Comparative microanatomy of the branchial sieve in three sympatric cyprinid species,related to filter-feeding mechanisms

The chimaeroid holocephalian fishes are distinguished among extant chondrichthyans by the possession of three pairs of tooth plates, evergrowing and partially hypermineralized, that are not shed and replaced like the teeth of living elasmobranchs. Although derivation of the chimaeroid tooth plate from the fusion of members of a plesiomorphic chondrichthyan tooth family has been proposed, evidence for this hypothesis has been lacking. A new analysis of the development and structure of the tooth plates in Callorhinchus milii (Holocephali, Chimaeriformes) reveals the compound nature of the tooth plates in a chimaeroid fish. Each tooth plate consists of an oral and aboral territory that form independently in the embryo and maintain separate growth surfaces through life. The descending lamina on the aboral surface of the tooth plate demarcates the growth surface of the aboral territory. Comparison with the tooth plates of Chimaera monstrosa indicates that compound tooth plates may be a feature of all chimaeroids in which a descending lamina is present. The tooth plates in these fishes represent the fusion of two members of a reduced tooth family. The condition of the tooth plates in C. milii is plesiomorphic for chimaeroids and is of evolutionary significance in that it provides further evidence to support a lyodont dentition in chimaeroid fishes similar to that found in other chondrichthyans. © 1994 Wiley-Liss, Inc.     

The postorbital palatoquadrate articulation in elasmobranchs

Although modern hexanchiforms are the only extant elasmobranchs with a postorbital articulation, according to most morphological and molecular cladistic analyses they are not basal, suggesting that Huxley ( 1876 Proc Zool Soc 1876;24–59) correctly identified this articulation as “an altogether secondary connection.” A postorbital articulation is present in many Paleozoic sharks, but differs from that found in hexanchiforms in its morphology, topographic position on the braincase, and inferred ontogenetic origins. Furthermore, a postorbital articulation is absent in hybodonts (the putative extinct sister group to neoselachians). It is proposed that the term amphistylic should be restricted to the modern hexanchiform condition, where the articular facet is located on the primary postorbital process. An identical articulation probably existed in some extinct galeomorphs (e.g., ?Synechodus dubrisiensis, ?Paraorthacodus), but is not widespread within elasmobranchs generally. The term archaeostylic (“ancient pillar”) is proposed here for the suspensorial arrangement in Paleozic sharks with a postorbital articulation on the ventrolateral part of the lateral commissure. Such an articulation is not known in other gnathostomes and may represent a basal chondrichthyan synapomophy (especially if ?Pucapampella is a stem chondrichthyan), suggesting that the autodiastylic pattern is not primitive for chondrichthyans and that holocephalans have secondarily lost a postorbital articulation. The amphistylic condition may have arisen from the archaeostylic, or it could have been acquired independently within neoselachians, but in either case it is most parsimoniously viewed as apomorphic. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

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.     

Interpretation of the toothplates of chimaeroid fishes

It has been argued that the toothplates of chimaeroid fishes exhibit a mode of growth that is fundamentally different from that of other chondrichthyans. Chimaeroid toothplates are supposed to be statodont, growing from the basal surface, whereas other chondrichthyan dentitions are lyodont, growing from the lingual towards the labial surface of the jaw. That idea is shown to be mistaken, because chimaeroid toothplates grow from the lingual surface, like other chondrichthyan dentitions. The mistake resulted from confusion about the nomenclature of toothplate surfaces, and on the choice of Chimaera as a Recent model. Callorhynchus is a more appropriate model, since it is shown to exhibit a primitive toothplate conformation, with the labial and symphysial margins of the occlusal surface bounded by a descending lamina which is applied to the margin of the jaw cartilage and grows basally throughout life. The descending lamina is well developed in toothplates of the extinct chimaeroid genera Ischyodus, Pachymylus and Brachymylus, but is much reduced in all Recent genera other than Callorhynchus. A basally-growing descending lamina also bounds the labial and symphysial margins of the principal toothplates in the Mesozoic myriacanthoids and Squaloraja. The toothplates of the Palaeozoic ‘cochliodonts' are reviewed; amongst them, the chondrenchelyids are the only forms with a basally growing descending lamina. So far as the dentition and its mode of growth arc concerned, the closest Palaeozoic relatives of chimaeroids seem to be the chondrenchelyids. The only statodont (basally growing) toothplates found in the course of this work are those of ptyctodont placoderms, which are therefore unlikely to be related to any chondrichthyans. Statodonty in its original sense (failure to shed teeth) is shown to be widespread and possibly primitive in chondrichthyans. Cochliodont and chimaeroid toothplates grow in a logarithmic spiral. Toothplates of primitive chimaeroid type, with basally growing marginal descending laminae, can develop only when the constant angle of the spiral is small (less than about 35°), and when the oral surface of the jaw grows to the same logarithmic spiral.     

Functional morphology of the feeding system in the ruff — Gymnocephalus cernua (L. 1758) — (Teleostei,Percidae)

The ruff, Gymnocephalus cernua, is a European freshwater fish that feeds by sucking up small invertebrates from the bottom of ponds and slow flowing rivers. The feeding movements have been studied by simultaneous electromyography of seventeen muscles of the head and cinematographic techniques. A theoretical model of movements imposes the functional demands of suction upon an abstraction of the form of a teleost head. Three phases in the feeding act, a preparatory phase, a suction phase and a transport phase, could be correlated with the observed movements and EMGs. Differences between the predicted and the actual movement are discussed. Two different types of feeding occur. The direction, magnitude and duration of the suction forces during feeding are modified, according to the position of the prey. A mechanism preventing early mandibular depression allows sudden and strong suction. Retardation of the suspensorial abduction during the overall expansion of the buccal cavity is ascribed to kinetic interrelations with the hyoid arch. Protrusion of the upper jaws also permits an earlier closure of the mouth and directs the food-containing waterflow posteriorly. When the fish is feeding on sinking prey, protrusion occurs later in the sequence of movements than when it is feeding from the bottom. As the protruded jaws produce a downwardly pointed mouth this retardation aims the suction force.     

Development of the cranium ofNeoceratodus forsteri,with a discussion of the suspensorium and the opercular apparatus in Dipnoi

Summary The ontogeny of the cranium of Dipnoi is restudied. The investigation especially refers to the basic components of the dipnoan cranium and several functional and developmental aspects of the structure of the larval skull ofNeoceratodus. There are fundamental differences even in the early development and composition of the chondrocranium ofNeoceratodus and Lepidosirenidae. This result, and comparison with several osteichthyans and Tetrapoda, requires a reinterpretation of the components of the dipnoan skull base. The pterygoid processes are not reduced, but incorporated into the cranial base early in ontogeny. The characteristic elongate trabecular rods, which in Gnathostomata usually bridge the ethmoidal plate and the orbito-temporal base of the chondrocranium, are much delayed in development inNeoceratodus, or even seem absent inLepidosiren andProtopterus. Accordingly, in Dipnoi no typical basitrabecular junction is formed in early ontogeny. Instead, the pars quadrata is fused to the mesodermal basis cranii posteriorly. InNeoceratodus a mesially directed basal process of the palatoquadrate is recognizable, which topographically corresponds to the basal process of Urodela and the pseudobasal process of anuran larvae. The ethmosphenoid region of the dipnoan skull also develops quite differently. In Lepidosirenidae, the palatoquadrates are interconnected anteriorly by a distinct commissura palatoquadrati, whereas inNeoceratodus a continuous planum ethmoidale (trabecular plate) is formed. The primary embryonic quadrato-trabecular connection persists as a commissura quadratocranialis anterior below the foramen opticum, at the root of the ectethmoid process. The formation of the skull base in living Amphibia appears to provide the best model for comparison, though it is difficult to propose any undisputable shared derived character states of the cranium of Dipnoi and Tetrapoda beyond this similarity. A similar difficulty presents the phylogenetic interpretation of the hyoid arch. In contrast to the absence of any dorsal hyoid arch elements inLepidosiren, the small hyomandibula ofNeoceratodus is surprisingly complete. In larvae it consists of a laterohyale, an epihyal part, and a processus symplecticus. A stylohyal cartilage is present, which forms rather late in ontogeny. The major chondral components of the hyoid arch are thus comparable to those of living Actinopterygii, except that a distinct symplecticum is not separated off, the components are relatively smaller, and they do not ossify. In view of the early-immobilized palatoquadrate, the hyomandibula ofNeoceratodus has no suspensorial function, but represents part of an opercular hinge and opening mechanism. The hamuloquadrate knob at the posterior face of the quadrate body is comparable to the processus hyoideus in some Urodela. It provides a pivoting joint for the ceratohyale, and therefore functions in buccal expansion. The closed spiracular canals include mechanoreceptive lateral line organs, which probably represent proprioreceptive organs for adjustment of mandibular, hyoid, and opercular movements. It is concluded that considerable differences between the skull architecture of Dipnoi and other Osteognathostomata (Teleostomi) can be assigned to the fact that palatoquadrate and trabecular anlagen fail to separate, resulting in a dramatic and highly adaptive change of palatoquadrate development in early ontogeny. Though these differences include several characters that suggest a plagiostomate condition of the jaw apparatus, this can be explained as a secondary acquisition. The multitude of retained plesiomorphies observed in the cranium of Dipnoi do not exclude a sister group-relationship to Tetrapoda. However, the ancestral osteognathostome suspensorial pattern still presents a problem of interpretation, for we lack a detailed survey of the development and significance of different quadrato-neurocranial connections.     

Embryonic staging and external features of development of the Chimaeroid fish,Callorhinchus milii (Holocephali,Callorhinchidae)

The development of Callorhinchus milii, a primitive chondrichthyan fish (Subclass Holocephali) is described in detail based on a complete series of embryos from stage 17 to hatching. The external features of these specimens, in comparison with other chondrichthyan embryos, are used to establish the first staging table for any chimaeroid species. Each stage of C. milii is defined by a suite of morphological characters in addition to total length, including the number of somites, extent of external pigmentation, eye size and shape, head flexure, heart morphology, and size and shape of paired and unpaired fins. Particular attention is given to features of the gill arches and associated structures, including external gill filaments and the opercular flap. Embryos of this species also possess a transient rostral bulb, a feature unique to chimaeroids. Embryological development of Callorhinchus milii is similar to that previously described for sharks and batoids (Subclass Elasmobranchii), including the spiny dogfish, Squalus acanthias, the Japanese bullshark, Heterodontus japonicus, the lesser spotted dogfish, Scyliorhinus canicula, the frill shark, Chlamydoselachus anguineus, the guitarfish, Rhinobatus halavi, and the skate, Raja brachyura. Callorhinchus milii is also similar in overall development to another holocephalan, Hydrolagus colliei. A review of previous staging schemes confirms that early morphological development in all three major chondrichthyan lineages (sharks, batoids, and chimaeras) can be correlated using a common set of stages. A uniform staging system is provided that should prove useful in continuing ontogenetic and phylogenetic studies of this entire clade of fishes. J. Morphol. 236:25–47, 1998. © 1998 Wiley-Liss, Inc.     

Feeding mechanics of teleost fishes (Labridae; Perciformes): A test of four-bar linkage models

The feeding mechanisms of two labrid fishes (Cheilinus chlorurus and C. diagrammus: Labridae: Perciformes) are modeled using four-bar linkage theory from mechanical engineering. The actions of the feeding mechanisms are simulated by a computer program that uses morphometric data to calculate the geometry of mechanism structure. The predictions of three different four-bar linkages regarding the kinematics of feeding are compared to the movements observed through hign speed (200 fps) cinematography. A previously unidentified four-bar chain was found to be an accurate model of the mechanism by which upper jaw protrusion, maxillary rotation, and gape increase occur in Cheilinus. This mechanism involves the anterior jaws including the mandible, maxilla, premaxilla, palatine, and suspensorium. The accuracy of two previously described four-bar linkages was also tested by comparison of model predictions and film results. The opercular linkage proposed by Anker ('74) as a mechanism of jaw depression via opercular levation was found to be a poor predictor of feeding movements. This four-bar chain involves the opercle, suspensorium, interopercle, and mandible. Muller ('87) proposed a mechanism of hyoid depression involving cranial elevation due to epaxial muscle contraction as input motion The links in this mechanism include the neurocranium and hyomandibula, hyoid, sternohyoideus muscle, and pectoral girdle. This model was an accurate predictor of hyoid depression in Cheilinus when simultaneous cranial elevation and sternohyoideus contraction were simulated. Quantitative kinematic models involve simplifying assumptions when applied to complex musculoskeletal systems, but such models have a wide range of applications to vertebrate functional morphology.     

Sharks, Rays and A Chimaeroid from the Kimmeridgian (Late Jurassic) of Ringstead, Southern England

Sampling of a lenticular concentration of vertebrate debris and associated sediments from the lower Kimmeridgian of southern England has allowed the study of a diverse and abundant assemblage of chondrichthyan remains. A number of previously undescribed species are recorded, of which three new species are named; Squatina? frequens, Synechodus plicatus and Protospinax planus. Additional diagnosis of the genus Paracestracion Koken is given to allow its identification from dental remains. Several nominal batoid species are synonymised with Spathobatis bugesiacus Thiolliere. This assemblage is considered to be typical of Middle–Late Jurassic neritic environments, and is compared to other contemporaneous selachian faunas.     

Vertebral Pneumaticity in the Ornithomimosaur Archaeornithomimus (Dinosauria: Theropoda) Revealed by Computed Tomography Imaging and Reappraisal of Axial Pneumaticity in Ornithomimosauria

Among extant vertebrates, pneumatization of postcranial bones is unique to birds, with few known exceptions in other groups. Through reduction in bone mass, this feature is thought to benefit flight capacity in modern birds, but its prevalence in non-avian dinosaurs of variable sizes has generated competing hypotheses on the initial adaptive significance of postcranial pneumaticity. To better understand the evolutionary history of postcranial pneumaticity, studies have surveyed its distribution among non-avian dinosaurs. Nevertheless, the degree of pneumaticity in the basal coelurosaurian group Ornithomimosauria remains poorly known, despite their potential to greatly enhance our understanding of the early evolution of pneumatic bones along the lineage leading to birds. Historically, the identification of postcranial pneumaticity in non-avian dinosaurs has been based on examination of external morphology, and few studies thus far have focused on the internal architecture of pneumatic structures inside the bones. Here, we describe the vertebral pneumaticity of the ornithomimosaur Archaeornithomimus with the aid of X-ray computed tomography (CT) imaging. Complementary examination of external and internal osteology reveals (1) highly pneumatized cervical vertebrae with an elaborate configuration of interconnected chambers within the neural arch and the centrum; (2) anterior dorsal vertebrae with pneumatic chambers inside the neural arch; (3) apneumatic sacral vertebrae; and (4) a subset of proximal caudal vertebrae with limited pneumatic invasion into the neural arch. Comparisons with other theropod dinosaurs suggest that ornithomimosaurs primitively exhibited a plesiomorphic theropod condition for axial pneumaticity that was extended among later taxa, such as Archaeornithomimus and large bodied Deinocheirus. This finding corroborates the notion that evolutionary increases in vertebral pneumaticity occurred in parallel among independent lineages of bird-line archosaurs. Beyond providing a comprehensive view of vertebral pneumaticity in a non-avian coelurosaur, this study demonstrates the utility and need of CT imaging for further clarifying the early evolutionary history of postcranial pneumaticity.     

Observations on Ctenurella (Ptyctodontida) and the classification of placoderm fishes

The structure of the hyoid arch of the ptyctodont Ctenurella is described and discussed with reference to theories of jaw suspension in placoderm fishes. It is concluded that primitive placoderms had a modified hyoid arch but that the hyomandibular took no direct role in supporting the jaws. The relationships of ptyctodonts are discussed and it is concluded that they are placoderm fishes. Several different classifications of placoderm fishes are evaluated and are shown to be weakly based, chiefly because of lack of precise knowledge of character distribution. An attempt is made to produce a classification by using a simple cladistic computer analysis. The result highlights homoplasy in character distribution amongst traditionally recognized placoderm groups.     

Muscles of the masticatory apparatus in two genera of hyraces (Procavia and Heterohyrax)

Subungulate hyraces are similar to the condition assumed to have characterized primitive ungulates and subungulates by virtue of their small body size, relatively unspecialized cranial and postcranial anatomy, and primitive type of lophodont dentition. The muscles of mastication of Procavia habessinica and Heterohyrax brucei are here compared with those of other mammals, both with ungulates, as an example of more specialized mammals, and with opossums, as an example of more generalized mammals, to determine aspects of hyrax myology that represent the retention of a condition primitive for herbivorous mammals. The masticatory muscles of hyraces retain the primitive ungulate/subungulate condition in the large, complexly subdivided temporalis, and in the enlarged, pinnated, bilayered medial pterygoid. The medial pterygoid originates from the pterygoid hamulus, a condition that may also be primitive for this assemblage. The large complex superficial masseter is derived compared with the condition in ruminant artiodactyls, but may represent the condition primitive for perissodactyls. The architectural modifications of this muscle in hyraces may represent adaptations to allow a wide gape threat display. Hyraces possess a posterior belly of the digastric alone, paralleling the condition in some perissodactyls. They possess a large and complexly subdivided styloglossus, which may be a shared derived character of subungulates. Hyraces are unique among ungulates and subungulates in the extreme reduction of the anterior hyoid cornua, and may be unique among mammals in the development of paired lingual processes from the ceratohyal ossifications.     

Comparative morphology of the tentorium and hypopharyngeal–premental sclerites in sporophagous and non‐sporophagous adult Aleocharinae (Coleoptera: Staphylinidae)

We elucidate the configuration of the tentorium and the sclerites of the hypopharynx–prementum complex in selected spore‐ (pollen‐) and non‐spore‐feeding Aleocharinae (Staphylinidae) by presenting the first comparative 3D reconstructions of these structures for 19 staphylinoid beetle species (six outgroups, 13 Aleocharinae). General organization of the tentorium follows the groundplan previously proposed for adult Staphylinidae, although some taxa have reduced or lost the dorsal (all Aleocharinae studied, Agathidium mandibulare [Leiodidae]) or anterior (Omalium rivulare [Omaliinae], Anotylus sculpturatus [Oxytelinae]) tentorial arms. All species investigated have premental and hypopharyngeal sclerites that are partly homologizable across taxa. We clarified that Musculus praementopalpalis externus originates from the premental sclerite, resolving its unclear origin reported in our previous publications. Eight of 13 investigated Aleocharinae species are spore/pollen feeders, six obligatorily. Three of these six (Eumicrota, Gyrophaena fasciata, G. gentilis) have grinding pseudomolae and a fully developed hypopharyngeal suspensorium with posterior bridge and anterior elongations; the remaining three (Oxypoda, Pagla, Polylobus) lack pseudomolae and suspensorial bridge, but have the suspensorium elongated anteriorly. The dorsolateral side of the hypopharyngeal sclerite interacts with the pseudomola. Obligate sporophagy/pollinivory apparently arose at least three times in Aleocharinae, not always involving the pseudomola–hypopharynx grinding mechanism.     

Hyoid and pharyngeal arch function during ventilation and feeding in elasmobranchs: Conservation and modification in function

The hypothesis that the mandibular and hyoid arches evolved from anterior pharyngeal arches to increase ventilation performance and subsequently became adapted for feeding is widely accepted. As jaws evolved, the morphology of the hyoid arch changed notably from that of a pharyngeal arch. Furthermore, hyoid arch morphology varies considerably among elasmobranch taxa and has been shown to be related to feeding style. The goal of this study is to determine whether the function (direction of movement or change in cavity cross‐section) of the hyoid arch is altered from that of the pharyngeal arch, and whether function is altered between ventilation, the basal behavior, and feeding, the derived behavior. Similar effects and associations of the pharyngeal arches by orientation to feeding or ventilation are also investigated. The kinematics of the hyoid and second pharyngeal arch during ventilation and feeding are quantified using sonomicrometry and hyomandibular angle measured in five shark and one skate species representing widely divergent hyomandibular morphologies. Hyoid and pharyngeal cavity width follows the same pattern of movement during ventilation; therefore the hyoid arch retains the ancestral function of the pharyngeal arches. The orientation of the hyomandibular cartilage appears to influence the pattern of arch movement during ventilation: anterior directed elements decrease in cavity width; laterally directed elements increase in cavity width; while posterior directed elements increase in cavity width or do not change; while cavity depth increases in all species. Hyoid and pharyngeal cavity width movement differs among the species during feeding and also appears to be related to hyoid arch orientation as well as feeding style. There appears to be a division between those species with hyomandibular angles less than 110° from those that are greater between feeding mode and hyoid cavity width movement. Primarily suction feeding species decrease hyoid cavity width whereas primarily bite feeding species increase hyoid cavity width during feeding while all species increase hyoid cavity depth.