Morphology and Function of the Vertebral Column in Remingtonocetus domandaensis (Mammalia, Cetacea) from the Middle Eocene Domanda Formation of Pakistan

The archaeocete family Remingtonocetidae is a group of early cetaceans known from the Eocene of India and Pakistan. Previous studies of remingtonocetids focused primarily on cranial anatomy due to a paucity of well-preserved postcranial material. Here we describe the morphology of the known vertebral column in Remingtonocetus domandaensis based largely on a single well-preserved partial skeleton recovered from the upper Domanda Formation of Pakistan. The specimen preserves most of the precaudal vertebral column in articulation and includes seven complete cervical vertebrae, ten partial to complete thoracic vertebrae, six complete lumbar vertebrae, and the first three sacral vertebrae. Cervical centra are long and possess robust, imbricating transverse processes that stabilized the head and neck. Lumbar vertebrae allowed for limited flexibility and probably served primarily to stabilize the lumbar column during forceful retraction of the hind limbs. Vertebral evidence, taken together with pelvic and femoral morphology, is most consistent with interpretation of Remingtonocetus domandaensis as an animal that swam primarily by powerful movement of its hind limbs rather than dorsoventral undulation of its body axis.     

A nearly complete skeleton of an early juvenile diplodocid (Dinosauria: Sauropoda) from the Lower Morrison Formation (Late Jurassic) of north central Wyoming and its implications for early ontogeny and pneumaticity in sauropods

A nearly complete skeleton of a juvenile sauropod from the Lower Morrison Formation (Late Jurassic, Kimmeridgian) of the Howe Ranch in Bighorn County, Wyoming is described. The specimen consists of articulated mid-cervical to mid-caudal vertebrae and most appendicular bones, but cranial and mandibular elements are missing. The shoulder height is approximately 67 cm, and the total body length is estimated to be less than 200 cm. Besides the body size, the following morphological features indicate that this specimen is an early juvenile; (1) unfused centra and neural arches in presacral, sacral and first to ninth caudal vertebrae, (2) unfused coracoid and scapula, (3) open coracoid foramen, and (4) relatively smooth articular surfaces on the limb, wrist, and ankle bones. A large scapula, short neck and tail and elongate forelimb bones relative to overall body size demonstrate relative growth. A thin-section of the mid-shaft of a femur shows a lack of annual growth lines, indicating an early juvenile individual possibly younger than a few years old. Pneumatic structures in the vertebral column of the specimen SMA 0009 show that pneumatisation of the postcranial skeleton had already started in this individual, giving new insights in the early ontogenetic development of vertebral pneumaticity in sauropods.

The specimen exhibits a number of diplodocid features (e.g., very elongate slender scapular blade with a gradually dorsoventrally expanded distal end, a total of nine dorsal vertebrae, presence of the posterior centroparapophyseal lamina in the posterior dorsal vertebrae). Although a few diplodocid taxa, Diplodocus, cf. Apatosaurus, and cf. Barosaurus, are known from several fossil sites near the Howe Ranch, identification of this specimen, even at a generic level, is difficult due to a large degree of ontogenetic variation.     

Additional materials of Myanmarpithecus yarshensis (Amphipithecidae, Primates) from the middle Eocene Pondaung Formation

Myanmarpithecus yarshensis is an amphipithecid primate from the middle Eocene Pondaung Formation in Myanmar. It was previously known based on maxillary fragments with P⁴–M³ and mandibular fragments with C–P₃ and M_2–3. This study reports new materials for the genus, including a humeral head fragment, a lingual fragment of the right M², a lingual fragment of the right M³, and a left I¹. These new materials were collected from approximately the same point, and likely belonged to the same individual. The upper molar morphology and size of the new materials show similarity to those of the type specimen, indicating that the new materials can be assigned to M. yarshensis. The humeral head is the first postcranial element that is associated with dental materials for amphipithecids. The morphological similarity between the previously reported larger humerus and this specimen confirms the assignment of the former specimen to Amphipithecidae and suggests common locomotor adaptations in the family. The upper central incisor is large relative to the molar fragments, but is within the variation among extant platyrrhines. The tooth is spatulate-shaped and high crowned, and lacks the mesial process, indicating similarity to I¹ of haplorhines and clear differences from that of adapoids. It has been suggested that amphipithecids, including Myanmarpithecus, have affinities with notharctine adapoids, but the morphology of I¹ does not support the notharctine hypothesis of the Amphipithecidae.     

Cross-sectional morphology of the SK 82 and 97 proximal femora

Computed tomography scans of the proximal femoral shaft of the South African “robust” australopithecine, A. robustus, reveal a total morphological pattern that is similar to the specimen attributed to A. boisei in East Africa but unlike that of Homo erectus or modern human femora. Like femora attributed to H. erectus, SK 82 and 97 have very thick cortices, although they do not have the extreme increase in mediolateral buttressing that is so characteristic of H. erectus. And unlike H. erectus or modern humans, their femoral heads are very small relative to shaft strength. These features are consistent with both increased overall mechanical loading of the postcranial skeleton and a possibly slightly altered pattern of bipedal gait relative to that of H. erectus and modern humans. Am J Phys Anthropol 109:509–521, 1999. © 1999 Wiley-Liss, Inc.     

An oreopithecid proximal humerus from the middle miocene of Maboko Island,Kenya

A proximal humerus, recently recovered from the middle Miocene of Maboko Island, Kenya, provides the earliest evidence of postcranial structure and adaptation of Oreopithecidae. Provisionally attributed toNyanzapithecus pickfordi (Harrison, 1986), the specimen manifests a globose head, subequally large tuberosities, and a board, shallow bicipital groove. Although readily distinguished from the fundamentally cercopithecoid proximal humeral morphology ofVictoriapithecus (Senut, 1986), the Maboko Island oreopithecid, shows none of the derived features that are characteristic of the proximal humeri of extant hominoids. It is inferred from proximal humeral anatomy that the Maboko Island oreopithecid was an active arboreal scansor with moderate mobility at the shoulder but lacking adaptations for circumduction of the arm. In combination with craniodental evidence, proximal humeral morphology indicates that Oreopithecidae was a clade of hominoids which originated before the last common ancestor of extant apes and went extinct, without issue, in the later Miocene.     

Femoral neck structure and function in early hominins

All early (Pliocene–Early Pleistocene) hominins exhibit some differences in proximal femoral morphology from modern humans, including a long femoral neck and a low neck‐shaft angle. In addition, australopiths (Au. afarensis, Au. africanus, Au. boisei, Paranthropus boisei), but not early Homo, have an “anteroposteriorly compressed” femoral neck and a small femoral head relative to femoral shaft breadth. Superoinferior asymmetry of cortical bone in the femoral neck has been claimed to be human‐like in australopiths. In this study, we measured superior and inferior cortical thicknesses at the middle and base of the femoral neck using computed tomography in six Au. africanus and two P. robustus specimens. Cortical asymmetry in the fossils is closer overall to that of modern humans than to apes, although many values are intermediate between humans and apes, or even more ape‐like in the midneck. Comparisons of external femoral neck and head dimensions were carried out for a more comprehensive sample of South and East African australopiths (n = 17) and two early Homo specimens. These show that compared with modern humans, femoral neck superoinferior, but not anteroposterior breadth, is larger relative to femoral head breadth in australopiths, but not in early Homo. Both internal and external characteristics of the australopith femoral neck indicate adaptation to relatively increased superoinferior bending loads, compared with both modern humans and early Homo. These observations, and a relatively small femoral head, are consistent with a slightly altered gait pattern in australopiths, involving more lateral deviation of the body center of mass over the stance limb. Am J Phys Anthropol, 2013. © 2013 Wiley Periodicals, Inc.     

Functional morphology of the postcranium and locomotor behavior of Neosaimiri fieldsi,a Saimiri-like Middle Miocene platyrrhine

A number of postcranial specimens of Neosaimiri fieldsi, a Middle Miocene platyrrhine, were discovered in 1988, 1989, and 1990 at La Venta, Colombia. Until recently only three postcranial specimens of this species had been discovered and the present material adds further information about this taxon's postcranial morphology. In overall skeletal dimensions and in postcranial features, Neosaimiri is most similar to Saimiri among extant medium-sized platyrrhines, but differs from Saimiri in having more rugose surface markings, a longer olecranon, a smaller anterior process of the distal tibia, an absence of a distal surface extension on the anterior tibial shaft, an absence of an anterior midtrochlear depression of the talus, and a shorter distal calcaneus relative to the calcaneal tuberosity. These differences suggest that Neosaimiri was relatively heavily built, possessed a more dominant forelimb in quadrupedal progression, and utilized a less stabilized upper ankle joint, and a shorter power arm for plantarflexion. Neosaimiri is interpreted as an arboreal quadruped with frequent leaping across arboreal gaps, as in extant Saimiri, with perhaps less frequent running and leaping than in Saimiri. As with the dentition, the postcranial specimens suggest the close relationship between Neosaimiri and extant Saimiri. Am J Phys Anthropol 102:515–544, 1997. © 1997 Wiley-Liss, Inc.     

Skeletal anatomy of the extinct shark Paraorthacodus jurensis (Chondrichthyes; Palaeospinacidae), with comments on synechodontiform and palaeospinacid monophyly

The skeletal morphology of Paraorthacodus jurensis, a Late Jurassic neoselachian from Nusplingen, is described based on the incomplete holotype and a newly discovered almost complete specimen. For the first time, the postcranial skeleton could be investigated. Paraorthacodus is characterized by a monognath dental heterodonty and tearing‐type dentition. The number of lateral cusplets in the lateral teeth differs between the holotype and the new specimen, possibly indicating sexual dimorphism. Clasper organs are not preserved in either of the two specimens. The notochord is sheathed by about 123 well‐calcified vertebral centra. The posterior‐most caudal vertebrae are lacking. The transition from monospondylous thoracic to diplospondylous abdominal vertebrae occurs at centra 48 and 49. The origin of the caudal fin is at the 80th centrum. Most conspicuous is the presence of a single spineless dorsal fin. In this respect, Paraorthacodus differs from most palaeospinacids, but resembles Macrourogaleus. Palidiplospinax possibly is sister to a group comprising Synechodus, Paraorthacodus, and Macrourogaleus (the Palaeospinacidae). A reinterpretation of dental and skeletal characters of synechodontiform taxa indicates that Synechodontiformes and Palaeospinacidae are monophyletic groupings of basal neoselachians. Synechodontiformes is probably sister to all living elasmobranchs.     

A new basilosaurid (Cetacea, Pelagiceti) from the Late Eocene to Early Oligocene Otuma Formation of Peru

Cynthiacetus peruvianus nov. sp. is a new basilosaurid species, from Late Eocene to Early Oligocene Otuma Formation of Peru. It is the first described archaeocete in South-America and is represented by a sub-complete skeleton. C. peruvianus differs from C. maxwelli (middle to Late Eocene of Egypt and United States) principally in having one cuspid less on both mesial and distal sides of p3 and p4. Cynthiacetus is among the largest basilosaurids. Its more characteristic features are located on its postcranial skeleton: large vertebrarterial foramina on cervical vertebrae and absence of ventral expansion of the transverse process on C3-C5. Besides, C. peruvianus presents the greatest number of thoracic vertebrae (20) and ribs observed in Cetacea and the first thoracics have an almost vertical neural spine. A preliminary parsimony analysis establishes the monophyly of the Basilosauridae on the basis of three unambiguous cranial synapomorphies. However, within the Basilosauridae, the most diagnostic characters are observed on the postcranial skeleton.     

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.     

The Smallest Known Tetrapod Footprints: Batrachichnus Salamandroides from the Carboniferous of Joggins,Nova Scotia,Canada

A new specimen of Batrachichnus salamandroides from the classic Carboniferous section at Joggins, Nova Scotia, is the smallest set of tetrapod footprints known in the fossil record. The trackmaker was a juvenile, quadrupedal temnospondyl or microsaur with a trunk length of 3.55 mm and an estimated body length of 8 mm (skull, presacral vertebrae, and caudal vertebrae). The 48 mm-long trackway preserves a high degree of extramorphological variation along its course, including a gait change associated with a change in direction along with an increased stride and pace, and the appearance of overstepped imprints, in the latter portion of the trackway. These morphological changes suggest the tetrapod changed from a walking gait to a running gait.     

The first specimen of the Middle Triassic Phalarodon atavus (Ichthyosauria: Mixosauridae) from South China,showing postcranial anatomy and peri‐Tethyan distribution

Phalarodon atavus from the Germanic Muschelkalk Basin was previously represented only by cranial elements. Here we report a nearly complete and articulated specimen of P. atavus from the Middle Triassic Luoping Biota, Yunnan, South China. This is the first specimen of P. atavus from outside the Germanic Basin. This discovery demonstrates a peri‐Tethyan distribution of P. atavus. The new specimen is also the first one preserving the postcranial anatomy of this species, providing the opportunity to evaluate its sustained swimming ability. Inferences made on its functional morphology suggest that this species was probably adapted for active foraging. Tooth crown morphology suggests that P. atavus may have preferred externally soft prey.     

The petrosal and bony labyrinth of Diplobune minor,an enigmatic Artiodactyla from the Oligocene of Western Europe

Anoplotheriinae are Paleogene European artiodactyls that present a unique postcranial morphology with a tridactyl autopodium and uncommon limb orientation. This peculiar morphology led to various hypotheses regarding anoplotheriine locomotion from semiaquatic to partly arboreal or partly bipedal. The petrosal bone, housing the organs of balance, and hearing, offers complementary information to postcranial morphology on the ecology of this uncommon artiodactyl. Here, we investigate the middle ear and bony labyrinth of the small anoplotheriine Diplobune minor based on four specimens from the Early Oligocene locality of Itardies (Quercy, France). A macroscopic study coupled with a μCT scan investigation of the petrosal anatomy provides novel information on the bony labyrinth, stapes, and innervation and vasculature of the inner ear of this enigmatic taxon. The petrosal of D. minor exhibits a mosaic of plesiomorphic characters and peculiar features that shed new light into the anatomy of this poorly studied taxon of an obscure taxonomic clade. We can confidently reject that D. minor was a semiaquatic species based on the petrosal morphology: presence of a large mastoid process and nonpachyostotic tegmen tympani do not support underwater hearing. On the other hand, the average semicircular canal radius points to a slow or medium slow agility for D. minor , and fully rejects it was a fast moving animal, which is congruent with its postcranial anatomy.     

Evidence of Spondyloarthropathy in the Spine of a Phytosaur (Reptilia: Archosauriformes) from the Late Triassic of Halberstadt,Germany

Pathologies in the skeleton of phytosaurs, extinct archosauriform reptiles restricted to the Late Triassic, have only been rarely described. The only known postcranial pathologies of a phytosaur are two pairs of fused vertebrae of “Angistorhinopsis ruetimeyeri” from Halberstadt, Germany, as initially described by the paleontologist Friedrich von Huene. These pathologic vertebrae are redescribed in more detail in this study in the light of modern paleopathologic methods. Four different pathologic observations can be made in the vertebral column of this individual: 1) fusion of two thoracic vertebral bodies by new bone formation within the synovial membrane and articular capsule of the intervertebral joint; 2) fusion and conspicuous antero-posterior shortening of last presacral and first sacral vertebral bodies; 3) destruction and erosion of the anterior articular surface of the last presacral vertebra; and 4) a smooth depression on the ventral surface of the fused last presacral and first sacral vertebral bodies. Observations 1–3 can most plausibly and parsimoniously be attributed to one disease: spondyloarthropathy, an aseptic inflammatory process in which affected vertebrae show typical types of reactive new bone formation and erosion of subchondral bone. The kind of vertebral shortening present in the fused lumbosacral vertebrae suggests that the phytosaur acquired this disease in its early life. Observation 4, the smooth ventral depression in the fused lumbosacral vertebrae, is most probably not connected to the spondyloarthropathy, and can be regarded as a separate abnormality. It remains of uncertain origin, but may be the result of pressure, perhaps caused by a benign mass such as an aneurysm or cyst of unknown type. Reports of spondyloarthropathy in Paleozoic and Mesozoic reptiles are still exceptional, and our report of spondyloarthropathy in fossil material from Halberstadt is the first unequivocal occurrence of this disease in a Triassic tetrapod and in a phytosaur.     

Cross-Sectional Morphology of the Femoral Neck of Wild Chimpanzees

To understand the mechanical effects of different modes of locomotion on the femoral neck of chimpanzees, we investigated the cross-sectional morphology of the femoral neck of 4 chimpanzees (Pan troglodytes schweinfurthii) collected from the Mahale Mountains, Tanzania. We performed serial computed tomography (CT) scans of the neck from the femoral head to the base of the neck perpendicular to the long axis of the neck. We measured the cortical thickness of the serial 5 cross sections of the neck region every 45° around the circumference, i.e., 8 points per section, and examined the cross-sectional properties of the mid-section. When we compared the superior and inferior parts of the cortical thickness of the femoral neck, the inferior part exhibited the greatest cortical thickness whereas the superior part had the smallest values in every specimen. Researchers have also observed such regional differences between superior and inferior cortical thicknesses in bipedal humans and other primates, although these differences are not as large in the chimpanzee as in bipedal hominini. The present study differed from the past study on hominini and chimpanzees in that the superior anterior (SA) part exhibited greater cortical thickness in chimpanzees. We believe these observations reflect the structural strengthening of parts of the chimpanzee femoral neck that is needed to accommodate the mechanical loads imposed by arboreal vertical climbing and terrestrial quadrupedal and bipedal locomotion.     

A New Titanosaurian Sauropod from the Hekou Group (Lower Cretaceous) of the Lanzhou-Minhe Basin,Gansu Province,China

Increased excavation of dinosaurs from China over the last two decades has enriched the record of Asian titanosauriform sauropods. However, the relationships of these sauropods remain contentious, and hinges on a few well-preserved taxa, such as Euhelopus zdanskyi. Here we describe a new sauropod, Yongjinglong datangi gen. nov. et sp. nov., from the Lower Cretaceous Hekou Group in the Lanzhou Basin of Gansu Province, northwestern China. Yongjinglong datangi is characterized by the following unique combination of characters, including seven autapomorphies: long-crowned, spoon-shaped premaxillary tooth; axially elongate parapophyses on the cervical vertebra; very deep lateral pneumatic foramina on the lateral surfaces of the cervical and cranial dorsal vertebral centra; low, unbifurcated neural spine fused with the postzygapophyses to form a cranially-pointing, triangular plate in a middle dorsal vertebra; an “XI”-shaped configuration of the laminae on the arches of the middle dorsal vertebrae; a very long scapular blade with straight cranial and caudal edges; and a tall, deep groove on the lateral surface of the distal shaft of the radius. The new specimen shares several features with other sauropods: a pronounced M. triceps longus tubercle on the scapula and ventrolaterally elongated parapophyses in its cervical vertebra as in Euhelopodidae. Based on phylogenetic analyses Yongjinglong datangi is highly derived within Titanosauria, which suggests either a remarkable convergence with more basal titanosauriform sauropods in the Early Cretaceous or a retention of plesiomorphic features that were lost in other titanosaurians. The morphology and remarkable length of the scapulocoracoid reveal an unusual relationship between the shoulder and the middle trunk: the scapulocoracoid spans over half of the length of the trunk. The medial, notch-shaped coracoid foramen and the partially fused scapulocoracoid synostosis suggest that the specimen is a subadult individual. This specimen sheds new light on the diversity of Early Cretaceous Titanosauriformes in China.     

A New Sail-Backed Styracosternan (Dinosauria: Ornithopoda) from the Early Cretaceous of Morella,Spain

A new styracosternan ornithopod genus and species is here described based on a partial postcranial skeleton and an associated dentary tooth of a single specimen from the Arcillas de Morella Formation (Early Cretaceous, late Barremian) at the Morella locality, (Castellón, Spain). Morelladon beltrani gen. et sp. nov. is diagnosed by eight autapomorphic features. The set of autapomorphies includes: very elongated and vertical neural spines of the dorsal vertebrae, midline keel on ventral surface of the second to fourth sacral vertebrae restricted to the anterior half of the centrum, a posterodorsally inclined medial ridge on the postacetabular process of the ilium that meets its dorsal margin and distal end of the straight ischial shaft laterally expanded, among others. Phylogenetic analyses reveal that the new Iberian form is more closely related to its synchronic and sympatric contemporary European taxa Iguanodon bernissartensis and Mantellisaurus atherfieldensis, known from Western Europe, than to other Early Cretaceous Iberian styracosternans (Delapparentia turolensis and Proa valdearinnoensis). The recognition of Morelladon beltrani gen. et sp. nov. indicates that the Iberian Peninsula was home to a highly diverse medium to large bodied styracosternan assemblage during the Early Cretaceous.