The suction mechanism of the pipid frog,Pipa pipa (Linnaeus, 1758)

Most suction‐feeding, aquatic vertebrates create suction by rapidly enlarging the oral cavity and pharynx. Forceful enlargement of the pharynx is powered by longitudinal muscles that retract skeletal elements of the hyoid, more caudal branchial arches, and, in many fish, the pectoral girdle. This arrangement was thought to characterize all suction‐feeding vertebrates. However, it does not exist in the permanently aquatic, tongueless Pipa pipa , an Amazonian frog that can catch fish. Correlating high‐speed (250 and 500 fps) video records with anatomical analysis and functional tests shows that fundamental features of tetrapod body design are altered to allow P. pipa to suction‐feed. In P. pipa , the hyoid apparatus is not connected to the skull and is enclosed by the pectoral girdle. The major retractor of the hyoid apparatus arises not from the pectoral girdle but from the femur, which lies largely within the soft tissue boundaries of the trunk. Retraction of the hyoid is coupled with expansion of the anterior trunk, which occurs when the hypertrophied ventral pectoral elements are depressed and the urostyle and sacral vertebra are protracted and slide forward on the pelvic girdle, thereby elongating the entire trunk. We suggest that a single, robust pair of muscles adduct the cleithra to depress the ventral pectoral elements with force, while modified tail muscles slide the axial skeleton cranially on the pelvic girdle. Combined hyoid retraction, axial protraction, and pectoral depression expand the buccopharyngeal cavity to a volume potentially equal to that of the entire resting body of the frog. Pipa may be the only tetrapod vertebrate clade that enlarges its entire trunk during suction‐feeding.     

Pectoral fin and girdle development in the basal actinopterygians Polyodon spathula and Acipenser transmontanus

The pectoral fins of Acipenseriformes possess endoskeletons with elements homologous to both the fin radials of teleosts and the limb bones of tetrapods. Here we present a study of pectoral fin development in the North American paddlefish, Polyodon spathula, and the white sturgeon, Acipenser transmontanus, which reveals that aspects of both teleost and tetrapod endoskeletal patterning mechanisms are present in Acipenseriformes. Those elements considered homologous to teleost radials, the propterygium and the mesopterygial radials, form via subdivision of an initially chondrogenic plate of mesenchymal cells called the endoskeletal disc. In Acipenseriformes, elements homologous to the sarcopterygian metapterygium develop separately from the endoskeletal disc as an outgrowth of the endoskeletal shoulder girdle that extends into the posterior margin of the finbud. As in tetrapods, the elongating metapterygium and the metapterygial radials form in a proximal to distal order as discrete condensations from initially nonchondrogenic mesenchyme. Patterns of variation seen in the Acipenseriform fin also correlate with putative homology: all variants from the "normal" fin bauplan involved the metapterygium and the metapterygial radials alone. The primary factor distinguishing Polyodon and Acipenser fin development from each other is the composition of the endoskeletal extracellular matrix. Proteoglycans (visualized with Alcian Blue) and Type II collagen (visualized by immunohistochemistry) are secreted in different places within the mesenchymal anlage of the fin elements and girdle and at different developmental times. Acipenseriform pectoral fins differ from the fins of teleosts in the relative contribution of the endoskeleton and dermal rays. The fins of Polyodon and Acipenser possess elaborate endoskeletons overlapped along their distal margins by dermal lepidotrichia. In contrast, teleost fins generally possess relatively small endoskeletal radials that articulate with the dermal fin skeleton terminally, with little or no proximodistal overlap.     

Anatomy and early development of the pectoral girdle,fin, and fin spine of sturgeons (Actinopterygii: Acipenseridae)

Acipenseriformes hold an important place in the evolutionary history of bony fishes. Given their phylogenetic position as extant basal Actinopterygii, it is generally held that a thorough understanding of their morphology will greatly contribute to the knowledge of the evolutionary history and the origin of diversity for the major osteichthyan clades. To this end, we examined comparative developmental series from the pectoral girdle in Acipenser fulvescens, A. medirostris, A. transmontanus, and Scaphirhynchus albus to document, describe, and compare ontogenetic and allometric differences in the pectoral girdle. We find, not surprisingly, broad congruence between taxa in the basic pattern of development of the dermal and chondral elements of the pectoral girdle. However, we also find clear differences in the details of structure and development among the species examined in the dermal elements, including the clavicle, cleithrum, supracleithrum, posttemporal, and pectoral‐fin spine. We also find differences in the internal fin elements such as the distal radials as well as in the number of fin rays and their association with the propterygium. Further, there are clear ontogenetic differences during development of the dermal and chondral elements in these species and allometric variation in the pectoral‐fin spine. The characters highlighted provide a suite of elements for further examination in studies of the phylogeny of sturgeons. Determining the distribution of these characters in other sturgeons may aid in further resolution of phylogenetic relationships, and these data highlight the role that ontogenetic and comparative developmental studies provide in systematics. J. Morphol. 276:241–260, 2015. © 2014 Wiley Periodicals, Inc.     

Comparative morphology of the pectoral fin spine of the Persian sturgeon Acipenser persicus,the Russian sturgeon Acipenser gueldenstaedtii,and the Starry sturgeon Acipenser stellatus in Iranian waters of the Caspian Sea

The morphological characteristics of the pectoral fin spine were compared in three species of sturgeon, the Persian sturgeon (Acipenser persicus), the Russian sturgeon (Acipenser gueldenstaedtii), and the Starry sturgeon (Acipenser stellatus), all sampled from the Caspian Sea. On the basis of morphological characters of the pectoral fin spine, 62.2% of the individuals were correctly classified into separate groups. The cluster analysis also divided the three species into two major subgroups. Acipenser persicus and A. gueldenstaedtii were grouped together, suggesting a similar evolutionary basis. Significant morphological heterogeneity in pectoral fin spine characteristics was observed among the three sturgeon species. Principal component analysis identified the largest differences were in the pectoral fin spine size and the angle between distal pectoral fin spine and the horizontal line (A°). The first and second principal components (PC1 and PC2) of all observations accounted for 64.19% and 14.33% of the total variation, respectively. The combination of all analyses showed the relevance of applying pectoral fin spine shape for interspecific distinction of the three species of sturgeons.     

Development of the mandibular, hyoid arch and gill arch skeleton in the Chinese barb Puntius semifasciolatus: comparisons of ossification sequences among Cypriniformes

The morphogenesis and sequence of ossification and chondrification of skeletal elements of the jaws, and hyoid arch and gill arches of Puntius semifasciolatus are described. These data provide a baseline for further studies and enable comparisons with other described cypriniforms. Some general patterns of ossification in the hyoid arch and branchial arches in cypriniforms were notable. First, the overall development is from anterior to posterior, with the exception of the fifth ceratobranchial bone, which ossifies first. Second, where ossification of iterated elements is sequential, it tends to proceed from posterior to anterior, even when more posterior chondrifications are the smallest in the series. Ossification of the ceratobranchial, epibranchial and pharyngobranchial bones tends to proceed from ventral to dorsal. The comparisons revealed small sets of skeletal elements whose ossification sequence appears to be relatively conserved across cyprinid cypriniforms. Several potentially key timing changes in the ossification sequence of the jaws, hyoid arch and gill arches were identified, such as the accelerated timing of ossification of the fifth ceratobranchial bone, which may be unique to cypriniforms.     

Development of the muscles associated with the mandibular and hyoid arches in the Siberian sturgeon,Acipenser baerii (Acipenseriformes: Acipenseridae)

The skeleton of the jaws and neurocranium of sturgeons (Acipenseridae) are connected only through the hyoid arch. This arrangement allows considerable protrusion and retraction of the jaws and is highly specialized among ray‐finned fishes (Actinopterygii). To better understand the unique morphology and the evolution of the jaw apparatus in Acipenseridae, we investigated the development of the muscles of the mandibular and hyoid arches of the Siberian sturgeon, Acipenser baerii. We used a combination of antibody staining and formalin‐induced fluorescence of tissues imaged with confocal microscopy and subsequent three‐dimensional reconstruction. These data were analyzed to address the identity of previously controversial and newly discovered muscle portions. Our results indicate that the anlagen of the muscles in A. baerii develop similarly to those of other actinopterygians, although they differ by not differentiating into distinct muscles. This is exemplified by the subpartitioning of the m. adductor mandibulae as well as the massive m. protractor hyomandibulae, for which we found a previously undescribed portion in each. The importance of paedomorphosis for the evolution of Acipenseriformes has been discussed before and our results indicate that the muscles of the mandibular and the hyoid may be another example for heterochronic evolution.     

A novel pharyngeal expansion mechanism in the yellow-spotted fanray, Platyrhina tangi (Elasmobranchii: Batoidea), with special reference to the function of the fifth ceratobranchial cartilage in batoids

Expansion of the ‘pharynx’ during breathing or capturing prey in fishes generally involves posteroventral retraction of the hyoid arch. However, the hyoid arch structure of batoid fishes (skates, rays, guitarfishes, and sawfishes) is unique, and how they expand the pharyngeal cavity is poorly understood. To investigate the mechanism of pharyngeal expansion during breathing in the yellow-spotted fanray, Platyrhina tangi, we conducted anatomical and kinematic investigations of the pharyngeal region. Our study revealed that the yellow-spotted fanray and sharks have different skeletal linkage systems for pharyngeal expansion. During pharyngeal expansion in the yellow-spotted fanray, the hyoid bar and branchial apparatus rotate ventrally around the hinge joint between the fifth ceratobranchial cartilage and the pectoral girdle. This pharyngeal expansion mechanism appears to be widespread among batoid fishes and is unique among cartilaginous fishes (sharks, batoids, and holocephalans). Batoid fishes possibly developed this pharyngeal expansion mechanism during early batoid evolution.     

Study of the pectoral girdle and fins of the Late Carboniferous sibyrhynchid iniopterygians (Vertebrata,Chondrichthyes, Iniopterygia) from Kansas and Oklahoma (USA) by means of microtomography,with comments on iniopterygian relationships

The latest works on iniopterygians question their monophyly when considering only the neurocranium of the two families (Sibyrhynchidae and Iniopterygidae), which have different conditions of preservation. Some of the synapomorphies of the Iniopterygia concern the pectoral girdle and fins. However, the anatomy of these different elements is still poorly known in this taxon. Here we describe in details three dimensionally preserved cartilages of the pectoral girdle and fins of the sibyrhynchid Iniopera sp. These structures have been extracted virtually from phosphatised nodules thanks to conventional and synchrotron microtomography, using absorption and phase contrast based techniques in the later case. The pectoral girdle of Iniopera sp. consists of three elements, which are, from dorsal to ventral, a paired suprascapular cartilage, a pair of robust scapulocoracoids and an unpaired intercoracoid cartilage. The scapular part of the scapulocoracoids is extremely reduced and the suprascapular cartilages link the scapulcoracoids to the rear of the neurocranium. These characters may be iniopterygian synapomorphies. Iniopterygians, stem and crown-holocephalans share a basipterygium that articulates with the pectoral girdle and bears an enlarged first pectoral fin radial. Posteriorly, the basipterygium articulates with either a well-defined metapterygium (in crown-holocephalans) or a metapterygial axis (in stem-holocephalans).     

Cephalic muscle development in the Australian lungfish,Neoceratodus forsteri

Lungfishes are the extant sister group of tetrapods. As such, they are important for the study of evolutionary processes involved in the water to land transition of vertebrates. The evolution of a true neck, that is, the complete separation of the pectoral girdle from the cranium, is one of the most intriguing morphological transitions known among vertebrates. Other salient changes involve new adaptations for terrestrial feeding, which involves both the cranium and its associated musculature. Historically, the cranium has been extensively investigated, but the development of the cranial muscles much less so. Here, we present a detailed study of cephalic muscle development in the Australian lungfish, Neoceratodus forsteri, which is considered to be the sister taxon to all other extant lungfishes. Neoceratodus shows several developmental patterns previously described in other taxa; the tendency of muscles to develop from anterior to posterior, from their region of origin toward insertion, and from lateral to ventral/medial (outside‐in), at least in the branchial arches. The m.protractor pectoralis appears to develop as an extension of the most posterior m.levatores arcuum branchialium, supporting the hypothesis that the m.cucullaris and its derivatives (protractor pectoralis, levatores arcuum branchialium) are branchial muscles. We present a new hypothesis regarding the homology of the ventral branchial arch muscles (subarcualis recti and obliqui, transversi ventrales) in lungfishes and amphibians. Moreover, the morphology and development of the cephalic muscles confirms that extant lungfishes are neotenic and have been strongly influenced via paedomorphosis during their evolutionary history.     

Developmental structure of the vertebral column,fins, scutes and scales in bester sturgeon,a hybrid of beluga Huso huso and sterlet Acipenser ruthenus

In the larval bester, a hybrid sturgeon of beluga Huso huso and sterlet Acipenser ruthenus, development of cartilage around the notochord began 7 days post hatch (dph) (14·0 mm, total length, L_T). The vertebral cartilage develops in the following sequence: basidorsals and basiventrals, neural canals, neural spines and ribs. The development of ribs remained incomplete in the largest specimen (181 dph, 179 mm L_T) that was examined. Endoskeletal development of the fins began 4 dph for the dorsal and anal fins, 6 dph for the pectoral fin and 10 dph for the caudal and pelvic fins. Complete elements of all fins were observed by 91 dph and complete ossification of fin rays was observed by 122 dph in the double‐stained specimens. Observation of the histological sections, however, suggested that ossification occurred soon after the formation of the organic matrix in the fin rays. Dorsal scutes were first visible by 25 dph, followed by the lateral and ventral scutes, which were visible by 37 and 44 dph, respectively. The number of scutes was fixed at 44, 59 and 91 dph and ossification was complete by 59 (dorsal) and 91 dph (lateral and ventral scutes) in the double‐stained specimens. Ossification occurred soon after the formation of the scute organic matrix in the histological sections. Four types of scales were observed in the H. huso×A. ruthenus hybrid. Median predorsal, preanal and small scales on the anterior section of the head were visible by 59 dph. Scales on the caudal fin were visible by 91 dph and a variable assemblage of scales anterior to the anal fin was visible by 122 dph. Both the scutes and scales developed in a process that is similar to that of intramembranous ossification.     

The pectoral anatomy of selected Ostariophysi. II. The Cypriniformes and siluriformes

The pectoral myology and osteology of the cyprinoids Notemigonus crysoleucas, the golden shiner, and Catostomus commersonnii, the common white sucker, resemble those of generalized, lower teleosts in structure and function, except in features related to the manipulation of the massive fifth ceratobranchial of cyprinoids by muscles attaching on the girdle. Catostomus is more specialized in having unique intercostal muscles to the girdle, complex subclavian arteries and lack of a superficial trapezius muscle. The bony pectoral anatomy of the siluriform, Ictalurus nebulosus, the brown bullhead, is highly specialized in relation to the presence and locking of the massive pectoral spine which is formed of fused dorsal and ventral propterygial rays; there is consolidation of the girdle through fusion of bones, presence of unique stabilizing bony structures, firm symphyseal union of bilateral girdles and the presence of friction-surfaces of girdle and spine for locking. The movements of the spine are specialized in the greater guidance offered by the girdle. Myological specializations are related mainly to ventral appendicular muscles which lock the spine. The nervous and arterial systems are generalized.     

Comparative anatomy and development of pectoral and pelvic girdles in hylid anurans

The development of the tetrapod pectoral and pelvic girdles is intimately linked to the proximal segments of the fore‐ and hindlimbs. Most studies on girdles are osteological and provide little information about soft elements such as muscles and tendons. Moreover, there are few comparative developmental studies. Comparative data gleaned from cleared‐and‐stained whole mounts and serial histological sections of 10 species of hylid frogs are presented here. Adult skeletal morphology, along with bones, muscles, and connective tissue of both girdles and their association with the proximal portions of the anuran fore‐ and hindlimbs are described. The data suggest that any similarity could be attributable to the constraints of their ball‐and‐socket joints, including incorporation of the girdle and stylopodium into a single developmental module. An ancestral state reconstruction of key structures and developmental episodes reveals that several development events occur at similar stages in different species, thereby preventing heterochronic changes. The medial contact of the halves of the pectoral girdle coincides with the emergence of the forelimbs from the branchial chamber and with the total differentiation of the linkage between the axial skeleton and the girdles. The data suggest that morphogenic activity in the anterior dorsal body region is greater than in the posterior one, reflecting the evolutionary sequence of the development of the two girdles in ancient tetrapods. The data also document the profound differences in the anatomy and development of the pectoral and pelvic girdles, supporting the proposal that the pectoral and pelvic girdles are not serially homologous, as was long presumed.     

THE ORIGIN OF MORPHOLOGICAL CHARACTERS AND THE BIOLOGICAL BASIS OF HOMOLOGY

A homolog is a part of the phenotype that is homologous to equivalent parts in other species. A biological homology concept is expected to explain three properties of homologs: 1) the conservation of those features that are used to define a homolog, 2) the individualization of the homolog with regard to the rest of the body, and 3) the uniqueness of homologs, i.e., their specificity for monophyletic groups. The main obstacle to describing a mechanistic basis for homology is the variability of the developmental pathways of undoubtedly homologous characters. However, not all aspects of the developmental pathway are of equal importance. The only organizational features of the developmental system that matter are those that have been historically acquired and cause developmental constraints on the further evolutionary modification of the characters. Two main factors contribute to historically acquired developmental constraints: generative rules of pattern formation and ontogenetic networks. In particular, hierarchical and cyclical inductive networks have the required properties to explain homology. How common such networks are is an open empirical question. The development and variation of pectoral fin hooks in blenniid fishes is presented as a model for the study of a simple ontogenetic network.     

Characterization of vitellogenin from white sturgeon,Acipenser transmontanus

Sturgeon are an ancient family (Acipenseridae) of fishes that lie close to the divergence of fish that eventually evolved into terrestrial animals and those that evolved into modern teleost species. Therefore, white sturgeon vitellogenin sequences fill a gap in the current understanding of the functional domains of this protein family. Vitellogenin cDNA was sequenced and used to investigate gene expression in white sturgeon, Acipenser transmontanus. Estrogen-induced vitellogenin mRNA was detected in the livers of both males and females and was also detected in undifferentiated gonads of estrogen-treated fish. Low levels of vitellogenin mRNA were also detected in the testis of both control and estrogen-treated males. The cDNA encoded a 186-kDa protein that was missing only six to seven of the amino-terminal amino acids. Comparisons to silver lamprey, Xenopus, and chicken vitellogenin sequences indicate that the overall structure of the yolk protein domains were highly conserved. There was considerable homology in three regions of the lipovitellin I domain. These conserved sequences are likely to be involved in vitellogenin receptor binding. The phosvitin domain of white sturgeon vitellogenin contained fewer and shorter serine repeats as predicted from yolk protein phosphate content of fish compared to Xenopus and chicken. However, the vitellogenin of white sturgeon had a lower serine content as compared with silver lamprey, indicating that an increased serine content is not strictly a function of evolutionary age. Correspondence to: C. A. Bidwell     

新型有原始肢突胴甲鱼的发现及胴甲鱼早期演化的初步探讨

本文记述的曲靖始突鱼(Procondylolepis qujingensis gen.et sp.nov.)是近几年在云南曲靖早泥盆世地层中发现的有肢突胴甲鱼一原始类型。它和已知胴甲鱼(包括早泥盆世无肢突的和中、晚泥盆世有“盔”状肢突的)不同的最大特点是在其肩带与胸鳍相接的肩关节处有原始的肢突和简单的关节窝;胸鳍甲近端的关节区很小。它展现出胴甲鱼类这一高度特化、长期使人迷惑不解的肢突,在胴甲鱼演化史上发展变化的梗概,填补了肢突从无到有中间的缺环,使人了解到胸鳍的具体结构。文中主要根据肢突的有无和特化程度等,对胴甲鱼早期演化史作了初步探讨,将胴甲鱼分为无肢突超目(Abrachicondylia)和有肢突超目(Brachicondylia)两大部分。始突鱼则代表有肢突超目一早期成员。