A new Patagonian species of Cricosaurus (Crocodyliformes,Thalattosuchia): first evidence of Cricosaurus in Middle–Upper Tithonian lithographic limestone from Gondwana

Abstract:  Upper Jurassic (Tithonian) deposits in north‐western Patagonia, Argentina, have yielded rich and taxonomically diverse assemblages of marine reptiles. These assemblages are also remarkable by their quality of preservation and are represented by ichthyosaurs, plesiosaurs, turtles and crocodyliforms. Despite the abundant crocodyliform record, only two metriorhynchid taxa have been identified: Cricosaurus araucanensis and Dakosaurus andiniensis. Here we described a new species of Cricosaurus, which represents the second species of Cricosaurus in the Tithonian of the Neuquén Basin, and the first metriorhynchid found in lithographic limestone from Gondwana. Furthermore, this specimen has the most complete postcranial remains of any metriorhynchid from South America. The new species is characterized by a short distance between the premaxilla and the nasal, a relatively narrow interorbital width, 23–25 mandibular teeth, bicarinated teeth with fine apicobasally aligned ridges, interalveolar spaces between the first seven teeth approximately 1.5 times longer than the anteroposterior diameter of the respective alveoli. To test the assignment of the new species to Cricosaurus, we carried out two phylogenetic analyses. In both analyses, Cricosaurus lithographicus sp. nov. is nested with other species referred to this genus. This new species has peculiar enamel ornamentation, characterized by numerous, fine apicobasally aligned ridges, when compared to other species of the genus.     

Cranial variation in Meriones tristrami (Rodentia: Muridae: Gerbillinae) and its morphological comparison with Meriones persicus,Meriones vinogradovi and Meriones libycus: a geometric morphometric study

Jirds (genus Meriones) comprise a group of rodents, of which the biodiversity is still poorly known. Reason for this is that several species of similar morphologies are known to occur sympatrically. In the north‐west of Iran, four such species occur: Meriones tristrami, Meriones persicus, Meriones vinogradovi and Meriones libycus, prone to several issues of taxonomical ambiguity. A proper characterization of morphological distinctiveness between these species, in relation to the variation within species, could provide the required information for species diagnosis and identification. As some cranial characters of M. tristrami, M. persicus and M. vinogradovi are quite similar, demarcations of species‐specific phenotypic variation have proven to be difficult. To tackle this problem, this study involves a geometric morphometric analysis of skull shape and size, incorporating a large representative sample of these four species, originating from most parts of their natural distribution range (especially for M. tristrami). It is first tested whether M. tristrami can be distinguished from the other sympatric species, and if so, to what degree the species shows a geoclimatic pattern in its skull shape and size when comparing different populations. The shape and size analyses show that M. libycus can be distinguished because of its largest skull and the relatively largest tympanic bulla, and that M. tristrami can be distinguished from the other species. At an intraspecific level in M. tristrami, the Iranian groups (Qazvin and west Iran) do not differ in shape among them, but do so in skull size. They could, however, be distinguished in skull shape from the non‐Iranian populations included (Turkey and Jordan). To what degree this continuous data can now be translated into discrete and diagnostic features, useful for taxonomic purposes, remains to be studied.     

A new stem-sperm whale (Cetacea, Odontoceti, Physeteroidea) from the Latest Miocene of Peru

A finely preserved skull with mandible and teeth associated, from the Latest Miocene beds (ca. 6 Ma) of the Pisco Formation, Sud-Sacaco, Peru, represents a new physeteroid genus and species, Acrophyseter deinodon. This moderate size sperm whale is characterized, among others, by: the short rostrum, the mandible distinctly curved upwards, large teeth very close together (12 on each upper tooth row and 13 on each lower tooth row), the lateral margin of the maxilla along the rostrum base much lower than the orbit roof, a wide supracranial basin dorsally overhanging the right orbit and limited to the cranium and a large temporal fossa dorsomedially elevated. A preliminary cladistic analysis provides a phylogenetic position of Acrophyseter nested within the stem-Physeteroidea, more basal than the clade Kogiidae + Physeteridae. The morphology of the oral apparatus and of the temporal fossa suggests that Acrophyseter was able to feed on large preys.     

A NEW SPECIES OF DICERORHINUS (RHINOCEROTIDAE) FROM THE PLIO‐PLEISTOCENE OF MYANMAR

Abstract: A skull and mandible of the new species Dicerorhinus gwebinensis sp. nov. of Rhinocerotidae (Mammalia, Perissodactyla) is described. The material is collected from the upper part of the Irrawaddy sediments (Plio‐Pleistocene) in central Myanmar. D. gwebinensis sp. nov. is morphologically more similar to the extant species D. sumatrensis (Sumatran rhinoceros) than to other species of the genus but differs from D. sumatrensis in having the comparatively shorter nasal, the more concave dorsal profile of the skull, the more elevated occiput and presence of molar crista in M3/. This is the first discovery of Dicerorhinus in the upper Miocene to lower Pleistocene of the Indian subcontinent and Mainland Southeast Asia, and fills the chronological and geographical gap of this lineage in Asia. The Dicerorhinus clade probably migrated into Southeast Asia from East Asia by the Pliocene or early Pleistocene. This hypothesis is supported by the scarcity or absence of this clade in the Neogene mammalian fauna of the Indian Subcontinent.     

Evidence for the teleosaurid crocodylomorph genus Machimosaurus in the Kimmeridge Clay Formation (Late Jurassic) of England

We report the only definite specimen of the teleosaurid crocodylomorph genus Machimosaurus from the Kimmeridge Clay Formation of England. This specimen (an isolated tooth) is now the only evidence of Machimosaurus in the Kimmeridge Clay Formation because a large skull and mandible, previously considered to be of Machimosaurus mosae, was recently shown to pertain to a metriorhynchid crocodylomorph. The tooth described herein was originally figured in 1884 as a tooth crown from a metriorhynchid crocodylomorph. However, its conical shape, blunt apex and distinctive enamel ornamentation are characteristic of the teleosaurid genus Machimosaurus. That Machimosaurus, and teleosaurids in general, were so rare in the Kimmeridge Clay Formation suggests that these marine crocodylomorphs did not commonly use this seaway. Their rarity is in contrast to contemporaneous deposits from continental Europe where teleosaurid remains, including Machimosaurus, are far more common. These continental deposits were deposited in shallow-marine/brackish ecosystems, suggesting that teleosaurids were largely restricted to coastal marine environments.     

A new species of the enigmatic archosauromorph Doswellia from the Upper Triassic Bluewater Creek Formation,New Mexico,USA

Abstract: Doswellia sixmilensis is a new species of the doswelliid archosauromorph genus Doswellia named for an incomplete skeleton from the Upper Triassic Bluewater Creek Formation of the Chinle Group in west‐central New Mexico, USA. D. sixmilensis differs from D. kaltenbachi Weems, the type and only other known species of Doswellia, in its larger size, higher tooth count and greater heterodonty, possession of keels on the cervical centra and the presence of discrete knobs or spikes on some osteoderms. The holotype of D. sixmilensis is the fourth occurrence of Doswellia and only the second occurrence of a Doswellia skull, which includes the previously unknown premaxilla and maxilla (and therefore the best dentition) and has the best‐preserved cervical vertebrae. Although it adds to our knowledge of the anatomy of Doswellia, this new information does not alter previous concepts of the phylogenetic relationships of the doswelliid genera, largely because they are so poorly known anatomically. The genus Doswellia is known from the Newark Supergroup in Virginia, and the Chinle Group in Texas, New Mexico and Utah, in strata of Otischalkian–Adamanian age. The type locality of D. sixmilensis is c. 43 m stratigraphically below a bed from which U‐Pb dating of detrital zircons yields a maximum depositional age of c. 220 Ma, so this is a reasonable approximate numerical age for D. sixmilensis.     

A new species of Acynodon (Crocodylia) from the upper cretaceous (Santonian–Campanian) of Villaggio del Pescatore,Italy

Abstract: The new species Acynodon  adriaticus is described on the basis of remains from the Santonian–Campanian of Villaggio del Pescatore (Trieste, NE Italy). This species differs in several cranial features from Acynodon  iberoccitanus, the only other Acynodon species whose cranial osteology is known in detail. The absence of maxillary and dentary caniniform teeth coupled with the presence of enlarged molariform teeth suggests that Acynodon probably fed on slowly moving hard‐shelled prey. Moreover, the new materials reveal for the first time the morphology of some postcranial elements of Acynodon: in particular, medial‐most paravertebral osteoderms that are characterized by two keels. A new cladistic phylogenetic analysis resolves the previously reported polytomy among the basal Globidonta: Acynodon is recognized as the most primitive globidontan. This genus may represent the geologically oldest known globidontan. The fact that Acynodon has been found only in Europe and that the outgroup of Globidonta, the Diplocynodontinae, is mainly known from Europe, suggests that globidontans may have originated in Europe and not in North America as previously supposed.     

The snout of Cricosaurus araucanensis: a case study in novel anatomy of the nasal region of metriorhynchids

Metriorhynchids are the only crocodyliforms adapted to pelagic marine life. Snout natural endocasts of the Tithonian (Late Jurassic) metriorhynchid Cricosaurus araucanensis indicated that skeletal changes defining the peculiar metriorhynchid body plan were coupled with changes of the soft cephalic anatomy such as the enlarged salt glands and restructuring of the paranasal sinus system. Seven new natural endocasts of the snout and a 3‐D reconstruction of C. araucanensis are described. Data from these casts and the reconstruction are congruent, and they are combined into an accurate reconstruction that improves our knowledge of the pre‐orbital anatomy. The olfactory tract, bulbs, olfactory nasal region and the anterior extension of the antorbital sinus within the maxilla are recognized. Osteological correlates of the salt gland body are also proposed. Palaeobiological inferences are erected based on the integration of natural endocasts and 3‐D reconstruction data. It is proposed that C. araucanensis nasal salt glands were highly vascularized with a blood supply comparable with those of extant marine birds. Reduced olfactory bulbs and olfactory nasal region indicate that the aerial olfaction, differing from extant crocodilians, was not well developed.     

Uncovering cryptic diversity of Lyngbya: the new tropical marine cyanobacterial genus Dapis (Oscillatoriales)

Cyanobacteria comprise an extraordinarily diverse group of microorganisms and, as revealed by increasing molecular information, this biodiversity is even more extensive than previously estimated. In this sense, the cyanobacterial genus Lyngbya is a highly polyphyletic group composed of many unrelated taxa with morphological similarities. In this study, the new genus Dapis was erected from the genus Lyngbya, based on a combined molecular, chemical, and morphological approach. Herein, two new species of cyanobacteria are described: D. pleousa and D. pnigousa. Our analyses found these species to be widely distributed and abundant in tropical and subtropical marine habitats. Seasonally, both species have the ability to form extensive algal blooms in marine habitats: D. pleousa in shallow‐water, soft bottom habitats and D. pnigousa on coral reefs below depths of 10 m. Electron microscopy showed that D. pleousa contains gas vesicles, a character not previously reported in Lyngbya. These gas vesicles, in conjunction with a mesh‐like network of filaments that trap oxygen released from photosynthesis, provide this species with an unusual mechanism to disperse in coastal marine waters, allowing D. pleousa to be present in both benthic and planktonic forms. In addition, both D. pleousa and D. pnigousa contained nitrogen‐fixing genes as well as bioactive secondary metabolites. Several specimens of D. pnigousa biosynthesized the secondary metabolite lyngbic acid, a molecule that has also been isolated from many other marine cyanobacteria. Dapis pleousa consistently produced the secondary metabolite malyngolide, which may provide a promising chemotaxonomic marker for this species.     

The <Emphasis Type="Italic">Eumyarion</Emphasis> (Mammalia,Rodentia, Muridae) assemblage from Sandelzhausen (Miocene,Southern Germany): a test on homogeneity

A large sample of Eumyarion cheek teeth (N = 569) from the Early/Middle Miocene boundary locality Sandelzhausen (MN5, Southern Germany), type locality E. bifidus Fahlbusch, 1964, is studied. It is concluded that this collection contains two species: E. bifidus and E. weinfurteri Schaub and Zapfe, 1953. The similarity in size and morphology of the cheek teeth of these two species is so great that only the M1 and M2 can be recognised with certainty. Eumyarion bifidus seems to be a descendant of E. orhani de Bruijn et al., 2006 from the Early Miocene (MN3) of Southwestern Anatolia and is therefore considered to be an immigrant into Central Europe.      

Cranial anatomy,taxonomic implications and palaeopathology of an Upper Jurassic Pliosaur (Reptilia: Sauropterygia) from Westbury,Wiltshire, UK

Abstract: Complete skulls of giant marine reptiles of the Late Jurassic are rare, and so the discovery of the 1.8‐m‐long skull of a pliosaur from the Kimmeridge Clay Formation (Kimmeridgian) of Westbury, Wiltshire, UK, is an important find. The specimen shows most of the cranial and mandibular anatomy, as well as a series of pathological conditions. It was previously referred to Pliosaurus brachyspondylus, but it can be referred reliably only to the genus Pliosaurus, because species within the genus are currently in need of review. The new specimen, together with another from the same locality, also referred to P. brachyspondylus, will be crucial in that systematic revision, and it is likely that the genus Pliosaurus will be found to include several genera. The two Westbury Pliosaurus specimens share many features, including the form of the teeth, but marked differences in the snout and parietal crest suggest sexual dimorphism; the present specimen is probably female. The large size of the animal, the extent of sutural fusion and the pathologies suggest this is an ageing individual. An erosive arthrotic condition of the articular glenoids led to prolonged jaw misalignment, generating a suite of associated bone and dental pathologies. Further damage to the jaw joint may have been the cause of death.     

Tooth and cranial disparity in the fossil relatives of Sphenodon (Rhynchocephalia) dispute the persistent ‘living fossil’ label

The tuatara (Sphenodon punctatus) is the only living representative of Rhynchocephalia, a group of small vertebrates that originated about 250 million years ago. The tuatara has been referred to as a living fossil; however, the group to which it belongs included a much greater diversity of forms in the Mesozoic. We explore the morphological diversity of Rhynchocephalia and stem lepidosaur relatives (Sphenodon plus 13 fossil relatives) by employing a combination of geometric morphometrics and comparative methods. Geometric morphometrics is used to explore cranium size and shape at interspecific scale, while comparative methods are employed to test association between skull shape and size and tooth number after taking phylogeny into account. Two phylogenetic topologies have been considered to generate a phylomorphospace and quantify the phylogenetic signal in skull shape data, the ancestral state reconstruction as well as morphological disparity using disparity through time plots (DTT). Rhynchocephalia exhibit a significant phylogenetic signal in skull shape that compares well with that computed for other extinct vertebrate groups. A consistent form of allometry has little impact on skull shape evolution while the number of teeth significantly correlates with skull shape also after taking phylogeny into account. The ancestral state reconstruction demonstrates a dramatic shape difference between the skull of Sphenodon and its much larger Cretaceous relative Priosphenodon. Additionally, DTT demonstrates that skull shape disparity is higher between rather than within clades while the opposite applies to skull size and number of teeth. These results were not altered by the use of competing phylogenic hypotheses. Rhynchocephalia evolved as a morphologically diverse group with a dramatic radiation in the Late Triassic and Early Jurassic about 200 million years ago. Differences in size are not marked between species whereas changes in number of teeth are associated with co‐ordinated shape changes in the skull to accommodate larger masticatory muscles. These results show that the tuatara is not the product of evolutionary stasis but that it represents the only survivor of a diverse Mesozoic radiation whose subsequent decline remains to be explained.     

A new cryptoclidid Plesiosaur from the Oxfordian (Late Jurassic) of Cuba

Cryptocleidus ? cuervoi caroliDE LA TORRE & ROJAS, 1949, a partial skull with associated mandible and atlas-axis from the Oxfordian of Cuba, has been completely prepared for the first time. It is here redescribed and assigned to a new cryptoclidid genus, Vinialesaurus, for which the species caroli is retained. Vinialesaurus is mainly diagnosed by palatal characters such as double internal nares, an anteriorly rounded vomer and a lack of anterior interpterygoid vacuities. It shares with other cryptoclidids large orbits and external nares, a small vertical jugal and a reduced tooth ornamentation. The occurrence of Vinialesaurus caroli in association with pliosauroids, ophthalmosaurian ichthyosaurs, metriorhynchid crocodilians and pleurodiran marine turtles, strongly suggests that a marine seaway was present in the Caribbean during the Oxfordian, connecting the western Tethys with the Oriental Pacific.     

A SKULL OF THE GIANT BONY‐TOOTHED BIRD DASORNIS (AVES: PELAGORNITHIDAE) FROM THE LOWER EOCENE OF THE ISLE OF SHEPPEY

Abstract: The first substantial skull of a very large Paleogene bony‐toothed bird (Pelagornithidae) is described from the Lower Eocene London Clay of the Isle of Sheppey in England. The specimen is assigned to Dasornis emuinus (Bowerbank), based on a taxonomic revision of the large London Clay Pelagornithidae. Very large bony‐toothed birds from the London Clay were known previously from fragmentary remains of non‐comparable skeletal elements only, and Dasornis londinensis Owen, Argillornis emuinus (Bowerbank), A. longipennis Owen, and Neptuniavis miranda Harrison and Walker are considered junior synonyms of D. emuinus. The new specimen allows a definitive assignment of Dasornis to the Pelagornithidae and documents that this taxon closely resembles other bony‐toothed birds in cranial morphology. It is hypothesized that giant size (i.e. a wingspan above 4 m) evolved only once within Pelagornithidae and that Dasornis emuinus is the sister taxon of the giant Neogene bony‐toothed birds, which share a derived wing morphology.     

A new large philisid (Mammalia,Chiroptera, Vespertilionoidea) from the late Early Eocene of Chambi,Tunisia

Abstract: Among the new dental remains from the late Early Eocene of Chambi (Kasserine area, Tunisia) is a large‐sized upper molar of a new bat species, Witwatia sigei nov. sp. (Chiroptera, Vespertilionoidea, Philisidae), described herein. The locality of Chambi has revealed evidence for an early appearance of two modern microchiropteran superfamilies in Africa: Dizzya exsultans, a Philisidae, which is considered to be an archaic Vespertilionoidea, and an indeterminate Rhinolophoidea. In addition to D. exsultans, the new species, W. sigei, is the second representative of the Philisidae in this locality. W. sigei extends back to the late Early Eocene the occurrence of the genus Witwatia, which was previously only reported from the early Late Eocene of the Fayum (BQ‐2, Egypt). By analogy with the largest extant microbats, the large size of Witwatia suggests a tendency to the opportunistic diet of this taxon, thereby contrasting with the strict insectivory characterizing primitive bats found in other continents in the same epoch.     

Plastid phylogenetics of Oceania yams (Dioscorea spp., Dioscoreaceae) reveals natural interspecific hybridization of the greater yam (D. alata)

Phylogenetic relationships of Oceanian staple yams (species of Dioscorea section Enantiophyllum) were investigated using plastid trnL‐F and rpl32‐trnL^(UAG) sequences and nine nuclear co‐dominant microsatellites. Analysis of herbarium specimens, used as taxonomic references, allowed the comparison with samples collected in the field. It appears that D. alata, D. transversa and D. hastifolia are closely related species. This study does not support a direct ancestry from D. nummularia to D. alata as previously hypothesized. The dichotomy in D. nummularia previously described by farmers in semi‐perennial and annual types was reflected by molecular markers, but the genetic structure of D. nummularia appears more complex. Dioscorea nummularia displayed two haplotypes, each corresponding to a different genetic group. One, including a D. nummularia voucher from New Guinea, is closer to D. tranversa, D. alata and D. hastifolia and encompasses only semi‐perennial types. The second group is composed of semi‐perennial and annual yams. However, some of these annual yams also displayed D. alata haplotypes. Nuclear markers revealed that some annual yams shared alleles with D. alata and semi‐perennial D. nummularia, suggesting a hybrid origin, which may explain their intermediate morphotypes and the difficulty met in classifying them.     

Environment and space as drivers of variation in skull shape in two widely distributed South‐American Tayassuidae,Pecari tajacu and Tayassu pecari (Mammalia: Cetartiodactyla)

The influence of the environment on the geographical variation of morphological traits has been recognized in a number of taxa. Pecari tajacu and Tayassu pecari are ideal models to investigate intraspecific geographic variation in skull because of their wide and heterogeneous geographical distribution in South America. We used geometric morphometric procedures to examine the geographical variation in skull shape of 294 adult specimens of these species from 134 localities. We quantified to what extent skull shape variation was explained by environment, skull size and geographical space using variation partitioning analysis. We detected a strong pattern of geographic variation for P. tajacu skull shape, but not for T. pecari. The environment seems to be the major selective force that drives skull shape variation in both species. Nevertheless, other spatially structured processes (e.g. genetic drift, gene flow) might also have affected variation in the skull shape of the more widespread species P. tajacu. Allometric relationships might reflect the biomechanical constraints that are thought to be strong enough to limit size‐related changes in T. pecari skull shape.     

Speciation and dietary specialization in Drupa,a genus of predatory marine snails (Gastropoda: Muricidae)

Claremont, M., Reid, D.G. & Williams, S.T. (2012) Speciation and dietary specialization in Drupa, a genus of predatory marine snails (Gastropoda: Muricidae). —Zoologica Scripta, 41, 137–149. We test the competing predictions of allopatric speciation and of ecological speciation by dietary specialization in Drupa, an Indo‐Pacific genus of carnivorous marine gastropods in the family Muricidae. We use a well‐resolved molecular phylogeny (reconstructed from one nuclear and two mitochondrial genes) to show the validity of the traditional species D. elegans, D. rubusidaeus, D. clathrata, D. morum and D. speciosa.‘Drupa ricinus’ is shown to consist of three species: D. ricinus s. s., D. albolabris and a new species, possibly endemic to Japan. ‘Purpura’aperta is transferred to Drupa. Despite potential widespread dispersal and a high degree of range overlap among sister species, range sizes between sister species are highly asymmetric, suggesting that speciation has been predominately peripatric. The exception is the sister pair D. ricinus s. s. and D. albolabris, which have symmetric range sizes and are sympatric over broad Indo‐Pacific ranges. Such symmetry and extensive sympatry are contrary to the predictions of the (peripatric) allopatric model of speciation. Nevertheless, contrary to the predictions of an ecological speciation model based upon dietary specialization, broad dietary range appears to be identical between the species. Small differences in microhabitat preferences (or hypothetical dietary specialization at a fine taxonomic scale) may have been significant in the speciation process or, if initial divergence was allopatric, in permitting subsequent sympatry. Broad dietary shifts appear to have accompanied more ancient divergences within the genus Drupa.     

Molecular phylogenetic and morphological studies on the systematic position of Heracula discivitta reveal a new subfamily of Pseudobistonidae (Lepidoptera: Geometroidea)

Heracula discivitta Moore is an uncommon moth species currently recorded from India, Nepal and China. Although this species has traditionally been placed in Lymantriinae, its systematic position in Macroheterocera has been enigmatic due to its unique morphological features. Here we used molecular and morphological data to explore the systematic position of H. discivitta. Our molecular phylogenetic analyses indicate that this species is sister to Pseudobiston pinratanai Inoue, a member of a recently established monotypic family Pseudobistonidae. The examinations of morphological features further show that H. discivitta shares synapomorphies with Pseudobistonidae. Based on the analysis results, we propose a new subfamily of Pseudobistonidae (Heraculinae subfam.n. ) to accommodate H. discivitta. The resemblance of the habitus to that of the brahmaeid genus Calliprogonos Mell & Hering is discussed. This published work has been registered on ZooBank, http://zoobank.org/urn:lsid:urn:lsid:zoobank.org:pub:63D17850‐6D51‐4E03‐A5D6‐F9EF6E7AF402 .