of herbivory in early tetrapods. L. hamatus shares several trademark features of herbivorous adaptation with moradisaurines, which suggest that this captorhinid species was omnivorous. As such, it represents a transitional taxon between faunivorous basal reptiles and the herbivorous moradisaurines.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 237–262.     

A taxonomic revision of Encyclia (Orchidaceae: Laeliinae) in Costa Rica

A taxonomic revision of Encyclia (Orchidaceae) in Costa Rica is presented. The taxonomic history of the genus and its phylogenetic position are discussed. Characters of vegetative and floral morphology are described and their taxonomic significance is discussed. The genus is treated as comprising nine species in the country and a key to species is provided. Each taxon is described on the basis of Costa Rican material, illustrated in a composite plate, and its distribution within the country is assessed. Distribution maps for all the taxa are given. Overall distribution, derivation of name, synonymy, notes on species ecology and diagnostic features are presented for each taxon. The names Encyclia tonduziana and Epidendrum mooreanum are typified. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 395–448.     

The differentiation of bunodont Listriodontinae (Mammalia,Suidae) of Africa: new data from Kalodirr and Moruorot,Kenya

The early Miocene sites of Moruorot and Kalodirr (Kenya, 17.5 Myr) have yielded a rich collection of mammals. New listriodont material from these localities, including a complete skull and a partial mandible, provide long awaited information on cranial features of early bunodont Listriodontinae. The evolution and systematics of the group are highly debated, especially regarding its first representatives. The new material described here sheds light on the differentiation of bunodont Listriodontinae in Africa and clarifies the systematics of the group. The first phylogenetic analysis of the Listriodontinae is here performed and supports close relationships between Kubanochoerus and a clade (Eurolistriodon, Listriodon). Lopholistriodon is the most basal representative of the listriodontine clade. These first results stress the role of the African continent in the biogeographical history of the Listriodontinae. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 653–678.     

A reassessment of Eocene parrotlike fossils indicates a previously undetected radiation of zygodactyl stem group representatives of passerines (Passeriformes)

In the past years, various Eocene fossil birds were described as stem group representatives of the zygodactyl Psittaciformes (parrots). These birds show quite disparate morphologies, which cast some doubt on the correct assignment of all of them to the psittaciform stem group. A reassessment of their affinities is further needed, because it was recently proposed that among extant birds, Psittaciformes and Passeriformes (passerines) form a clade and that passerines possibly derived from a zygodactyl ancestor. Here, phylogenetic analyses are performed, which for the first time also include representatives of the Zygodactylidae, the extinct zygodactyl sister taxon of the Passeriformes. The early Eocene Psittacopes was originally described as a stem group representative of Psittaciformes. However, none of the present analyses supported psittaciform affinities for Psittacopes and instead recovered this taxon in a clade together with zygodactylids and passerines. Also part of this clade are the early Eocene taxa Pumiliornis and Morsoravis, and it is detailed that Psittacopes and the long‐beaked and presumably nectarivorous Pumiliornis, with which it has not yet been compared, are very similar in their postcranial osteology. The present analysis corroborates the hypothesis of a zygodactyl stem species of passerines. To account for these results, Psittacopes is here assigned to a new higher‐level taxon and a new name is also introduced for the clade including Zygodactylidae and Passeriformes.     

Cranial osteology of Exostinus serratus (Squamata: Anguimorpha), fossil sister taxon to the enigmatic clade Xenosaurus

Within the squamate clade Anguimorpha, Xenosaurus is an enigmatic taxon combining several apparently primitive features with a highly specialized set of autapomorphies. This combination makes the fossil record along the Xenosaurus stem particularly important for resolving the relationships of Xenosaurus with other anguimorphs, and between species of Xenosaurus.Exostinus serratus Cope, 1873 from the Oligocene of the western United States is currently hypothesized to be the immediate sister taxon to Xenosaurus. The cranial osteology of this pivotal taxon is described here for the first time from all known material, using high‐resolution X‐ray computerized tomography (CT) scanning to visualize individual elements. Exostinus serratus displays a mosaic of ancestral anguimorph features that are transformed in Xenosaurus, as well as unique synapomorphies shared with Xenosaurus. The region of the external nares is less transformed than in Xenosaurus, as are the general proportions of the bones surrounding the cartilaginous nasal capsule. However, the forms of the teeth and of the osteodermal sculpture, as well as several details of maxillary and mandibular morphology, are distinctly Xenosaurus‐like. Several autapomorphies are also present, including an abbreviated tooth row and an unusually wide palatal shelf of the maxilla. An understanding of the ways in which the non‐ossified tissues of the head influenced the development of the bones is crucial to interpreting their morphology. Furthermore, subtle anatomical features often provide important comparative information, and are emphasized herein. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 921–953.     

First Potwarmus from the Miocene of Saudi Arabia and the early phylogeny of murines (Rodentia: Muroidea)

Potwarmus is recorded for the first time in Saudi Arabia. The material comes from the Middle Miocene of the Hofuf Formation at Al‐Jadidah. This species, Potwarmus flynni sp. nov. , is described and compared with all species currently placed in the genera Potwarmus, Dakkamys, Paradakkamys, Vallaris, and some species of ‘Myocricetodon’ (i.e. the so‐called ‘primitive dendromurids’) as well as with two of the most primitive definitive murines (namely Antemus chinjiensis and Progonomys debruijni). Potwarmus flynni sp. nov. differs from the above‐mentioned species in having the anterior part of the m1 strongly reduced. A cladistic analysis provides evidence that this new Arabian taxon is close to Potwarmus primitivus and Potwarmus sp. nov. from Jebel Zelten (Libya), and also that this genus is unlikely to be the sister taxon to Antemus. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 664–679.