Postembryonic ontogeny of the spadefoot toad,Scaphiopus intermontanus (Anura: Pelobatidae): Skeletal morphology

Postembryonic skeletal ontogeny of the pelobatid frog Scaphiopus intermontanus is described based on a developmental series of cleared-and-stained, whole-mount specimens. The focus is on laboratory-reared individuals fed a herbivorous diet as larvae. Although there is variation in the timing of ossification of individual skeletal elements relative to developmental stages based on external morphological criteria, the sequence of skeletal development generally is conservative. Compared with its close relative, S. bombifrons, ossifications that occur during prometamorphosis tend to be slightly delayed in S. intermontanus; however, cranial bones that ossify during late metamorphic climax in S. intermontanus are delayed until postmetamorphosis in S. bombifrons. The differences in timing between the two species are consistent, however, with differences observed between two developmental series of S. intermontanus raised at two different temperatures. Noteworthy features of skeletal development in S. intermontanus include: 1) presence of palatine ossifications that form from independent centers of ossification and soon fuse with the postnarial portion of the vomers to form the compound vomeropalatine bones; 2) compound sphenethmoid that may arise from four or more endochondral centers of ossification and one dorsal, dermal center of ossification; and 3) presence of transverse processes and vestigal prezygapophyses on the first postsacral vertebra. The morphology of the larval orbitohyoideus and interhyoideus muscles is compared. The record of skeletal ontogeny and muscle morphology presented herein for the herbivorous larval morph can serve as a baseline for comparisons with the ontogeny of the carnivorous larval morph of Scaphiopus. J. Morphol. 238:179–244, 1998. © 1998 Wiley-Liss, Inc.     

Ontogeny of the osteocranium in the African catfish,Clarias gariepinus Burchell (1822) (Siluriformes: Clariidae): Ossification sequence as a response to functional demands

The ontogeny of the bony skull of the African catfish, Clarias gariepinus, is studied from initial ossification until a complete skull is formed. The ossification sequence in C. gariepinus seems to be related to the functional demands that arise in a developing larva. Early ossification of the opercular bone coincides with the initiation of opercular skin movements. Early ossifications involve several dentulous bones, formed shortly before the transition phase from endogenous to exogenous feeding. The enlarging branchiostegal membrane becomes supported by the gradual adding of branchiostegal rays. Parasphenoid ossification may be related to protection of the brain during prey transport, whereas the several hyoid bones, including the parurohyal, are formed in relation to the increasing loads exerted onto the tendons of the sternohyoideus and consequently onto the hyoid bar. Overall skull reinforcement occurs almost simultaneously, with a whole set of perichondral bones arising especially at places of high mechanical load. The suspensorium becomes protected against dislocation in an anteroposterior direction through a ligamentous connection, which even becomes partially ossified, forming the sesamoid entopterygoid. Later, the cranial lateral-line system becomes enclosed by a set of gutters, which close, frequently becoming plate-like later in ontogeny. The brain also becomes covered dorsally. Additional dentition (prevomeral tooth plates) formation seems to coincide with formation of the opercular four-bar system, as well as with the time the digestive system becomes completely functional. Eventually, unossified regions between the bones become closed off, fortifying and completely covering the skull. J. Morphol. 235:183–237, 1998. © 1998 Wiley-Liss, Inc.     

Primary homologies of the circumorbital bones of snakes

Some snakes have two circumorbital ossifications that in the current literature are usually referred to as the postorbital and supraorbital. We review the arguments that have been proposed to justify this interpretation and provide counter‐arguments that reject those conjectures of primary homology based on the observation of 32 species of lizards and 81 species of snakes (both extant and fossil). We present similarity arguments, both topological and structural, for reinterpretation of the primary homologies of the dorsal and posterior orbital ossifications of snakes. Applying the test of similarity, we conclude that the posterior orbital ossification of snakes is topologically consistent as the homolog of the lacertilian jugal, and that the dorsal orbital ossification present in some snakes (e.g., pythons, Loxocemus, and Calabaria) is the homolog of the lacertilian postfrontal. We therefore propose that the terms postorbital and supraorbital should be abandoned as reference language for the circumorbital bones of snakes, and be replaced with the terms jugal and postfrontal, respectively. The primary homology claim for the snake “postorbital” fails the test of similarity, while the term “supraorbital” is an unnecessary and inaccurate application of the concept of a neomorphic ossification, for an element that passes the test of similarity as a postfrontal. This reinterpretation of the circumorbital bones of snakes is bound to have important repercussions for future phylogenetic analyses and consequently for our understanding of the origin and evolution of snakes. J. Morphol. 274:973–986, 2013. © 2013 Wiley Periodicals, Inc.     

The forgotten remains of a leptocleidid plesiosaur (Sauropterygia: Plesiosauroidea) from the Early Cretaceous of Gronau (Münsterland, Westphalia, Germany)

Gronausaurus wegneri n. gen. n. sp. represents a newly discovered leptocleidid sauropterygian based on one individual from the Early Cretaceous (Berriasian) of Gronau in Westphalia, Germany. The holotype and only known specimen consists of a skeleton, which lacks most of the dermal skull bones, a large number of cervical vertebrae and distal limb elements. Gronausaurus wegneri is unique in having distinct cavities, the subdiapophyseal fossae, below the transverse processes of the pectoral and anterior dorsal vertebrae, that probably stabilised the bones against tensile forces of the rotator and levator muscles in the living animal.     

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.     

Histochemical study of jaw muscle fibers in the American alligator (Alligator mississippiensis)

The ontogenetic development of caudal vertebrae and associated skeletal elements of salmonids provides information about sequence of ossification and origin of bones that can be considered as a model for other teleosts. The ossification of elements forming the caudal skeleton follows the same sequence, independent of size and age at first appearance. Dermal bones like principal caudal rays ossify earlier than chondral bones; among dermal bones, the middle principal caudal rays ossify before the ventral and dorsal ones. Among chondral bones, the ventral hypural 1 and parhypural ossify first, followed by hypural 2 and by the ventral spine of preural centrum 2. The ossification of the dorsal chondral elements starts later than that of ventral ones. Three elements participate in the formation of a caudal vertebra: paired basidorsal and basiventral arcocentra, chordacentrum, and autocentrum; appearance of cartilaginous arcocentra precedes that of the mineralized basiventral chordacentrum, and that of the perichordal ossification of the autocentrum. Each ural centrum is mainly formed by arcocentral and chordacentrum. The autocentrum is irregularly present or absent. Some ural centra are formed only by a chordacentrum. This pattern of vertebral formation characterizes basal teleosts and primitive extant teleosts such as elopomorphs, osteoglossomorphs, and salmonids. The diural caudal skeleton is redefined as having two independent ural chordacentra plus their arcocentra, or two ural chordacentra plus their autocentra and arococentra, or only two ural chordacentra. A polyural caudal skeleton is identified by more than two ural centra, variably formed as given for the diural condition. The two ural centra of primitive teleosts may result from early fusion of ural centra 1 and 2 and of ural centra 3 and 4, or 3, 4, and 5 (e.g., elopomorphs), respectively. The two centra may corespond to ural centrum 2 and 4 only (e.g., salmonids). Additionally, ural centra 1 and 3 may be lost during the evolution of teleosts. Additional ural centra form late in ontogeny in advanced salmonids, resulting in a secondary polyural caudal skeleton. The hypural, which is a haemal spine of a ural centrum, results by growth and ossification of a single basiventral ural arococentrum and its haemal spine. The proximal part of the hypural always includes part of the ventral ural arcocentrum. The uroneural is a modification of a ural neural arch, which is demonstrated by a cartilaginous precursor. The stegural of salmonids and esocids originates from only one paired cartilaginous dorsal arcocentrum that grows anteriorly by a perichondral basal ossification and an anterodorsal membranous ossification. The true epurals of teleosts are detached neural spines of preural and ural neural arches as shown by developmental series; they are homologous to the neural spines of anterior vertebrae. Free epurals without any indication of connection with the dorsal arococentra are considered herein as an advanced state of the epural. Caudal distal radials originate from the cartilaginous distal portion of neural and haemal spines of preural and ural (epurals and hypurals) vertebrae. Therefore, they result from distal growth of the cartilaginous spines and hypurals. Cartilaginous plates that support rays are the result of modifications of the plates of connective tissue at the posterior end of hypurals (e.g., between hypurals 2 and 3 in salmonids) and first preural haemal spines, or from the distal growth of cartilaginous spines (e.g., epural plates in Thymallus). Among salmonids, conditions of the caudal skeleton such as the progressive loss of cartilaginous portions of the arcocentra, the progressive fusion between the perichondral ossification of arcocentra and autocentra, the broadening of the neural spines, the enlargement and interdigitation of the stegural, and other features provide evidence that Prosopium and Thymallus are the most primitive, and that Oncorhynchus and Salmo are the most advanced salmonids respectively. This interpretation supports the current hypothesis of phylogenetic relationships of salmonids. © 1992 Wiley-Liss, Inc.     

Ontogeny of cranial ossification in the small-mouthed salamander, Ambystoma texanum (Matthes)

The ontogenetic sequence of cranial bony structure from initial ossifications through metamorphosis in Ambystoma texanum is described on the basis of 128 cleared and stained specimens. For convenience of discussion nine stages are recognized on the basis of conspicuous events. Cranial bones ossify and are modified in a definite sequence, and comparisons of complete sequences among groups of salamanders may prove useful in classification and in better understanding of relationships.     

Osteogenesis,homology, and function of the intercostal plates in ornithischian dinosaurs (Tetrapoda,Sauropsida)

Intercostal plates are bony structures positioned lateral to the anterior dorsal ribs in some ornithischian dinosaurs. Some propose these plates are homologous, or functionally analogous, with the uncinate processes of extant avian dinosaurs that assist in breathing, while others suggest they served a defensive function. To elucidate their osteogenesis, homology, and function, a histological survey of intercostal plates from three taxa (Hypsilophodon, Talenkauen, and Thescelosaurus) was undertaken. This study reveals that osteogenesis of intercostal plates closely resembles that of secondary centers of ossification in endochondral bone, typically present in the epiphyses of mammalian long bones. In contrast, ossification of avian uncinate processes begins at a primary ossification center via the development of a bony collar around a cartilaginous model. Based on these data, intercostal plates and avian uncinate processes are likely not evolutionary homologs. Dense packets of obliquely oriented Sharpey’s fibers within the parallel-fibered bone of somatically mature intercostal plates indicate these plates were positioned medial to at least a portion of the hypaxial musculature, which does not support their use as bony armor. Rather, we propose that intercostal plates performed some biomechanical function, either assisting in breathing in a way analogous to avian uncinate processes, or working together with the sternal ribs and sternal plates of these ornithischian taxa to provide increased rigidity to the anterior portion of the ribcage.     

Appendicular skeletal morphology in minute salamanders,genus Thorius (amphibia: Plethodontidae): Growth regulation,adult size determination,and natural variation

Patterns of growth and variation of the appendicular skeleton were examined in Thorius, a speciose genus of minute terrestrial plethodontid salamanders from southern Mexico. Observations were based primarily on ontogenetic series of each of five species that collectively span the range of adult body size in the genus; samples of adults of each of seven additional species provided supplemental estimates of the full range of variation of limb skeletal morphology. Limbs are generally reduced, i.e., pedomorphic, in both overall size and development, and they are characterized by a pattern of extreme variation in the composition of the limb skeleton, especially mesopodial elements, both within and between species. Fifteen different combinations of fused carpal or tarsal elements are variably present in the genus, producing at least 18 different overall carpal or tarsal arrangements, many of which occur in no other plethodontid genus. As many as four carpal or tarsal arrangements were observed in single population samples of each of several; five tarsal arrangements were observed in one population of T. minutissimus. Left-right asymmetry of mesopodial arrangement in a given specimen is also common. In contrast, several unique, nonpedomorphic features of the limb skeleton, including ossification of the typically cartilaginous adult mesopodial elements and ontogenetic increase in the degree of ossification of long bones, are characteristic of all species and distinguish Thorius from most related genera. They form part of a mechanism of determinate skeletal growth that restricts skeletal growth after sexual maturity. Interspecific differences in the timing of the processes of appendicular skeletal maturation relative to body size are well correlated with interspecific differences in mean adult size and size at sexual maturity, suggesting that shifts in the timing of skeletal maturation provide a mechanism of achieving adult size differentiation among species. Processes of skeletal maturation that confer determinate skeletal growth in Thorius are analogous to those typical of most amniotes – both groups exhibit ontogenetic reduction and eventual disappearance of the complex of stratified layers of proliferating and maturing cartilage in long bone epiphyses – but, unlike most amniotes, Thorius lacks secondary ossification centers. Thus, the presence of secondary ossification centers cannot be used as a criterion for establishing determinate skeletal growth in all vertebrates.     

Ontogeny of the Early Carboniferous acanthodian <Emphasis Type="Italic">Acanthodes lopatini</Emphasis> Rohon

The sequence of ossification of skeletal elements in the growth series of the acanthodian Acanthodes lopatini Rohon from the Lower Tournaisian of the Minusa Trough (Siberia) is described. Four formal ontogenetic stages are recognized. It is shown that even 20?30-mm-long juveniles have well-developed fin spines, scapulocoracoids, mandibular bones, branchiostegal rays, and incipient gill rakers. At this stage, scales cover the posterior part of the body and the orbits are marked by dark “eye stains” of uncertain nature. Further ontogenetic changes are connected with the development of sclerotic ring elements, sensory line system, squamation of the head region and fin webs, perichondral ossifications of the Meckel’s, quadrate, and palatine cartilages, and, finally, mineralization of the hyoid and gill arches. Ontogenetic features of A. lopatini are compared with those in other members of the genus and other acanthodiforms. Some developmental features shared by A. lopatini and Lodeacanthus gaujicus Upeniece suggest that acanthodids could have evolved from mesacanthids.     

Development of the integument of Dasypus hybridus and Chaetophractus vellerosus,and asynchronous events with respect to the postcranium

The integument of extant armadillos (Xenarthra, Cingulata) is a unique organ in which complex glandular systems are associated with pilose follicles, dermal ossifications, and cornified scales. Up to date, papers have focused on neither comparative morphology of the skin (dorsal and ventral) nor chronology of the development of interspecific homolog structures. In order to clarify the way in which events occur during development of the integument structures, maturity of other tissues (e.g. skeletal tissues) should be considered. Therefore, we will be able to identify events that have been pre- or post-displaced during ontogenetic development. The aim of this paper is to describe in a developmental and comparative framework the integumentary system of neonates of Dasypus hybridus and Chaetophractus vellerosus. In order to understand the morphology of the different integumentary structures serial histological sections were prepared. Staining techniques included H–E, Masson Trichrome, PAS, orcein and reticulin. To study ossification of postcranial elements, the specimens were cleared and double-stained with alcian blue and alizarin red. Determinations of ossification centers and their progress were recorded through the early uptake of alizarin. The dorsal dermis of neonates from D. hybridus is clearly differentiated into a superficial and deep layer, as in fetuses of Dasypus novemcinctus. In C. vellerosus, however, these layers could not be identified. This suggests a less connective tissue differentiation in the latter species at this stage. Osteoderms in D. hybridus are well differentiated unlike C. vellerosus where no condensations of osteoprogenitory cells are observed. Conversely, pilose follicles and glandular tissues are less developed in D. hybridus. Regarding postcranial elements, ossification centers are less advanced in C. vellerosus than D. hybridus, this is particularly notorious for the vertebral column, sternal, and pelvic girdle elements. Asynchronies between neonates of both species observed on integumentary and postcranial skeletal tissues could match with specific adaptive strategies related to distribution in different environments, and/or different postnatal care.     

The Rutland Cetiosaurus: the anatomy and relationships of a Middle Jurassic British sauropod dinosaur

A relatively well–preserved specimen of Cetiosaurus oxoniensis, from the Middle Jurassic (Bajocian) of Rutland, United Kingdom, is described in detail. The material includes a nearly complete cervical series, representative dorsal vertebrae, a fragment of sacrum, anterior caudals, the right femur, and numerous rib and limb fragments. Contrary to previous suggestions that this specimen possesses 14 cervical and ten dorsal vertebrae, it seems more probable that there were at most 13 cervicals and at least 12 dorsals. The vertebral column displays several autapomorphic features which supplement the generic diagnosis of Cetiosaurus, including: (1) a stout, anteriorly directed process located at the top of the neural spine of the twelfth (?) cervical vertebra; and (2) the presence of lateral pits, separated by a thin midline septum, below the transverse processes of middle dorsal vertebrae. Cladistic analysis indicates that Cetiosaurus is probably the sister–taxon to the advanced neosauropod clade. This relationship affects the distribution of particular character states that have played an important role in determining sauropod phylogeny.     

Senile features in the skeleton of an aged Japanese monkey

The senile features in the skeleton of a male Japanese monkey, who is presumed to be about 40 years old, were examined in comparison with younger individuals. As for the skull, every part is constructed solidly, and the sutures around the temporal and parietal bones are for the most part closed. In the dentition many of the front teeth are destroyed or lost, and the cheek teeth are severely worn. In the vertebrae, the annular epiphyseal discs unite completely with the body at its anterior and posterior surfaces, and the porosity and deformation of the bodies are remarkable. The hip bones, in the pelvis, unite with each other by solid ossification of the pubic symphysis. The long bones of the anterior and posterior limbs are marked by their general thickness, the rugged increase of bone around the joints, especially in the arms, and the complete union of each epiphysis with the shaft through the ossification of the epiphyseal cartilage. These senile features furnish a clue as to the establishment of a criterion for age estimation in Japanese monkeys.This observation was briefly reported inMonkey Vol. 13, No. 1, 1969, and Fig. 9 was used there.     

Development and diversity of the suspensorium of trichomycterids and comparison with loricarioids (Teleostei: Siluriformes)

Studies of ontogenetic series of trichomycterids and other catfishes reveal that the suspensorium of siluroids is highly specialized; several synapomorphies separate siluroids from other teleosts. In siluroids, the palatoquadrate is divided into pars autopalatina and pars pterygoquadrata and both are usually connected by the autopaiatine-metapterygoid ligament. The pterygoquadrate is broadly joined to the dorsal limb of the hyoid arch, forming a cartilaginous hyomandibular-symplectic-pterygoquadrate plate in early ontogeny. This produces a special alignment of the hyomandibula and quadrate which is characteristic of siluroids. A symplectic bone is absent. The interhyal is absent in trichomycterids and astroblepids. Dorsal and ventral limbs of the hyoid arch are connected by a ligament. A rudimentary interhyal and this ligament are present in primitive siluroids such as diplomystids and nematogenyids as well as loricariids. The metapterygoid arises as an anterior ossification of the pars pterygoquadrata in siluroids. The formation and position of the metapterygoid exhibit two patterns: (1) the metapterygoid develops as an ossification of a cartilaginous projection positioned between the future hyomandibula and quadrate in primitive catfishes (e.g., Diplomystes) as well as in Nematogenys, callichthyids, loricariids, and astroblepids; (2) the metapterygoid arises as an ossification of the cartilaginous projection (pterygoid process) positioned just above the articular facet of the quadrate for the lower jaw. An ossified anterior chondral pterygoid process of the complex quadrate is present in trichomycterids, whereas the process is absent (simple quadrate) in catfishes such as diplomystids, nematogenyids, callichthyids, and loricariids. The anterior membranous process of the quadrate of Astroblepus is non-homologous with the chondral pterygoid process of trichomycterids; both structures arose independently within the loricarioids. Despite topological relationships, the origin and development of bones reveal the presence of a chondral hyomandibula which develops a large meinbranous outgrowth during ontogeny and a chondral metapterygoid in trichomycterids. The presence of a compound hyomandibula + metapterygoid or a compound metapterygoid + ectopterygoid + entopterygoid have no developmental support in trichomycterines or other siluroids. The “entopterygoid” of Nematogenys and Diplomystes arises as an ossification of a ligament. The dermal entopterygoid of other ostariophysans and the “entopterygoid” are homologous. An ectopterygoid or tendon bone “ectopterygoid” is absent in loricarioids. The suspensorium is an important structural system which has significant evolutionary transformations which characterize loricarioid subgroups; however, no character, of the suspensorium supports the monophyly of the loricarioids.     

Static osteogenesis does not precede dynamic osteogenesis in periosteal ossification of Pleurodeles (Caudata,Amphibia) and Pogona (Squamata,Lepidosauria)

Two successive mechanisms have been described in perichondral ossification: (1) in static osteogenesis, mesenchymal cells differentiate into stationary osteoblasts oriented randomly, which differentiate into osteocytes in the same site; (2) in dynamic osteogenesis, mesenchymal cells differentiate into osteoblasts that are all oriented in the same direction and move back as they secrete collagen fibers. This study is aimed at testing the hypothesis that the ontogenetic sequence static then dynamic osteogenesis observed in the chicken and in the rabbit is homologous and was acquired by the last common ancestor of amniotes or at a more inclusive node. For this we analyze the developmental patterns of Pleurodeles (Caudata, Amphibia) and those of the lizard Pogona (Squamata, Lepidosauria). We processed Pleurodeles larvae and Pogona embryos, prepared thin and ultrathin sections of appendicular bones, and analyzed them using light and transmission electron microscopy. We show that static osteogenesis does not precede dynamic osteogenesis in periosteal ossification of Pleurodeles and Pogona . Therefore, the null hypothesis is rejected and according to the parsimony method the ontogenetic sequence observed in the chicken and in the rabbit are convergent. In Pleurodeles and Pogona dynamic osteogenesis occur without a previous rigid mineralized framework, whereas in the chicken and in the rabbit dynamic osteogenesis seems to take place over a mineralized support whether bone (in perichondral ossification) or calcified cartilage (in endochondral ossification). Interestingly, in typical dynamic osteogenesis, osteoblasts show an axis (basal nucleus—distal endoplasmic reticulum) perpendicular to the front of secreted unmineralized bone matrix, whereas in Pleurodeles and Pogona this axis is parallel to the bone matrix.     

New information on a juvenile sauropod specimen from the Morrison Formation and the reassessment of its systematic position

Abstract: Morphological changes in the ontogeny of sauropods are poorly known, making difficult to establish the systematic affinities of very young individuals. New information on an almost complete juvenile sauropod (SMA 0009) with an estimated total length of about 2 m is here presented. The specimen was described as a diplodocid owing to the presence of some putative synapomorphies of this group. However, recent further preparation revealed the absence of diplodocid characters and the presence of macronarian derived characters. To test the affinities of this specimen, a phylogenetic analysis was conducted. The strict consensus tree recovers the specimen as a basal titanosauriform, in an unresolved relation with Brachiosaurus and Giraffatitan. Nevertheless, a brachiosaurid assignment is here suggested in base of the widely accepted monophyly of this group (only recovered when SMA 0009 is placed within this group). Although the existence of a new taxon cannot be completely ruled out, the combination of derived and plesiomorphic characters in the specimen suggests its assignment to Brachiosaurus. Sixteen extra steps are needed to place this specimen within Diplodocidae. The high cost to place this specimen within this group is owing to the fact that several diplodocid characters are absent in SMA 0009, such as the absence of divided centroprezygapophyseal lamina in cervical vertebrae, procoelous anterior caudal centra, composed lateral lamina in anterior caudal vertebrae, elongated middle caudal vertebrae, short cervical ribs and caudolateral projection of distal condyle of metatarsal I. Finally, the systematic position reveals few major ontogenetic transformations. These affect the pneumatic structures (e.g. change from simple pleurocoels in the cervical vertebrae to complex pleurocoels and the development of lateral excavations in the dorsal vertebrae) but also include unrecorded transformations of the neural spine (e.g. the development of the spinodiapophyseal lamina, the widening of the neural spines in the dorsal vertebrae) and allometric growth in some limb bones.     

New material of Natchitochia jonesi and a comparison of the innominata and locomotor capabilities of Protocetidae

New material of Natchitochia from the Bartonian Archusa Marl Member is described here, including thoracic, lumbar, sacral, and caudal vertebrae, an innominate, proximal femur, and pedal? phalanx. The vertebrae and innominate are similar to those of Qaisracetus and Georgiacetus. The structure of the caudal vertebrae support previous observations that as sacral vertebrae disconnect from the sacrum, they become caudalized, developing hemal processes on the posteroventral margins of the bodies, reminiscent of chevron bones associated with true caudal vertebrae. The innominate of Natchitochia shares an elongate ilium and pubis with Qaisracetus and Georgiacetus, which differ from the innominata of the more apomorphic archaeocetes. Comparison of archaeocete innominata and sacra in a phylogenetic context indicates that the apomorphic sacrum composed of 4 vertebrae (Pakicetus, Ambulocetus, Rodhocetus, Maiacetus) was reduced to 3 (Qaisracetus) to 2 (Protocetus?, Natchitochia) to 0 (Georgiacetus, Basilosauridae), while the innominata remained robust, supporting a large hind limb until the origin of the Basilosauridae. In Georgiacetus, the innominate is large but detached from the vertebral column, preventing the use of the hind limb in terrestrial locomotion. More crownward cetaceans for which the innominate is known display greatly reduced innominata and hind limbs are disconnected from the vertebral column.     

On the Sacral Plexus and Sacral Vertebra of Lizards.

The authors mention that of late it has been recognized that, in any attempt to answer the question as to which vertebra of any lower animal answers to the first sacral vertebra of Man, the nervous no less than the osteological relations of the parts should be carefully investigated. And it has been considered that the nervous rather than the osteological relations should be deemed the more important: in fact it has been sometimes asserted that the nerves must be taken as the fixed points, and that the bones must rather have their homology decided by the nerves, than vice versa. Should it be possible to show that in any group of reptiles, both the nervous and osteological relatious of any vertebra constautly agree with the nervous and osteological relation of Man's first sacral vertebra, the homology between such two parts may well be taken as thereby established; but if either of these sets of relations exhibit discrepancy, then of course such homology cannot be considered satisfactorily determined. Nor can we justly set aside osteological in favour of nervous resemblances if it should turn out that the nerves themselves exhibit notable variations of conditions as we pass from one allied form to another–a fortiori if there should be variations in this respect even within the limits of a species. It might surely be anticipated that more or less variation would be found to exist inner‐vous as well as in skeletal structures; and in the event of such anticipations being justified, the determination of sacral homology must depend upon a comparison of the values of the conflicting claims of different degrees of resemblance in both the osseous and nervous systems–unless we prefer to consider the osteological sacrum and the nervous sacrum as two distinct structures, which may or may not completely coincide, and may or may not widely diverge. The authors afterwards discuss the opinions held by Professor Gegenbaur with regard to the pelvic relations in birds and some reptiles, also those of Professor Hoffmann concerning the lumbar and sacral plexuses of Batrachians and Reptiles. Then follows an account of dissections of the Chameleon (Cha‐mceleo vulgaris), the Green Lizard (Laeerta viridis), the common Teguexin (Teius teguexin), the Bearded Lizard (Grammatophora barbata), the Agama colonorum, the Tuberculated Lizard (Iguana tuberculata), and of the Monitor (M. arenaria). On these dissections are based some remarks on the general condition of the nervous and osseous structures of the sacral region in Lizards, according to their views and as compared with those held by G‐egenbaur and Hoffmann. To this succeed other chapters devoted to a consideration of the sacral region of Batraehians, to the sacral region of Mammals, and to the sacral region of Birds, each discussed in a similar spirit. Their generalizations to the foregoing may be thus summarized:– It appears, then, that in Lizards generally, the lumbar plexus may be formed by from two to three roots, aud that the most pre‐axial of these is here in advance of the fourth presacral nerve, while the most postaxial root is never more postaxial than the first presacral nerve. But Monitor and Ohamwleo present a slight exception in certain respects. In all the Eeptilia examined and enumerated by the authors, the transverse processes which abut against the ilium are wholly or in part parapophysial, and are in serial relation (serial liomo‐logues) with the capitular processes (or the capitular parts of the transverse processes) of the more preaxial vertebrae. The junction of the sacral vertebrae with the ilium is much postacetabular in Saurians; but in Crocodilia and Tortoises (some at least) it is about opposite the acetabulum. In Batrachians the transverse processes abutting against the ilium are parapophysial, but diapophysial in nature like those of Eeptiles. In Mammals as compared with Lizards, it would seem, with respect to nerves, that the first and second sacral vertebra? (say, for instance, of the Cat), answer very well to the two vertebrae with enlarged transverse processes of Lizards, while osteologically they of course also answer very well to them. There can be little doubt, however, that the first two sacral vertebrae of the Cat are to be considered homologous with the anterior human sacral vertebra¹; and therefore it would seem that the two ilium‐joining vertebrse of Lizards should be considered homologous with the anterior human sacral vertebrae. In Man, the Cat, and also in other Mammals down to the Echidna, the transverse processes abutting against the ilium are parapophysial, like those of Eeptiles and Batrachians. In all the Mammals examined by the authors, however, the junction of the sacral transverse processes with the ilia is preacetabular, although that junction is much less preacetabular in position in Man than it is in most Mammals. Altogether, from the osseous and nervous conditions evinced together in the groups hitherto referred to, the authors propose the following definition of a “Sacral Vertebra” in Mammals, Eeptiles, and Batrachians:–“ vertebra'ivithparapophysial transverse processes winch abut against the ilium, preaxial or post‐axial or opposite to the acetabulum, and having a root of the sciatic plexus coming forth either immediately preaxiad or postaxiad of it.” This definition will exclude from the sacrum, as not abutting against the ilium, of Man, the more posterior vertebrse called “ sacral” in anthropotomy. But in the lower mammals (even already in Apes) the number of so‐called “ sacral ” vertebrre augments more or less with age by the ankylosis of the sacral vertebras, so as not to render the extent of the “ sacrum ” very variable. It would surely be well, then, to distinguish the human sacral vertebra, like the ribs, into true and false, those being the true sacral vertebrae which abut against the ilium. In Birds the determination of the homological relations of the different parts of the postdorsal part of the spinal column is a matter of much difficulty. On the whole, and seeing on the one hand the manifest homology between the sacral vertebrae of Man and Lizards by the help of Crocodiles and Tortoises, and on the other hand the manifest homology between the sacral vertebrae of Lizards and the posterior parapophysial vertebras of most Birds, the authors think it better to regard the latter vertebras in Birds as alone truly sacral, and to regard such forms as Bwceros, Pica, and certain Parrots as differing from the rule of the Class in the suppression of their parapophysial processes, sm&Fregatta as differing from the same rule by the development of parapophyses in all the vertebras of this region. The sacral vertebra? in Birds may be defined, then, as “vertebrce having one of the more postaxial roots of the sciatic plexus coming forth either immediately preaxiad or postaxiad, and having parapophysial transverse processes abutting against the ilium, such vertebra being placed immediately postaxiad to vertebra which are devoid of such parapophyses, or else being the homologues of a vertebra so conditioned in most birds. By the combination of these two definitions, as given above (the one for Mammals, Eeptiles, and Batrachians, and the other for Birds), it seems to the authors that the sacral vertebras may be defined in all Vertebrata above Pishes which have pelvic limbs.     

The anatomy of the palatoquadrate in the Lower Triassic Proterosuchus fergusi (Reptilia,Archosauromorpha) and its morphological transformation within the archosauriform clade

The anatomy of the palatoquadrate ossifications of the Lower Triassic archosauromorph Proterosuchus fergusi from South Africa is described. It consists of two ossifications, the epipterygoid and the quadrate, which were joined by cartilage in life. The margins of the cartilage are clearly indicated by ridges and grooves on the dorsal surface of the pterygoid. The epipterygoid ossification consists of two structures: the anteroposteriorly expanded basal portion and, dorsally from it, an extending, slender, ascending process. From the anterior margin of the basal portion of the epipterygoid, a plate‐like structure, herein called the lamina epipterygoidea anteromedialis, extends anteromedially to form the anterolateral wall of the cavum epiptericum. Comparisons with the similarly constructed embryonal and adult epipterygoid components of Sphenodon punctatus show that the anteromedial lamina of the epipterygoid of P. fergusi is an additional component of the epipterygoid of this species and that this lamina is absent in the former species. However, a structure in a topologically similar position to the anteromedial lamina of the epipterygoid of P. fergusi is present in the palatoquadrate of Alligator mississippiensis. In the latter species, the structure is called the lamina palatoquadrati anterior; it ossifies in membrane and forms the dorsolateral cover of the huge trigeminal ganglion. It is hypothesized here that the anteromedial lamina of the epipterygoid of P. fergusi and the anterior lamina of the palatoquadrate of A. mississippiensis are most probably homologous structures and are present in both the basal and one of the crown taxa of the archosauromorph clade, respectively.     

Demineralization of the vertebral skeleton in Atlantic salmon Salmo salar L. during spawning migration

In Atlantic salmon, Salmo salar, the mineral rate of vertebrae in a given fish varies according to the position of the vertebra along the rachidian axis. Indeed, the mean rate goes from 49% in the anterior vertebrae and raises to 51% in post-truncal vertebrae. Although no significant difference in the mineral rate was noticed between males and females either in the lower river basin or after spawning, the mineral rate of vertebral bone decreased significantly (1–2%) during spawning migration. Vertebrae, like scales, are an important reservoir of calcium from which fasting salmon draws the minerals and organic materials necessary for the substantial remodeling of cranial bones in males and for sexual maturation. We hypothesize that mineral decrease in vertebrae may be the result of a halastasic demineralization of the vertebral tissues.