Functional morphology of the hindlimb of fossilized pygmy hippopotamus from Ayia Napa (Cyprus).

We studied the functional morphology of the postcranial skeleton of the endemic hippopotamus Phanourios minor, derived from the Upper Pleistocene site of Ayia Napa. The deposit, which consists of a hard limestone substrate on which the species moved, has yielded a great abundance of hippopotamus material, making the Ayia Napa locality one of the most important paleontological sites in Cyprus. The immigration of the large-sized mainland Hippopotamus to Cyprus led to the emergence of a new insular species with its main characteristic being the extremely reduced body size. In this study, all the hindlimb elements of the Cypriot hippo are described in detail and compared with those of the modern species, with the extant Hippopotamus amphibius being considered similar to the possible ancestor of P. minor. In some cases, the morphological comparison is reinforced using bones of other extinct insular and mainland hippos. Additionally, we provided a functional analysis of the hindlimb joints, suggesting specific locomotor habits for the species. The anatomical examination reveals that the elements in P. minor are robust with marked muscular insertion areas resembling those found in Hippopotamus. However, there are also similarities with Choeropsis liberiensis in certain morphofunctional traits. P. minor adapted to slow but powerful locomotion with remarkable stabilization, particularly in the zeugopodium and the autopodium. The knee was less mobile in the craniocaudal direction compared with that in recent hippos, while the abduction-adduction movements of the thigh were advanced. The pes presented good mobility in the sagittal plane and limitation in transversal movements. Thus, P. minor displayed modifications to its limbs, influenced by the mountainous island environment and the body size reduction, resulting in specialized locomotion, which was different from that of extant hippopotamuses.

Picking up the threads: Comparative osteology and associated cartilaginous elements for members of the genus Trilepida Hedges, 2011 (Serpentes, Leptotyphlopidae) with new insights on the Epictinae systematics.

The threadsnakes of the family Leptotyphlopidae have been historically neglected in terms of their natural history, ecology, systematics, and morphology. Given the relevance of morphological data for resolving systematic, evolutionary, and functional issues, we aimed to provide a detailed comparative description of osteology and associated cartilaginous elements for members of the genus Trilepida. Data were obtained through high-resolution computed tomography images, cleared and stained specimens, and radiography images of a total of 47 specimens and 12 species. Both cranial and axial osteology characters exhibited a relevant degree of intraspecific variation regarding qualitative and quantitative data associated with skull and vertebrae foramina and the shape of bony sutures and processes. The high representativeness of examined species and specimens allows us to provide a comprehensive discussion on the inter- and intraspecific osteological variation, as well as a compelling osteological diagnosis for the entire genus. Trilepida spp. differ from all Epictinae by the presence of the following combination of characters: paired nasals, fused supraoccipitals (distinct from parietal, prootics, and otooccipitals), a single (fused) parietal without a dorsal fontanelle, and the basioccipital participating in the foramen magnum (except in Trilepida nicefori). Our results reinforce the need for integration of detailed anatomical traits to usually conserved external morphological data to provide accurate diagnostic features for Epictinae. In addition, new phylogenetic hypotheses or even taxonomic re-allocations may broadly benefit from these detailed comparative studies.

Postnatal ossification sequences in Acrocephalus scirpaceus and Chroicocephalus ridibundus (Aves: Neognathae): The precocial-altricial spectrum and evolution of compound bones in birds.

Although the development of the avian skeleton has attracted considerable attention, most of the studies have been concentrated on the embryonic period, while studies on the postnatal period are rare. We studied the postnatal development of the skeleton in two phylogenetically distant birds, an altricial passerine Acrocephalus scirpaceus and a semiprecocial charadriiform Chroicocephalus ridibundus. The neonates of the former, despite being altricial, have well-ossified skeleton-the degree of development approaches that of the semiprecocial gull. However, after hatching the limb bones (particularly those of the hind limb) ossify earlier in the gull which is probably related to faster acquisition of locomotor abilities. We have observed that, in contrast to previous reports from neognathous birds, in the ankle of the gull, the ascending process fuses with the astragalus rather than with the calcaneum. This type of development is present in palaeognaths and nonavian dinosaurs but has not yet been reported in neognaths. This indicates a greater diversity within Neognathae and suggests a more complex scenario for the evolution of the avian ankle. However, data from a greater number of species are needed to establish the developmental sequence ancestral for neognathous birds. Furthermore, the sequence of bone fusions in the wrist of Acrocephalus is similar to the fossil-documented evolutionary sequence observed in the phylogeny of early birds, with the semilunate carpal and major metacarpal fusing first, followed by the alular metacarpal fusing with the major metacarpal and then the major and minor metacarpal fusing proximally. These data underscore the importance of developmental studies for reconstructing the evolutionary history.

Femur associated with the Deep Skull from the West Mouth of the Niah Caves (Sarawak, Malaysia).

The skeletal remains of Pleistocene anatomically modern humans are rare in island Southeast Asia. Moreover, continuing doubts over the dating of most of these finds has left the arrival time for the region's earliest inhabitants an open question. The unique biogeography of island Southeast Asia also raises questions about the physical and cultural adaptations of early anatomically modern humans, especially within the setting of rainforest inhabitation. Within this context the Deep Skull from the West Mouth of the Niah Caves continues to figure prominently owing to its relative completeness and the greater certainty surrounding its geological age. Recovered along with this partial cranium in 1958 were several postcranial bones including a partial femur which until now has received little attention. Here we provide a description and undertake a comparison of the Deep Skull femur finding it to be very small in all of its cross-sectional dimensions. We note a number of size and shape similarities to the femora of Indigenous Southeast Asians, especially Aeta people from the Philippines. We estimate its stature to have been roughly 145-146 cm and body mass around 35 kg, confirming similarities to Aeta females. Its extreme gracility indicated by low values for a range of biomechanical parameters taken midshaft meets expectations for a very small (female) Paleolithic East Asian. Interestingly, the second moment of area about the mediolateral axis is enlarged relative to the second moment of area about the anteroposterior axis, which could potentially signal a difference in activity levels or lifestyle compared with other Paleolithic femora. However, it might also be the result of sexual dimorphism in these parameters as well as possibly reflecting changes associated with aging.

Vertebral Comparative Anatomy and Morphological Differences in Anguine Lizards With a Special Reference to Pseudopus apodus.

The article reports on the first detailed vertebral and rib morphology of anguine taxon Pseudopus apodus using micro-computed tomography. A comparison shows significant morphological differences of vertebrae of Pseudopus relative to those of Anguis and Ophisaurus. Usually, there are 55 presacral vertebrae, two sacral, and 95-97 caudal vertebrae. Pseudopus apodus can be defined by 23 diagnostic features concerning the vertebral column. Although zygapophyseal articulation between atlas and axis is well developed in limbed anguid gerrhonotine lizards like Abronia or Barisia, it is absent in the extant representatives of the clade Anguinae, which are limbless. Thus, our study brings further support to the hypothesis about the complete reduction of this articulation in forms with reduced or absent limbs. Comparison of adult and juvenile morphology of vertebrae of P. apodus was also analyzed. Heterochrony in the evolution of this taxon was previously confirmed by its skull morphology and it can be also documented on the basis of vertebrae. Our data suggest that a peramorphic heterochronic process played a role in the evolution of this largest extant anguine species. Geometric morphometric analyses revealed a pattern of high vertebral disparity among species. We found a clear separation of limbless forms in morphospace. Pseudopus apodus always clusters within Ophisaurus-species confirming molecular and some morphological phylogenies. Only the first tail vertebra shows a distinct difference to those of other anguids, which might be related to altered locomotion associated to the larger body size in this species. Anat Rec, 302:232-257, 2019. © 2018 Wiley Periodicals, Inc.

External morphology, postcranial and appendicular osteology of three southwestern Atlantic flatfishes ( Paralichthys, Paralichthyidae), and comparisons with other congeneric species

Comparisons of the external morphology and analysis of osteological features of the postcranial and appendicular skeletons of three southwestern Atlantic flatfish species of the genus Paralichthys (P. isosceles, P. orbignyanus and P. patagonicus) were carried out. Bones are described, and detailed morphological, morphometric and meristic characteristics of these flounders are given in order to provide information about the external and internal morphology of three species of Paralichthys occurring in the south-west Atlantic waters that add new information and will help regarding within the framework of a phylogenetic study of the group. Interspecific differences were found in the number of vertebrae and intermuscular bones, as well as in the morphology and morphometry of vertebrae, caudal skeletons, pectoral and pelvic girdle bones. Relationships between bones are discussed and bone characteristics compared with those found in other species of Paralichthys and in other pleuronectiform species. The position of Paralichthys isosceles within Paralichthys is discussed, along with other congeners such as P. triocellatus and P. oblongus.(AU)

Se llevaron a cabo comparaciones de la morfología externa y el análisis de las características osteológicas de los esqueletos postcraneal y apendicular de tres especies de peces planos del Atlántico sudoccidental del género Paralichthys (P. isosceles, P. orbignyanus y P. patagonicus). Se describen los huesos, y se proporcionan características morfológicas, morfométricas y merísticas detalladas de estos lenguados con el fin de aportar información sobre la morfología externa e interna de tres especies de Paralichthys presentes en el Atlántico sudoccidental. Esta nueva información contribuirá al marco de un estudio filogenético del grupo. Se encontraron diferencias interespecíficas en el número de vértebras y huesos intermusculares, así como en la morfología y morfometría de las vértebras, los esqueletos caudales, los huesos de las cinturas pectoral y pélvica. Se discuten las relaciones entre los huesos y las características óseas en comparación con las encontradas en otras especies de Paralichthys y de otros Pleuronectiformes. Se discute la posición de Paralichthys isosceles dentro del género Paralichthys, junto con otros congéneres como P. oblongus y P. triocellatus.(AU)

The postcranial skeleton of Boreogomphodon (Cynodontia: Traversodontidae) from the Upper Triassic of North Carolina, USA and the comparison with other traversodontids.

Postcranial remains of Boreogomphodon from the Upper Triassic of North Carolina are described and compared to those of other known traversodontid cynodonts. The postcranial skeleton of Boreogomphodon is characterized by four sacral ribs, simple ribs lacking costal plates, the extension of the scapular neck below the acromion process, a short scapular facet on the procoracoid, a concave anterior margin of the procoracoid, humerus entepicondyle with smooth corner, and the presence of a fifth distal carpal. Four types of ribs are identified among traversodontids: 'normal' form, tubercular rib, costal plate, and Y-shaped rib. Fossorial behavior is suggested for traversodontids with elaborate costal plates. Within Traversodontidae, the procoracoid is relatively small; the anterior process of the iliac blade extends anteroventrally to different degrees in different taxa, which facilitates retraction of the femur; and the limb bones show allometric growth in terms of length and width.

Reappraisal of Synagrops Günther, 1887 with rehabilitation and revision of Parascombrops Alcock, 1889 including description of seven new species and two new genera (Perciformes: Acropomatidae).

An ongoing review of the fishes of the basal percoid family Acropomatidae has revealed that the genus Synagrops Günther, 1887 as it is currently understood is not a natural group. Species with a serrated pelvic-fin spine are here placed in the resurrected genus Parascombrops Alcock, 1889 (type-species: Parascombrops pellucidus Alcock, 1889), and the new, monospecific genus Caraibops n. gen. (type-species: Synagrops trispinosus Mochizuki & Sano, 1984). Parascombrops is unique amongst Acropomatidae in the combination of the presence of vacant 8th interneural space, a predorsal formula /0+0/0+2/ and an epaxialis attachment type 1. Caraibops n. gen. shares none of these characters and further differs from Parascombrops by an anal-fin formula of III + 9 (vs II + 7 or III + 6), and the absence of denticles on the ectopterygoid. Parascombrops is revised and now contains a total of 13 species, including 7 new: P. analis (Katayama, 1957), P. argyreus (Gilbert & Cramer, 1897), P. glossodon n. sp., P. madagascariensis n. sp., P. mochizukii n. sp., P. nakayamai n. sp., P. ohei n. sp., P. parvidens n. sp., P. pellucidus Alcock, 1889, P. philippinensis (Günther, 1880), P. serratospinosus (Smith & Radcliffe, 1912), P. spinosus (Schultz, 1940) and P. yamanouei n. sp. Synagrops adeni Kotthaus, 1970 and S. malayanus Weber, 1913 are treated as synonyms of P. pellucidus and P. philippinensis, respectively. Lectotypes are designated for P. philippinensis and S. malayanus. The main characters used to distinguish between the species of Parascombrops are: serration of other fin spines, number of gill rakers and pseudobranchial filaments, head profile, presence or absence of ridges on the preopercle, shape of 1st anal-fin pterygiophore, dentition on vomer, palatines and ectopterygoids, orbit diameter, pectoral-fin length, maximal body depth and otolith morphology. The genus Synagrops is here confined to two species, S. japonicus (Döderlein, 1883) and S. bellus (Goode & Bean, 1896), characterized by the apomorphic character of an otic capsule with a posteriorly open myodome, a basioccipital fossa and a very specialized otolith morphology. Synagrops is also characterized by the absence of pelvic-fin spine serrations. Two other species without a serrated pelvic-fin spine, originally described in Synagrops, are removed from this genus. Synagrops microlepis Norman, 1935 is separated into the monotypic Kaperangus n. gen., the only genus in the family with two supraneurals (cf. three in all other taxa). The second, Synagrops pseudomicrolepis Schultz, 1940 is re-assigned to the genus Verilus.        The geographic distribution of Parascombrops as currently composed is discussed, and is shown to be primarily of West Pacific nature, with few species in the Indian Ocean and one in the tropical West-Atlantic (P. spinosus). The West Atlantic species Parascombrops spinosus is very closely related to P. mochizukii from the tropical northwestern Pacific, and the implications of this disjunct distribution are discussed. The high degree of speciation now recognized in Parascombrops species of the West-Pacific indicates that a diverse ecological adaptation within an overall pseudoceanic habitat may have played a major role in speciation, which would have remained obscured without adequate taxonomic resolution.        Fossil, otolith-based records are also briefly discussed in the context. The extant Parascombrops argyreus and P. ohei are reported from the Pliocene of Japan, and Caraibops trispinosus has been recorded from the Pliocene of Venezuela.

The postcranial skeletal maturation of Australopithecus sediba.

OBJECTIVES: In 2008, an immature hominin defined as the holotype of the new species Australopithecus sediba was discovered at the 1.9 million year old Malapa site in South Africa. The specimen (MH1) includes substantial post-cranial skeletal material, and provides a unique opportunity to assess its skeletal maturation. METHODS: Skeletal maturity indicators observed on the proximal and distal humerus, proximal ulna, distal radius, third metacarpal, ilium and ischium, proximal femur and calcaneus were used to assess the maturity of each bone in comparison to references for modern humans and for wild chimpanzees (Pan troglodytes). RESULTS: In comparison to humans the skeletal maturational ages for Au. sediba correspond to between 12.0 years and 15.0 years with a mean (SD) age of 13.1 (1.1) years. In comparison to the maturational pattern of chimpanzees the Au. sediba indicators suggest a skeletal maturational age of 9-11 years. Based on either of these skeletal maturity estimates and the body length at death of MH1, an adult height of 150-156 cm is predicted. DISCUSSION: We conclude that the skeletal remains of MH1 are consistent with an ape-like pattern of maturity when dental age estimates are also taken into consideration. This maturity schedule in australopiths is consistent with ape-like estimates of age at death for the Nariokotome Homo erectus remains (KMN-WT 15000), which are of similar postcranial immaturity to MH1. The findings suggest that humans may have distinctive and delayed post-cranial schedules from australopiths and H. erectus, implicating a recent evolution of somatic and possibly life history strategies in human evolution.

Osteological postcranial traits in hylid anurans indicate a morphological continuum between swimming and jumping locomotor modes.

Anurans exhibit a particularly wide range of locomotor modes that result in wide variations in their skeletal structure. This article investigates the possible correlation between morphological aspects of the hylid postcranial skeleton and their different locomotor modes and habitat use. To do so, we analyzed 18 morphometric postcranial variables in 19 different anuran species representative of a variety of locomotor modes (jumper, hopper, walker, and swimmer) and habitat uses (arboreal, bush, terrestrial, and aquatic). Our results show that the evolution of the postcranial hylid skeleton cannot be explained by one single model, as for example, the girdles suggest modular evolution while the vertebral column suggests other evolutionary modules. In conjunction with data from several other studies, we were able to show a relationship between hylid morphology and habitat use; offering further evidence that the jumper/swimmer and walker/hopper locomotor modes exhibit quite similar morphological architecture. This allowed us to infer that new locomotor modalities are, in fact, generated along a morphological continuum. J. Morphol. 278:403-417, 2017. © 2017 Wiley Periodicals, Inc.

A new species of Neobathyclupea from the northern Indian Ocean with comments on N. malayana (Teleostei, Perciformes, Bathyclupeidae).

A new species of Neobathyclupea is described from two specimens collected off Myanmar and off Socotra Island. The new species is most similar to N. malayana, but differs from it and other congeners in having jet-black coloured fins, larger pseudobranch, dorsal-pterygiophore pattern and some body proportions. Neobathyclupea malayana is re-diagnosed. Individual variations in armament of the preopercle and in the dorsal-pterygiophore patterns within Bathyclupeidae are discussed.

Taxonomic identification of Lower Pleistocene fossil hominins based on distal humeral diaphyseal cross-sectional shape.

The coexistence of multiple hominin species during the Lower Pleistocene has long presented a challenge for taxonomic attribution of isolated postcrania. Although fossil humeri are well-suited for studies of hominin postcranial variation due to their relative abundance, humeral articular morphology has thus far been of limited value for differentiating Paranthropus from Homo. On the other hand, distal humeral diaphyseal shape has been used to justify such generic distinctions at Swartkrans. The potential utility of humeral diaphyseal shape merits larger-scale quantitative analysis, particularly as it permits the inclusion of fragmentary specimens lacking articular morphology. This study analyzes shape variation of the distal humeral diaphysis among fossil hominins (c. 2-1 Ma) to test the hypothesis that specimens can be divided into distinct morphotypes. Coordinate landmarks were placed on 3D laser scans to quantify cross-sectional shape at a standardized location of the humeral diaphysis (proximal to the olecranon fossa) for a variety of fossil hominins and extant hominids. The fossil sample includes specimens attributed to species based on associated craniodental remains. Mantel tests of matrix correlation were used to assess hypotheses about morphometric relationships among the fossils by comparing empirically-derived Procrustes distance matrices to hypothetical model matrices. Diaphyseal shape variation is consistent with the hypothesis of three distinct morphotypes (Paranthropus, Homo erectus, non-erectus early Homo) in both eastern and southern Africa during the observed time period. Specimens attributed to non-erectus early Homo are unique among hominids with respect to the degree of relative anteroposterior flattening, while H. erectus humeri exhibit morphology more similar to that of modern humans. In both geographic regions, Paranthropus is characterized by a morphology that is intermediate with respect to those morphological features that differentiate the two forms of early Homo. This study demonstrates the utility of the humeral diaphysis for taxonomic identification of isolated postcranial remains and further documents a high degree of postcranial diversity in early Homo.

Osteology and Cartilaginous Elements of Trilepida salgueiroi (Amaral, 1954) (Scolecophidia: Leptotyphlopidae).

Morphological and anatomical studies on the Leptotyphlopidae have increased in the past 10 years, providing important data on the systematics of this group, mainly focused on the skull and lower jaw morphology. However, most studies are based on a single specimen, rarely combining more than one single method of assessing morphological information. Therefore, several data on postcranial morphology, sexual dimorphism, and ontogenetic and intraspecific variation of leptotyphlopids remain poorly understood. Herein, we provide a detailed description of the cranial and postcranial skeleton of Trilepida salgueiroi based on more than 20 specimens (including males, females and juveniles), as well as a description of osteology and cartilaginous elements through the use of combined methodologies such as X-ray scanning, high resolution CT-scanning, and clearing and staining of articulated and disarticulated specimens. We also provide data on the presence/absence of dimorphic and intraspecific variation of the observed characters. The presence of a statolithic mass in the cavum vestibuli differs from the pattern found in other scolecophidians. A correlation of dorsal vertebrae with ventral and subcaudal scales was found (1:1), as well as total number of vertebrae with middorsal scales (1:1), thoracolumbar vertebrae and ventral scales (1:1), and a higher number of caudal vertebra with subcaudal scales (1.23:1). Intraspecific variation was found in several elements of the skull, lower jaw, pelvic girdle and vertebral number, but no evidence of sexual dimorphism was found in any of the species characters analysed. The homologies of several elements are discussed, although still remaining poorly understood and unknown.

Shape, size, and maturity trajectories of the human ilium.

Morphological traits of the ilium have consistently been more successful for juvenile sex determination than have techniques applied to other skeletal elements, however relatively little is known about the ontogeny and maturation of size and shape dimorphism in the ilium. We use a geometric morphometric approach to quantitatively separate the ontogeny of size and shape of the ilium, and analyze interpopulation differences in the onset, rate and patterning of sexual dimorphism. We captured the shape of three traits for a total of 191 ilia from Lisbon (Portugal) and London (UK) samples of known age and sex (0-17 years). Our results indicate that a) there is a clear dissociation between the ontogeny of size and shape in males and females, b) the ontogeny of size and shape are each defined by non-linear trajectories that differ between the sexes, c) there are interpopulation differences in ontogenetic shape trajectories, which point to population-specific patterning in the attainment of sexual dimorphism, and d) the rate of shape maturation and size maturation is typically higher for females than males. Male and female shape differences in the ilium are brought about by trajectory divergence. Differences in size and shape maturation between the sexes suggest that maturity may confound our ability to discriminate between the sexes by introducing variation not accounted for in age-based groupings. The accuracy of sex determination methods using the ilium may be improved by the use of different traits for particular age groups, to capture the ontogenetic development of shape in both sexes.

Evolution of the dinosauriform respiratory apparatus: new evidence from the postcranial axial skeleton.

Examination of the thoracic rib and vertebral anatomy of extant archosaurs indicates a relationship between the postcranial axial skeleton and pulmonary anatomy. Lung ventilation in extant crocodilians is primarily achieved with a hepatic piston pump and costal rotation. The tubercula and capitula of the ribs lie on the horizontal plane, forming a smooth thoracic "ceiling" facilitating movement of the viscera. Although the parietal pleura is anchored to the dorsal thoracic wall, the dorsal visceral pleura exhibits a greater freedom of movement. The air sac system and lungs of birds are associated with bicapitate ribs with a ventrally positioned capitular articulation, generating a rigid and furrowed rib cage that minimizes dorsoventral changes in volume in the dorsal thorax. The thin walled bronchi are kept from collapsing by fusion of the lung to the thorax on all sides. Data from this study suggest a progression from a dorsally rigid, heterogeneously partitioned, multichambered lung in basal dinosauriform archosaurs towards the small entirely rigid avian-style lung that was likely present in saurischian dinosaurs, consistent with a constant volume cavum pulmonale, thin walled parabronchi, and distinct air sacs. There is no vertebral evidence for a crocodilian hepatic piston pump in any of the taxa reviewed. The evidence for both a rigid lung and unidirectional airflow in dinosauriformes raises the possibility that these animals had a highly efficient lung relative to other Mesozoic vertebrates, which may have contributed to their successful radiation during this time period.