Skeletal development of the direct-developing caecilian <Emphasis Type="Italic">Gegeneophis ramaswamii</Emphasis> (Amphibia: Gymnophiona: Caeciliidae)

A few previous studies of skeletal and especially skull development in Gymnophiona often provided contradictory results. We studied the development of the skull and vertebral column of Gegeneophis ramaswamii, a direct-developing Indian caeciliid, based on 13 specimens. The chondrocranium forms at (Brauer in Zool Jahrb Anat 12:477-508,1899) stage 38. First dermal and perichondral ossifications occur at stage 40. The first dermal bones to form are the mentomeckelian, dentary, angular, vomer, and premaxillary. These are followed by the coronoid, palatine, pterygoid, maxillary, and the skull-roofing bones. The last occurring dermal ossifications are the parasphenoid and the squamosal. We present evidence for the occurrence of a lacrimal bone. No ectopterygoid, basioccipital, supraoccipital, pleurosphenoid, postorbital, or supratemporal elements were found. We assess the homology of the bones constituting the caecilian skull and discuss the above-mentioned terminologies. The phylogenetic implications of our findings are briefly discussed and we conclude that the evidence from developmental morphology is at present consistent with a monophyletic Lissamphibia of temnospondyl origin.     

Cranial ontogeny in the direct-developing frog, Eleutherodactylus coqui (Anura: Leptodactylidae), analyzed using whole-mount immunohistochemistry.

Direct development in amphibians is an evolutionarily derived life-history mode that involves the loss of the free-living, aquatic larval stage. We examined embryos of the direct-developing anuran Eleutherodactylus coqui (Leptodactylidae) to evaluate how the biphasic pattern of cranial ontogeny of metamorphosing species has been modified in the evolution of direct development in this lineage. We employed whole-mount immunohistochemistry using a monoclonal antibody against the extracellular matrix component Type II collagen, which allows visualization of the morphology of cartilages earlier and more effectively than traditional histological procedures; these latter procedures were also used where appropriate. This represents the first time that initial chondrogenic stages of cranial development of any vertebrate have been depicted in whole-mounts. Many cranial cartilages typical of larval anurans, e.g., suprarostrals, cornua trabeculae, never form in Eleutherodactylus coqui. Consequently, many regions of the skull assume an adult, or postmetamorphic, morphology from the inception of their development. Other components, e.g., the lower jaw, jaw suspensorium, and the hyobranchial skeleton, initially assume a mid-metamorphic configuration, which is subsequently remodeled before hatching. Thirteen of the adult complement of 17 bones form in the embryo, beginning with two bones of the jaw and jaw suspensorium, the angulosplenial and squamosal. Precocious ossification of these and other jaw elements is an evolutionarily derived feature not found in metamorphosing anurans, but shared with some direct-developing caecilians. Thus, in Eleutherodactylus cranial development involves both recapitulation and repatterning of the ancestral metamorphic ontogeny.(ABSTRACT TRUNCATED AT 250 WORDS)     

五种蝌蚪口器及舌鳃骨的结构比较

采用体视显微镜和骨骼双染色法对5种不同栖息环境的无尾两栖动物蝌蚪的口器和舌鳃骨的形态结构特征进行了观察。5种蝌蚪口器由唇齿行、唇乳突和角质颌等组成。舌鳃骨是由关联骨Ⅰ、关联骨Ⅱ、角舌骨、舌鳃骨盘和角鳃骨等骨骼组成。蝌蚪的梅氏软骨若较发达,其摄食方式可能为刮食;蝌蚪的舌鳃骨发达,其摄食方式则可能为滤食。角质颌、唇齿以及角鳃骨上鳃耙的出现显著增强了蝌蚪主动摄食能力和对食物与非食物的主动选择性。     

Cranial ontogeny in Philautus silus (Anura: Ranidae: Rhacophorinae) reveals few similarities with other direct-developing anurans

Direct development has evolved in rhacophorine frogs independently from other anuran lineages, thereby offering an opportunity to assess features associated with this derived life history. Using a developmental series of the direct-developing Philautus silus (Ranidae: Rhacophorinae) from Sri Lanka, we examine features of cranial morphology that are part of a suite of adaptations that facilitate feeding in free-living tadpoles, but have been changed or lost in other direct-developing lineages. Larval-specific upper jaw cartilages, which are absent from many non-rhacophorine direct-developing species (such as Eleutherodactylus coqui), develop in embryos of P. silus. Similarly, lower jaw cartilages initially assume a larval morphology, which is subsequently remodeled into the adult jaw configuration before hatching. However, the cartilaginous jaw suspension and hyobranchial skeleton never assume a typical larval morphology. The palatoquadrate, which suspends the lower jaw, lacks the posterior connections to the braincase found in many metamorphosing species. Unlike in metamorphosing species, bone formation in P. silus begins before hatching. However, the sequence of bone formation resembles that of metamorphosing anurans more than that of other direct developers. In particular, P. silus does not exhibit precocious ossification of the lower jaw, which is characteristic of some frogs and caecilians that lack a free-living tadpole. These data reveal some similarities between Philautus and other direct-developing anurans. However, the departure of Philautus embryos from the generalized tadpole skeletal morphology is less pronounced than that observed in other direct-developing taxa.     

The retro-articular process, streptostyly and the caecilian jaw closing system

Caecilians have two functionally separate sets of jaw closing muscles. The jaw adductor muscles are parallel fibered muscles positioned close to the jaw joint and their lever mechanics suggests they are well suited to rapidly closing the jaws. A second set of muscles, the hypaxial interhyoideus posterior (IHP), levers the jaws closed by pulling on the retroarticular process (RA) of the lower jaw. Models of the lower jaw point out that the angle and length of the RA has a profound effect on the closure force exerted by the IHP. The caecilian skull is streptostylic – the quadrate-squamosal apparatus (QSA) moves relative to the rest of the skull, a condition that seems at odds with a well-ossified cranium. Modeling the contribution of this streptostylic suspension of the lower jaw shows that rotational freedom of the QSA amplifies the force of the IHP by redirecting force applied along the low axis of the lower jaw. Measurements from several species and life stages of preserved caecilians reveal a large variation in predicted bite force (as a multiple of IHP force) with age and phylogeny.     

The morphology of prehatching embryos of Caecilia orientalis (Amphibia: Gymnophiona: Caeciliidae)

The state of development of advanced embryos of the direct‐developing Ecuadorian caecilian Caecilia orientalis (Caeciliidae: Gymnophiona: Amphibia) was examined. Because it is established that development is correlated with reproductive modes in a number of features, we included comparison with taxa that represent the major reproductive modes and all of the modern normal tables and ossification sequences. The embryos of C. orientalis most closely resemble those of stage 47/48 Gegeneophis ramaswamii, an Indian caeciliid, and stage 47/48 Hypogeophis rostratus, a Seychellian caeciliid, both direct developers, in details of bone mineralization, chondrocranial degeneration, and vertebrogenesis. They are most like stage 45 H. rostratus in external features (gills, pigmentation, etc.). They are less similar to prehatchings of Ichthyophis kohtaoensis, an ichthyophiid with free‐living larvae, and to fetuses of the viviparous caeciliid Dermophis mexicanus and the viviparous typhlonectid Typhlonectes compressicauda at comparable total lengths in both skeletal development and external features. The similarity of developmental features among the direct‐developers suggests a correlation with mode of life history. A noteworthy feature is that C. orientalis has an armature of multiple rows of teeth on the lower jaw with tooth crowns that resemble the “fetal” teeth of viviparous taxa and that are covered with a layer of oral mucosal epithelium until full development and eruption, but the upper jaw bears a single row of widely spaced, elongate, slightly recurved teeth that resemble those of the adult. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.     

Metamorphosis of cranial design in tiger salamanders (Ambystoma tigrinum): A morphometric analysis of ontogenetic change

While ontogenetic analyses of skull development have contributed to our understanding of phylogenetic patterns in vertebrates, there are few studies of taxa that undergo a relatively discrete and rapid change in morphology during development (metamorphosis). Morphological changes occurring in the head at metamorphosis in tiger salamanders (Ambystoma tigrinum) were quantified by a morphometric analysis of cranial osteology and myology to document patterns of change during metamorphosis. We employed a cross-sectional analysis using a sample of larvae just prior to metamorphosis and a sample of transformed individuals just after metamorphosis, as well as larvae undergoing metamorphosis. There were no differences in external size of the head among the larval and transformed samples. The hyobranchial apparatus showed many dramatic changes at metamorphosis, including shortening of ceratobranchial 1 and the basibranchial. The subarcualis rectus muscle increased greatly in length at metamorphosis, as did hypobranchial length and internasal distance. A truss analysis of dorsal skull shape showed that at metamorphosis the snout becomes wider, the maxillary and squamosal triangles rotate posteromedially, and the neurocranium shortens (while maintaining its width), resulting in an overall decrease in skull length at metamorphosis. These morphometric differences are interpreted in light of recent data on the functional morphology of feeding in salamanders. Morphological reorganization of the hyobranchial apparatus and shape changes in the skull are related to the acquisition of a novel terrestrial feeding mode (tongue projection) at metamorphosis. Metamorphic changes (both internal and external) that can be used to judge metamorphic condition are discussed.     

Ontogenetic differences in the feeding biomechanics of oviparous and viviparous caecilians (Lissamphibia: Gymnophiona)

Caecilians have a unique dual jaw-closing system in that jaw closure is driven by the ancestral jaw-closing muscles (mm. levatores mandibulae) plus a secondarily recruited hyobranchial muscle (m. interhyoideus posterior). There is a variety of feeding habits (suction feeding, skin feeding, intrauterine scraping, and biting) during ontogeny that relate to reproductive modes in different caecilian species. This study examines the cranial biomechanics of caecilians in the suction-feeding larva of Ichthyophis cf. kohtaoensis, in the embryo and juvenile of the skin-feeding Boulengerula taitana, and in a newborn of the intrauterine feeder Typhlonectes natans. A lever arm model was applied to calculate effective mechanical advantages of jaw-closing muscles over gape angles and to predict total bite force in developing caecilians. In I. cf. kohtaoensis, Notable differences were found in the larval jaw-closing system compared to that of the adult. The suction-feeding larva of I. cf. kohtaoensis has comparatively large mm. levatores mandibulae that insert with an acute muscle fiber angle to the lower jaw and a m. interhyoideus posterior that has its optimal leverage at small gape angles. Conversely, the skin-feeding juvenile of B. taitana and the neonate T. natans are very similar in the feeding parameters considered herein compared to adult caecilians. Some ontogenetic variation in the feeding system of B. taitana before the onset of feeding was present. This study contributes to our understanding of the functional demands that feeding habits put on the development of cranial structures.     

The plagiosaurid temnospondyl Plagiosuchus pustuliferus (Amphibia: Temnospondyli) from the Middle Triassic of Germany: anatomy and functional morphology of the skull

The cranial anatomy of the plagiosaurid temnospondyl Plagiosuchus pustuliferus, from the Middle Triassic of Germany, is described in detail on the basis of a newly discovered skull and mandibular material. The highly derived skull is characterized by huge orbitotemporal fenestrae, a reduction of the circumorbital bones – the prefrontal, postfrontal and (probably) postorbital are lost – and the expansion of the jugal to occupy most of the lateral skull margin. Ventrally the extremely long subtemporal vacuities correlate with the elongate adductor fossa of the mandible. The dentition is feebly developed on both skull and mandible. Ossified ?ceratobranchials and ‘branchial denticles’ indicate the presence of open gills clefts in life. The remarkably divergent cranial morphology of P. pustuliferus highlights the extraordinary cranial diversity within the Plagiosauridae, probably unsurpassed within the Temnospondyli. Specific structural aspects of the skull – including an extremely short marginal tooth row, feeble dentition and an elongated chamber for adductor musculature – together with evidence for a hyobranchial skeleton, suggests that P. pustuliferus utilized directed suction feeding for prey capture. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 348–373.     

Morphology and metamorphosis of Eupsophus calcaratus tadpoles (Anura: Leptodactylidae)

Eupsophus calcaratus, a leptodactyloid frog from the austral Andean forests of Argentina and Chile, has endotrophic, nidicolous tadpoles. We studied a metamorphic series from Stages 31 to 46 of Gosner's developmental table (1960). Other than the scarce pigmentation, proportionately large eyes, and massive developing hindlimbs, the remaining external characters are similar to those of generalized, exotrophic larvae. At the same time, internal morphology does not reveal any character state attributable to the endotrophic-nidicolous way of life; conversely, structures such as the hyobranchial skeleton and the mandibular cartilages are similar to those of exotrophic-macrophagous tadpoles. The metamorphic process is characterized by the delayed development of diverse structures (e.g., ethmoid region, palatoquadrate, and hyobranchial apparatus), and the retention of some larval characters (e.g., parietal fenestrae, overall absence of ossification) with the absence of development of some "juvenile" characters (e.g., adult otic process, several bones) in metamorphosed individuals. These heterochronic processes and truncation of larval development are related to a shorter larval life (when compared to other species of the austral Andean region) and to the small size at metamorphosis.     

Evolution and mechanics of long jaws in butterflyfishes (family Chaetodontidae)

We analyzed the functional morphology and evolution of the long jaws found in several butterflyfishes. We used a conservative reanalysis of an existing morphological dataset to generate a phylogeny that guided our selection of seven short- and long-jawed taxa in which to investigate the functional anatomy of the head and jaws: Chaetodon xanthurus, Prognathodes falcifer (formerly Chaetodon falcifer), Chelmon rostratus, Heniochus acuminatus, Johnrandallia nigrirostris, Forcipiger flavissimus, and F. longirostris. We used manipulations of fresh, preserved, and cleared and stained specimens to develop mechanical diagrams of how the jaws might be protruded or depressed. Species differed based on the number of joints within the suspensorium. We used high-speed video analysis of five of the seven species (C. xanthurus, Chel. rostratus, H. acuminatus, F. flavissimus, and F. longirostris) to test our predictions based on the mechanical diagrams: two suspensorial joints should facilitate purely anteriorly directed protrusion of the lower jaw, one joint should allow less anterior protrusion and result in more depression of the lower jaw, and no joints in the suspensorium should constrain the lower jaw to simple ventral rotation around the jaw joint, as seen in generalized perciform fishes. We found that the longest-jawed species, F. longirostris, was able to protrude its jaws in a predominantly anterior direction and further than any other species. This was achieved with little input from cranial elevation, the principal input for other known lower jaw protruders, and is hypothesized to be facilitated by separate modifications to the sternohyoideus mechanism and to the adductor arcus palatini muscle. In F. longirostris the adductor arcus palatini muscle has fibers oriented anteroposteriorly rather than medial-laterally, as seen in most other perciforms and in the other butterflyfish studied. These fibers are oriented such that they could rotate the ventral portion of the quadrate anteriorly, thus projecting the lower jaw anteriorly. The intermediate species lack modification of the adductor arcus palatini and do not protrude their jaws as far (in the case of F. flavissimus) or in a purely anterior fashion (in the case of Chel. rostratus). The short-jawed species both exhibit only ventral rotation of the lower jaw, despite the fact that H. acuminatus is closely related to Forcipiger.     

Morphology and function of the head in foetal and juvenile Scolecomorphus kirkii (Amphibia: Gymnophiona: Scolecomorphidae)

The external and musculoskeletal morphology of the head is described for an ontogenetic series of the scolecomorphid caecilian Scolecomorphus kirkii . The rostral region of foetuses and juveniles is expanded into large, posterolaterally pointing paraoral processes that are formed by the maxilla. Extraoral teeth are present on the underside of the rostrum and laterally on the paraoral processes. In the foetuses, teeth are covered by epidermal tissue. The endoskeletal part of the foetal skull is largely cartilaginous, but all of the dermal bones, with the exception of the squamosal, are present. The foetal chondrocranium is extensively developed and shows a peculiar, posterolateral process of the nasal capsule that is connected to the trabecula cranii by a transverse bar posterior to the choana, and extends further posterior beyond the level of the posterior end of the pila antotica. Only two mm. adductor mandibulae are present, together with two pterygoideus muscles that insert onto the lower jaw. The palatoquadrate and quadrate of foetuses and juveniles, respectively, are highly mobile. It is suggested that the derived head morphology of Scolecomorphus foetuses and juveniles is an adaptation to specialized postparitive feeding.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 491–504.     

A NEW GENUS OF TAPIRIDAE FROM SHANWANG, SHANDONG

<正> While helping to curate the jumbled specimens accumulated at the Shanwang Paleontological Museum in 1985, the senior authors of the present paper were happy enough to find a tapirid specimen. Through careful preparation it turned out to be a part of articulated skeleton consisting of skull, lower jaw, cervical and some anterior dorsal vertebrae and a scapula. As most of the other fossils from Shanwang, it is laterally compressed, therefore, no width could actually be measured. The nasal bones of the skull have become vertically oriented through compression. Otherwise the skull is little deformed. In order to study the tooth morphology closely, we had to extract most of the teeth from the left side of the tightly occlused tooth battery. All this notwithstanding, the skull represents one of the few well preserved specimens among the tapirid fossils ever found.     

Tongue evolution in the lungless salamanders, family plethodontidae. I. Introduction, theory and a general model of dynamics.

Plethodontid salamanders capture prey by projecting the tongue from the mouth. An analysis of theoretical mechanics of the hyobranchial skeleton is used to formulate a working hypothesis of tongue movements. Predictions that the skeletal elements of the tongue are included in the projectile and that the hyobranchial skeleton is folded during projection are central to the analysis. When decapitated in a particular way, salamanders project the tongue, and it is not retracted. When these heads are fixed and sectioned, examination confirms the predications. In turn, these observations are used to refine the working hypothesis and to generate a general model of tongue dynamics for plethodontids. Muscles performing the major roles of projection (subarcualis rectus I) and retraction (rectus cervicis profundus) are identified. The skeleton is folded passively along a morphological track having the form of a tractrix. Predictions concerning the shape of the track and the exact configuration of the folded skeleton are confirmed by study of sectioned material. The skeleton unfolds along the track during retraction and is spread into the resting state. The model developed herein will be used as a basis for predictions concerning selection patterns in the family and for analytical purposes in comparative and evolutionary studies.     

Pecularities of bony skeleton development in Asian clawed salamanders (Onychodactylus, Hynobiidae) related to embryonization

We studied skull, vertebral column, and limb skeleton development in Japanese clawed salamander Onychodactylus japonicus (Hynobiidae). The study is based on the ontogenetic series of embryos and larvae obtained from wild-captured adults by artificial induction of breeding using hormonal stimulation. The first stages of the skeleton formation in O. japonicus are shifted to the late embryonic period and hatching larvae already possess a well-ossified vertebral column, large number of skull ossifications and show signs of ossification in the forelimb skeleton. Compared to the primitive pattern of the skeleton development typical for other hynobiid salamanders, O. japonicus shows a number of heterochronies related to embryonization. In particular, this species is characterized by an earlier ossification of the vertebral column compared to that of the skull and by the delayed development and early reduction of the coronoid. Our results, along with the previously reported data on the skeleton development in the Fischer’s clawed salamander O. fischeri (Smirnov and Vassilieva, 2002), indicate that the genus Onychodactylus is characterized by the loss or reduction of several skeletal features typically found at early larval stages in other Hynobiidae species. In particular, provisional bones (especially the coronoid) and their dentition are underdeveloped. In addition, it is corroborated that the first tooth generation is absent in Onychodactylus, whereas such monocuspid nonpedicellate tooth generation normally develops at the early larval stages of other caudate amphibians. Since similar patterns of skeleton ontogeny are observed in other caudate groups with different extent of embryonization, it is proposed that, in different lineages of Urodela, the evolution of ontogeny followed similar pathways and was accompanied by the same changes in skeletogenesis.     

Morphology of a fossil elephant calf (Archidiskodon,Elephantidae) from the Oldowan Muhkai IIa site

The skull and lower jaw morphology of a calf of Archidiskodon sp. from the Oldowan (Early Paleolithic) Muhkai IIa site (Akushinskii raion, Dagestan) is described. The Muhkai IIa site is dated more than 1.5 Ma. This is the first record of the skull and lower jaw of calf of this species from the northern Caucasus. A skull fragment and lower jaw with functioning teeth of the DP2/DP3 generation are preserved. The calf is at most 8–10 months of individual age. The finely plicate enamel and formation of a complete enamel loop on DP3 are evidence that the calf belongs to Archidiskodon rather than to the European Elephas lineage.     

A NEW PARASEMIONOTID-LIKE FISH FROM THE LOWER TRIASSIC OF JURONG, JIANGSU PROVINCE, SOUTH CHINA

Abstract:  Based on a well-preserved specimen from the Early Triassic Lower Qinglong Formation exposed at Qingshan Quarry, Jurong, Jiangsu Province of China, the new taxon Peia jurongensis gen. et sp. nov. is named and described with anatomical details of its dermal skull, neurocranium and postcranial skeleton. The new taxon is characterized by the following set of characters: shallow V-shaped, tuber-like rostral; anterior part of frontal widened to roughly same width as its posterior part; roughly squared parietal; supraorbital absent; wide and slightly posteriorly inclined preopercle; five small ossifications present in dorsal half of preopercle; preopercular canal shifting anteriorly in ventral half of the preopercle, and reaching the anteroventral corner of this bone; dermosphenotic joining the skull roof and with a descending lamina; numerous branchiostegal rays. Comparisons are made between the new taxon and several parasemionotids from the same locality; a discussion is carried out on the distribution and evolution of several characters in halecomorphs and other closely related neopterygians.