Functional morphology of the feeding apparatus of ten species of Pacific butterflyfishes (Perciformes,Chaetodontidae): an ecomorphological approach

Synopsis The functional morphology of the feeding apparatus and the feeding ecology of an assemblage of ten species of butterflyfishes was investigated using a comparative ecomorphological approach. Behavioral observations in situ and in vivo, morphological measurements of fresh-killed specimens, scanning electron microscopy, and kinematic modeling were utilized. The fishes show varying degrees of morphological and behavioral specialization and generalization. The more specialized species group according to how they feed, rather than what they feed on. The feeding guild concept is therefore not very instructive in understanding the functional significance of the feeding apparatus. Many of the morphologically specialized butterflyfishes demonstrate evolutionary convergence in feeding morphology. Whereas the more morphologically specialized species do generally have more ecologically specialized diets, these data do not particularly support the ecomorphological hypothesis in that similar diets do not correspond to similar morphologies.     

The morphology and evolution of the ventral gill arch skeleton in batoid fishes (Chondrichthyes: Batoidea)

The ventral gill arch skeleton was examined in some representatives of batoid fishes. The homology of the components was elucidated by comparing similarities and differences among the components of the ventral gill arches in chondrichthyans, and attempts were made to justify the homology by giving causal mechanisms of chondrogenesis associated with the ventral gill arch skeleton. The ceratohyal is present in some batoid fishes, and its functional replacement, the pseudohyal, seems incomplete in most groups of batoid fishes, except in stingrays. The medial fusion of the pseudohyal with successive ceratobranchials occurs to varying degrees among stingray groups. The ankylosis between the last two ceratobranchials occurs uniquely in stingrays, and it serves as part of the insertion of the last pair of coracobranchialis muscles. The basihyal is possibly independently lost in electric rays, the stingray genus Urotrygon (except U. daviesi) and pelagic myiiobatoid stingrays. The first hypobranchial is oriented anteriorly or anteromedially, and it varies in shape and size among batoid fishes. It is represented by rami projecting posterolaterally from the basihyal in sawfishes, guitarfishes and skates. It consists of a small piece of cartilage which extends anteromedially from the medial end of the first ccratobranchial in electric rays. It is a large cartilaginous plate in most of stingrays. It is absent in pelagic myliobatoid stingrays. The remaining hypobranchial cartilages also vary in shape and size among batoid fishes. Torpedo and possibly the Jurassic Belemnobalis and Spathobatis possess the generalized or typical chondrichthyan ventral gill arch structure in which the hypobranchials form a Σ-shaped pattern. In the electric ray Hypnos and narkinidid and narcinidid electric rays, the hypobranchial components are oriented longitudinally along the mid-portion of the ventral gill arches. They form a single cartilaginous plate in the narkinidid electric rays, Narcine and Diplobatis. In guitarfishes and skates, the second hypobranchial is unspecialized, and in skates, it does not have a direct contact with the second ceratobranchial. In both groups, the third and fourth hypobranchials are composed of a small cartilage which forms a passage for the afferent branches of the ventral aorta and serve as part of the insertion of the coracobranchialis muscle. In sawfishes and stingrays, the hypobranchials appear to be included in the medial plate. In sawfishes, the second and third components separately chondrify in adults, but the fourth component appears to be fused with the middle medial plate. In stingrays, a large medial plate appears to include the second through to the last hypobranchial and most of the basibranchial copulae. The medial plate probably develops independently in sawfishes and stingrays. Because the last basibranchial copula appears to be a composite of one to two hypobranchials and at least two basibranchial copulae, the medial plate may be formed by several developmental processes of chondrogenesis. More detailed comparative anatomical and developmental studies are needed to unveil morphogenesis and patternings of the ventral gill arch skeleton in batoid fishes.     

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.     

Molecular and karyological aspects of Batoidea (Chondrichthyes, Elasmobranchi) phylogeny

Although considerable progress has been made in elucidating the relationships within the Chondrichthyes, there is no agreement as it concerns the systematics of Batoidea, the most derived superorder among cartilaginous fishes, and many different interpretations exist. Our investigation provides the first assessment of relationships among the described batoid species using sequences from both mtDNA and nuclear genes as well as karyological morphology. Our work consists primarily in reconstructing the phylogenetic relationships of Batoidea by examining the mtDNA (16S) and nuclear gene (18S) sequences from 11 batoid species. The three analytical methods (NJ, MP and Bayesian analysis) grouped Rajiformes, Myliobatiformes and Rhinobatiformes. In these trees the two torpedoes diverge from the other batoid fishes. We also compare the molecular data with the available karyological evidence, which consist of the diploid number and the karyotype morphology of eight species belonging to the four orders examined. The results show that the karyological structure in the different species is generally consistent with the various phylogenetical trees, and that Torpediniformes confirm their unique genome organization.     

Convergent, parallel and correlated evolution of trophic morphologies in the subfamily schizothoracinae from the Qinghai-Tibetan plateau

Schizothoracine fishes distributed in the water system of the Qinghai-Tibetan plateau (QTP) and adjacent areas are characterized by being highly adaptive to the cold and hypoxic environment of the plateau, as well as by a high degree of diversity in trophic morphology due to resource polymorphisms. Although convergent and parallel evolution are prevalent in the organisms of the QTP, it remains unknown whether similar evolutionary patterns have occurred in the schizothoracine fishes. Here, we constructed for the first time a tentative molecular phylogeny of the schizothoracine fishes based on the complete sequences of the cytochrome b gene. We employed this molecular phylogenetic framework to examine the evolution of trophic morphologies. We used Pagel's maximum likelihood method to estimate the evolutionary associations of trophic morphologies and food resource use. Our results showed that the molecular and published morphological phylogenies of Schizothoracinae are partially incongruent with respect to some intergeneric relationships. The phylogenetic results revealed that four character states of five trophic morphologies and of food resource use evolved at least twice during the diversification of the subfamily. State transitions are the result of evolutionary patterns including either convergence or parallelism or both. Furthermore, our analyses indicate that some characters of trophic morphologies in the Schizothoracinae have undergone correlated evolution, which are somewhat correlated with different food resource uses. Collectively, our results reveal new examples of convergent and parallel evolution in the organisms of the QTP. The adaptation to different trophic niches through the modification of trophic morphologies and feeding behaviour as found in the schizothoracine fishes may account for the formation and maintenance of the high degree of diversity and radiations in fish communities endemic to QTP.     

Development and Evolution of Dentition Pattern and Tooth Order in the Skates And Rays (Batoidea; Chondrichthyes)

Shark and ray (elasmobranch) dentitions are well known for their multiple generations of teeth, with isolated teeth being common in the fossil record. However, how the diverse dentitions characteristic of elasmobranchs form is still poorly understood. Data on the development and maintenance of the dental patterning in this major vertebrate group will allow comparisons to other morphologically diverse taxa, including the bony fishes, in order to identify shared pattern characters for the vertebrate dentition as a whole. Data is especially lacking from the Batoidea (skates and rays), hence our objective is to compile data on embryonic and adult batoid tooth development contributing to ordering of the dentition, from cleared and stained specimens and micro-CT scans, with 3D rendered models. We selected species (adult and embryonic) spanning phylogenetically significant batoid clades, such that our observations may raise questions about relationships within the batoids, particularly with respect to current molecular-based analyses. We include developmental data from embryos of recent model organisms Leucoraja erinacea and Raja clavata to evaluate the earliest establishment of the dentition. Characters of the batoid dentition investigated include alternate addition of teeth as offset successional tooth rows (versus single separate files), presence of a symphyseal initiator region (symphyseal tooth present, or absent, but with two parasymphyseal teeth) and a restriction to tooth addition along each jaw reducing the number of tooth families, relative to addition of successor teeth within each family. Our ultimate aim is to understand the shared characters of the batoids, and whether or not these dental characters are shared more broadly within elasmobranchs, by comparing these to dentitions in shark outgroups. These developmental morphological analyses will provide a solid basis to better understand dental evolution in these important vertebrate groups as well as the general plesiomorphic vertebrate dental condition.     

Edgeworth's legacy of cranial muscle development with an analysis of muscles in the ventral gill arch region of batoid fishes (Chondrichthyes: Batoidea).

A series of studies by Edgeworth demonstrated that cranial muscles of gnathostome fishes are embryologically of somitic origin, originating from the mandibular, hyoid, branchial, epibranchial, and hypobranchial muscle plates. Recent experimental studies using quail-chick chimeras support Edgeworth's view on the developmental origin of cranial muscles. One of his findings, the existence of the premyogenic condensation constrictor dorsalis in teleost fishes, has also been confirmed by molecular developmental studies. Therefore, developmental mechanisms for patterning of cranial muscles, as described and implicated by Edgeworth, may serve as structural entities or regulatory phenomena responsible for developmental and evolutionary changes. With Edgeworth's and other studies as background, muscles in the ventral gill arch region of batoid fishes are analyzed and compared with those of other gnathostome fishes. The spiracularis is regarded as homologous at least within batoid fishes, but its status within elasmobranchs remains unclear; developmental modifications of the spiracularis proper are evident in some batoid fishes and in several shark groups. The peculiar ventral extension of the spiracularis in electric rays and some stingrays may represent convergence, probably facilitating ventilation and/or feeding in both groups. The evolutionary origin of the "internus" and "externus" remains uncertain, despite the fact that a variety of forms of the constrictor superficiales ventrales in batoid fishes indicates an actual medio-ventral extension of the "externus." The intermandibularis is probably present only in electric rays. The "X" muscle occurs only in electric rays and is considered to be Edgeworth's intermandibularis profundus. Its association with the adductor mandibular complex in narkinidid and narcinidid electric rays may relate to its functional role in lower jaw movement. Contrary to common belief, in most batoid fishes as well as some sharks, muscles that originate from the branchial muscle plate and extend medially in the ventral gill arches do exist: the medial extension of the interbranchiales in most batoid fishes and some sharks and the "Y" muscle in the pelagic stingrays Myliobatos and Rhinoptera. The latter is another example of the medial extension of the "internus." Whether the interbranchiales and "Y" muscle are homologous within elasmobranchs and whether homologous with the obliques ventrales and/or transversi ventrales of osteichthyan fishes await further research. Four hypobranchial muscles are recognized in batoid fishes: the coracomandibularis, coracohyoideus, coracoarcualis, and coracohyomandibularis. The coracohyoideus is discrete from the coracoarcualis; its complete structural separation from the latter occurs in several groups of batoid fishes.(ABSTRACT TRUNCATED AT 400 WORDS)     

Body plan convergence in the evolution of skates and rays (Chondrichthyes: Batoidea)

Skates, rays and allies (Batoidea) comprise more than half of the species diversity and much of the morphological disparity among chondrichthyan fishes, the sister group to all other jawed vertebrates. While batoids are morphologically well characterized and have an excellent fossil record, there is currently no consensus on the interrelationships of family-level taxa. Here we construct a resolved, robust and time-calibrated batoid phylogeny using mitochondrial genomes, nuclear genes, and fossils, sampling densely across taxa. Data partitioning schemes, biases in the sequence data, and the relative informativeness of each fossil are explored. The molecular phylogeny is largely congruent with morphology crownward in the tree, but the branching orders of major batoid groups are mostly novel. Body plan convergence appears to be widespread in batoids. A depressed, rounded pectoral disk supported to the snout tip by fin radials, common to skates and stingrays, is indicated to have been derived independently by each group, while the long, spiny rostrum of sawfishes similarly appears to be convergent with that of sawsharks, which are not batoids. The major extant batoid lineages are inferred to have arisen relatively rapidly from the Late Triassic into the Jurassic, with long stems followed by subsequent radiations in each group around the Cretaceous/Tertiary boundary. The fossil record indicates that batoids were affected with disproportionate severity by the end-Cretaceous extinction event.     

New insights into the enameloid microstructure of batoid fishes (Chondrichthyes)

Chondrichthyan teeth are capped with a hypermineralized tissue known as enameloid. Its microstructure displays a hierarchical organization that has increased in structural complexity from a homogenous single‐crystallite enameloid (SCE) in early Chondricthyans to the complex multilayered enameloid found in modern sharks (consisting of bundles of crystallites arranged in intriguing patterns). Recent analyses of the enameloid microstructure in batoid fishes, focused on Myliobatiformes and fossil taxa, point to the presence of a bundled (or fibred) multilayered enameloid, a condition proposed as plesiomorphic for Batoidea. In this work, we provide further enameloid analysis for a selection of taxa covering the phylogeny of batoids. Our SEM analysis shows a superficial layer of SCE, where individualized crystallites are clearly discernable, capping the teeth in most of the species studied. A bundled double‐layered enameloid was found only in a Rhinoidei, Rhina ancylostoma Bloch & Schneider, 1801. We conclude that the most widespread condition among extant batoids is a monolayer SCE lacking microstructural differentiation, probably plesiomorphic at least for crown batoidea. We suggest that the complex bundled enameloid present in other batoids is a convergent character that has appeared repeatedly during the evolution of batoids, probably as a mechanical adaptation towards moderate durophagous diets.     

Advances in the Study of Feeding Behaviors, Mechanisms, and Mechanics of Sharks

Sharks as a group have a long history as highly successful predatory fishes. Although, the number of recent studies on their diet, feeding behavior, feeding mechanism, and mechanics have increased, many areas still require additional investigation. Dietary studies of sharks are generally more abundant than those on feeding activity patterns, and most of the studies are confined to relatively few species, many being carcharhiniform sharks. These studies reveal that sharks are generally asynchronous opportunistic feeders on the most abundant prey item, which are primarily other fishes. Studies of natural feeding behavior are few and many observations of feeding behavior are based on anecdotal reports. To capture their prey sharks either ram, suction, bite, filter, or use a combination of these behaviors. Foraging may be solitary or aggregate, and while cooperative foraging has been hypothesized it has not been conclusively demonstrated. Studies on the anatomy of the feeding mechanism are abundant and thorough, and far exceed the number of functional studies. Many of these studies have investigated the functional role of morphological features such as the protrusible upper jaw, but only recently have we begun to interpret the mechanics of the feeding apparatus and how it affects feeding behavior. Teeth are represented in the fossil record and are readily available in extant sharks. Therefore much is known about their morphology but again functional studies are primarily theoretical and await experimental analysis. Recent mechanistic approaches to the study of prey capture have revealed that kinematic and motor patterns are conserved in many species and that the ability to modulate feeding behavior varies greatly among taxa. In addition, the relationship of jaw suspension to feeding behavior is not as clear as was once believed, and contrary to previous interpretations upper jaw protrusibility appears to be related to the morphology of the upper jaw-chondrocranial articulation rather than the type of jaw suspension. Finally, we propose a set of specific hypotheses including: (1) The functional specialization for suction feeding hypothesis that morphological and functional specialization for suction feeding has repeatedly arisen in numerous elasmobranch lineages, (2) The aquatic suction feeding functional convergence hypothesis that similar hydrodynamic constraints in bony fishes and sharks result in convergent morphological and functional specializations for suction feeding in both groups, (3) The feeding modulation hypothesis that suction capture events in sharks are more stereotyped and therefore less modulated compared to ram and bite capture events, and (4) The independence of jaw suspension and feeding behavior hypothesis whereby the traditional categorization of jaw suspension types in sharks is not a good predictor of jaw mobility and prey capture behavior. Together with a set of questions these hypotheses help to guide future research on the feeding biology of sharks.     

Ecomorphological relationships among Caribbean tetraodontiform fishes

The anatomy of the oral jaw apparatus, lever-arm mechanics and the diet of six species of Caribbean fishes in the order Tetraodontiformes were investigated to explore the relationships between trophic morphology and feeding habit in these fishes. Tetraodontiforms use their oral jaw apparatus to capture and reduce a broad range of prey types such as plankton, polychaete worms, holothuroids, sea urchins, crabs, molluscs, gorgonians and algae. The different feeding habits of tetraodontiforms are reflected by differences in the morphological and biomechanical features of their oral jaw apparatus that appear to enhance their abilities to feed on hard prey organisms. Species that bite and crush hard, benthic prey organisms had more massive bones and muscles, longer jaw-opening in-levers, and higher jaw-closing lever ratios than the planktivorous, suction-feeding species. Masses of the jaw and suspensorium bones and lower jaw adductor muscles as well as the jaw-opening in-levers and jaw-closing lever ratios of crushers were greater than those of biters. In contrast, the mass of the adductor muscle of the upper jaw did not vary among species with different diets, indicating that this muscle may not be central to the factors that determine patterns of prey use in these fishes. The diversity of feeding behaviours and the wide range of feeding habits among fishes in the order Tetraodontiformes illustrate the versatility of the oral jaw apparatus as a single functional feeding system in fishes.     

Determinants of tapeworm species richness in elasmobranch fishes: untangling environmental and phylogenetic influences

Parasite species richness is a fundamental characteristic of host species and varies substantially among host communities. Hypotheses aiming to explain observed patterns of richness are numerous, and none is universal. In this study, we use tapeworm parasites of elasmobranch fishes to examine the phylogenetic and environmental influences on the variation in species richness for this specific system. Tapeworms are the most diverse group of helminths to infect elasmobranchs. Elasmobranchs are cosmopolitan in distribution and their tapeworm parasites are remarkably host specific; therefore, making this an ideal system in which to examine global patterns in species diversity. Here, we 1) quantify the tapeworm richness in elasmobranch fishes, 2) identify the host features correlated with tapeworm richness, and 3) determine whether tapeworm richness follows a latitudinal gradient. The individual and combined effects of host size, factors associated with water temperatures (influenced by latitude and depth), host habitat, and type of elasmobranch (shark or batoid) on measures of species diversity were assessed using general linear models. These analyses included tapeworm host records for 317 different elasmobranch species (124 species were included in our analyses) and were conducted with and without taking into account phylogenetic relationships between host species. Since sharks and batoids differ substantially in body form, analyses were repeated for each host subset. On average, batoids harboured significantly more tapeworm species than shark hosts. Tapeworm richness in sharks was influenced by median depth, whereas no predictor variable included in our models could adequately account for interspecific variation in tapeworm richness in batoid hosts. The taxonomic diversity of tapeworm assemblages of sharks and batoids was influenced by median depth and median latitude, respectively. When the influence of host phylogeny is accounted for, larger hosts harbour a greater tapeworm richness, whereas hosts exploiting wider latitudinal ranges harbour more taxonomically distinct tapeworm assemblages. Species richness and taxonomic diversity of tapeworm assemblages in elasmobranch fishes are influenced by different evolutionary pressures, including host phylogenetic relationships, space constraints and geographical area. Our results suggest that ca 3600 tapeworm species have yet to be described from elasmobranch fishes.     

Histology and structural integration of the major morphologies of the Cypriniform Weberian apparatus

The Weberian apparatus, a diagnostic feature of otophysan fishes, is a novel hearing adaptation integrating several developmental and morphological systems (ear-vertebral column-swim bladder). Otophysan fishes are one of the largest and most successful freshwater clades, with over 10,000 species across most continents. The largest otophysan order, Cypriniformes, dominates the freshwaters of Asia, Europe, North America, and Africa. Spanning such a wide variety of environments, the Weberian apparatus undergoes morphological modifications to maintain functionality. Within Cypriniformes, we propose three distinct morphological classes of the Weberian apparatus based on the level of skeletal expansion around the swim bladder: simple (typical of most Cyprinidae), anterior plate (found in families such as Gyrinocheilidae, Catostomidae, and Botiidae), and encapsulated (either single-capsule as found, e.g., in Gobionidae and Cobitidae, or double-capsule as found, e.g., in Nemacheilidae and Balitoridae). Little ontological or comparative data exists regarding the construction or integration of these different morphologies, and less is known about the tissue level integration and variation within these morphologies. We used paraffin histology to document the hard and soft tissue anatomy of the Weberian apparatus in six species representing all morphological classes. We found sites of similarity across the morphologies including size and structure of the saccule, aspects of ossicle ossification, and swim bladder tunica composition, indicating potential sites of developmental and functional constraint. In contrast, we found differences across both auditory and nonauditory features in otic chamber size, ossification within ossicles and other vertebral elements, and composition of ligaments, indicating likely sites of adaptability. Some of these changes are likely evolutionary (taxonomic), but may be influenced by the environmental niche occupied by the clade. These results show a clear need for increased ontological and comparative study of the complete cypriniform Weberian apparatus, particularly histologically, as well as increased auditory studies across morphological types.     

Rare ecomorphological convergence on a complex adaptive landscape: Body size and diet mediate evolution of jaw shape in squirrels (Sciuridae)

Convergence is widely regarded as compelling evidence for adaptation, often being portrayed as evidence that phenotypic outcomes are predictable from ecology, overriding contingencies of history. However, repeated outcomes may be very rare unless adaptive landscapes are simple, structured by strong ecological and functional constraints. One such constraint may be a limitation on body size because performance often scales with size, allowing species to adapt to challenging functions by modifying only size. When size is constrained, species might adapt by changing shape; convergent shapes may therefore be common when size is limiting and functions are challenging. We examine the roles of size and diet as determinants of jaw shape in Sciuridae. As expected, size and diet have significant interdependent effects on jaw shape and ecomorphological convergence is rare, typically involving demanding diets and limiting sizes. More surprising is morphological without ecological convergence, which is equally common between and within dietary classes. Those cases, like rare ecomorphological convergence, may be consequences of evolving on an adaptive landscape shaped by many‐to‐many relationships between ecology and function, many‐to‐one relationships between form and performance, and one‐to‐many relationships between functionally versatile morphologies and ecology. On complex adaptive landscapes, ecological selection can yield different outcomes.     

Evolutionary dynamics of complex biomechanical systems: an example using the four-bar mechanism

Like many phenotypic traits, biomechanical systems are defined by both an underlying morphology and an emergent functional property. The relationship between these levels may have a profound impact on how selection for functional performance is translated into morphological evolution. In particular, complex mechanical systems are likely to be highly redundant, because many alternative morphologies yield equivalent functions. We suggest that this redundancy weakens the relationship between morphological and functional diversity, and we illustrate this effect using an evolutionary model of the four-bar lever system of labrid fishes. Our results demonstrate that, when traits are complex, the morphological diversity of a clade may only weakly predict its mechanical diversity. Furthermore, parallel or convergent selection on function does not necessarily produce convergence in morphology. Empirical observations suggest that this weak form-function relationship has contributed to the morphological diversity of labrid fishes, as functionally equivalent species may nevertheless possess morphologically distinct jaws. We suggest that partial decoupling of morphology and mechanics due to redundancy is a major factor in morphological diversification.     

Morphological-dietary relationships within two assemblages of marine demersal fishes

In this study we have used morphological characters related to feeding and prey capture and dietary data to investigate the trophic organization within two assemblages of marine demersal fishes. Morphological and dietary disparities within fish assemblages were estimated from species similarities based on Euclidean distances plus species projections on the principal axes from multivariate analyses. The analyses of the morphological variables indicated that species in each assemblage comprised morphologically distinct groups strongly influenced by trophically linked characters. Stomach content analyses revealed that fish species in each assemblage were classified into three basic feeding groups: polychaete-shrimp feeders, crab feeders and fish feeders. These results indicated that food resource partitioning was operating within each assemblage. However, when morphological and trophic data were compared no significant correlations were found. The results did not particularly support the ecomorphological hypothesis that dietary differences are due to morphological differences, since similar diets do not correspond to similar morphologies. The patterns of trophic organization within the fish assemblages examined, possibly reflect differences among species due not only to the effect of ecological demands on morphology but also by their evolutionary history and constructional constraints imposed by phylogeny. This revised version was published online in August 2006 with corrections to the Cover Date.     

Interspecific dietary diversity has little influence on pathways of glucose metabolism in liver and heart of piranhas and pacus (family Serrasalmidae)

The Serrasalmidae is a speciose family of Neotropical freshwater fishes with diverse interspecies dietary preferences, from the herbivorous or frugivorous pacus to the omnivorous or carnivorous piranhas. The Serrasalmidae is an ideal comparative model to explore the biochemical correlates of dietary preference. For example, debate exists about whether variation in dietary preference among fishes influences the capacities for de novo glucose synthesis (gluconeogenesis) or glucose use as metabolic fuel (glycolysis). We predicted that carnivory in piranhas would be associated with an enhanced gluconeogenic capacity compared with the herbivorous pacus, because carnivores consume less carbohydrate. As a corollary, we expected similar glycolytic capacities at a major site of glucose use (heart) in fishes with different diets. We measured activities of key enzymes of gluconeogenesis and the opposing reactions of glycolysis in liver (the primary gluconeogenic site) and heart of five serrasalmid species showing an interspecies dietary gradient from herbivory to omnivory to carnivory. The interspecific variation in enzyme activities was, in general, unrelated to dietary preference. Glycolytic capacity in heart was lower in herbivores but this may be related to non-diet factors. Among serrasalmid fishes, the enzymatic capacities of glucose metabolism are not overtly influenced by interspecific divergence of dietary preference.     

Convergence in trophic morphology and feeding performance among piscivorous natricine snakes

Piscivory has independently evolved numerous times amongst snakes, and therefore these animals provide a powerful opportunity to test for convergent evolution in a vertebrate feeding system. In this study, we integrate performance trials with comparative methods to test the hypothesis that piscivory drives convergence in trophic morphology and feeding performance among natricine snakes. Within and across species, increasing the relative length of the quadrate bone in the skull is positively and strongly linked to a reduction in the time needed to swallow large fish prey. Thus, our feeding experiments suggest that a longer quadrate bone enables snakes to better conform their head shape to the shape of the prey during swallowing. Ancestral diet reconstructions and phylogenetically corrected multiple regression analyses further reveal that evolutionary increases in piscivory are coupled to the evolution of relatively longer quadrates, and hence improved feeding performance on fish prey in these animals. The exploitation of similar dietary niches drives the evolution of convergent trophic morphologies and feeding performances in natricine snakes.