The postcranial system is composed of the axial and appendicular skeletons. The axial skeleton, w... more The postcranial system is composed of the axial and appendicular skeletons. The axial skeleton, which consists of serially repeating segments commonly known as vertebrae, protects and provides leverage for movement of the body. Across the vertebral column, much numerical and morphological diversity can be observed, which is associated with axial regionalization. The present article discusses this basic diversity and the early developmental mechanisms that guide vertebral formation and regionalization. An examination of vertebral numbers across the major vertebrate clades finds that actinopterygian and chondrichthyan fishes tend to increase vertebral number in the caudal region whereas Sarcopterygii increase the number of vertebrae in the precaudal region, although exceptions to each trend exist. Given the different regions of axial morphospace that are occupied by these groups, differential developmental processes control the axial patterning of actinopterygian and sarcopterygian species. It is possible that, among a variety of factors, the differential selective regimes for aquatic versus terrestrial locomotion have led to the differential use of axial morphospace in vertebrates.
Vertebrates exhibit tremendous diversity in body shape, though quantifying this variation has bee... more Vertebrates exhibit tremendous diversity in body shape, though quantifying this variation has been challenging. In the past, researchers have used simplified metrics that either describe overall shape but reveal little about its anatomical basis or that characterize only a subset of the morphological features that contribute to shape variation. Here, we present a revised metric of body shape, the vertebrate shape index (VSI), which combines the four primary morphological components that lead to shape diversity in vertebrates: head shape, length of the second major body axis (depth or width), and shape of the precaudal and caudal regions of the vertebral column. We illustrate the usefulness of VSI on a data set of 194 species, primarily representing five major vertebrate clades: Actinopterygii, Lissamphibia, Squamata, Aves, and Mammalia. We quantify VSI diversity within each of these clades and, in the course of doing so, show how measurements of the morphological components of VSI can be obtained from radiographs, articulated skeletons, and cleared and stained specimens. We also demonstrate that head shape, secondary body axis, and vertebral characteristics are important independent contributors to body shape diversity, though their importance varies across vertebrate groups. Finally, we present a functional application of VSI to test a hypothesized relationship between body shape and the degree of axial bending associated with locomotor modes in ray-finned fishes. Altogether, our study highlights the promise VSI holds for identifying the morphological variation underlying body shape diversity as well as the selective factors driving shape evolution.
Animal movement and its muscular control are central topics in functional morphology. As experime... more Animal movement and its muscular control are central topics in functional morphology. As experimentalists we often manipulate stimuli in a controlled setting or compare species to observe the degree of variation in movement and motor control of particular behaviors. Understanding and communicating the biological significance of these sources of variability requires a universal terminology that is presently lacking in the functional morphology literature. We suggest that ʻstereotypyʼ be used to refer to the degree of variability observed in a behavior across trials under a given set of conditions. The ability of an organism to alter its behavior across experimental treatments is referred to as ʻflexibilityʼ. We discuss how there has been a tendency to confound the phenomenon of a behavior exhibiting low variability, which we refer to as stereotyped, with inflexibility, or the inability to alter the behavior in response to a change in stimulus. The degree of stereotypy and flexibility in a behavior need not be correlated, nor need they have a common underlying basis. Coordination, a term used to describe the relationship between different body parts during movement, can be stereotyped and can show flexibility. Stereotypy of coordination can be assessed by the strength of correlations between movements of two body parts. The influence of coordination coherence on behavioral performance has rarely been considered, and could shed light on how taxa differ in their ability to perform behaviors. We suggest definitions of the terms stereotypy, flexibility and coordination, and provide examples of how and when these terms could be used when discussing behavioral changes in functional morphology.
Research on suction-feeding performance has mostly focused on measuring individual underlying com... more Research on suction-feeding performance has mostly focused on measuring individual underlying components such as suction pressure, flow velocity, ram or the effects of suction-induced forces on prey movement during feeding. Although this body of work has advanced our understanding of aquatic feeding, no consensus has yet emerged on how to combine all of these variables to predict prey-capture performance. Here, we treated the aquatic predator-prey encounter as a hydrodynamic interaction between a solid particle (representing the prey) and the unsteady suction flows around it, to integrate the effects of morphology, physiology, skull kinematics, ram and fluid mechanics on suction-feeding performance. We developed the suctioninduced force-field (SIFF) model to study suction-feeding performance in 18 species of centrarchid fishes, and asked what morphological and functional traits underlie the evolution of feeding performance on three types of prey. Performance gradients obtained using SIFF revealed that different trait combinations contribute to the ability to feed on attached, evasive and (strainsensitive) zooplanktonic prey because these prey types impose different challenges on the predator. The low overlap in the importance of different traits in determining performance also indicated that the evolution of suction-feeding ability along different ecological axes is largely unconstrained. SIFF also yielded estimates of feeding ability that performed better than kinematic traits in explaining natural patterns of prey use. When compared with principal components describing variation in the kinematics of suction-feeding events, SIFF output explained significantly more variation in centrarchid diets, suggesting that the inclusion of more mechanistic hydrodynamic models holds promise for gaining insight into the evolution of aquatic feeding performance.
When suction-feeding vertebrates expand their buccal cavity to draw water into their mouth, they ... more When suction-feeding vertebrates expand their buccal cavity to draw water into their mouth, they also exert a hydrodynamic force on their prey. This force is key to strike success, directly countering forces exerted by escaping or clinging prey. While the ability to produce high flow accelerations in front of the mouth is central to the predatorʼs ability to exert high forces on the prey, several mechanisms can contribute to the disparity between the potential and realized performance through their effect on flow and acceleration as experienced by the prey. In the present study, we test how interspecific variation in gape size, mouth displacement speed and the fishʼs ability to locate prey at the optimal position affect variation in the force exerted on attached prey. We directly measured these forces by allowing bluegill sunfish and largemouth bass to strike at ghost shrimp tethered to a load cell that recorded force at 5000 Hz, while synchronously recording strikes with a 500 Hz video. Strike kinematics of largemouth bass were slower than that of bluegill, as were estimated flow speeds and the force exerted on the prey. This difference in force persisted after taking into account the faster suction flows and accelerations of bluegill, and was only accounted for by considering interspecific differences in gape size, mouth displacement speed and fishʼs ability to locate the prey at the optimal position. The contribution to interspecific differences in the force exerted on the prey was estimated to be 42% for flow speed, 25% for strike efficiency, 3% for gape size and 30% for mouth displacement speed. Hence, kinematic diversity results in substantial differences in suction performance, beyond those expected based on the capacity to generate a high flow velocity. This functional complexity, in the form of biomechanically independent mechanisms that are recruited for one function, can potentially mitigate performance trade-offs in suction-feeding fishes.
Convergence is central to the study of evolution because it demonstrates the power of natural sel... more Convergence is central to the study of evolution because it demonstrates the power of natural selection to deterministically shape phenotypic diversity. However, the conditions under which a common morphology repeatedly evolves may be restrictive. Many factors, such as differing genetic and environmental backgrounds and many-to-one mapping of form to function, contribute to variability in responses to selection. Nevertheless, lineages may evolve similar, even if not identical, forms given a shared selective regime, providing opportunities to examine the relative importance of natural selection, constraint, and contingency. Here, we show that following 10 transitions to durophagy (eating hard-shelled prey) in moray eels (Muraenidae), cranial morphology repeatedly evolved toward a novel region of morphological space indicative of enhanced feeding performance on hard prey. Disparity among the resulting 15 durophagous species, however, is greater than disparity among ancestors that fed on large evasive prey, contradicting the pattern expected under convergence. This elevated disparity is a consequence of lineage-specific responses to durophagy, in which independent transitions vary in the suites of traits exhibiting the largest changes. Our results reveal a pattern of imperfect convergence, which suggests shared selection may actually promote diversification because lineages often differ in their phenotypic responses to similar selective demands.
Anguilliformes are an ecologically diverse group of predominantly marine fishes whose members are... more Anguilliformes are an ecologically diverse group of predominantly marine fishes whose members are easily recognized by their extremely elongate bodies, and universal lack of pelvic fins. Recent studies based on mitochondrial loci, including full mitogenomes, have called into question the monophyly of both the Anguilliformes, which appear to be paraphyletic without the inclusion of the Saccopharyngiformes (gulper eels and allies), as well as other more commonly known eel families (e.g., Congridae, Serrivomeridae). However, no study to date has investigated anguilliform interrelationships using nuclear loci. Here we present a new phylogenetic hypothesis for the Anguilliformes based on five markers (the nuclear loci Early Growth Hormone 3, Myosin Heavy Polypeptide 6 and Recombinase Activating Gene 1, as well as the mitochondrial genes Cytochrome b and Cytochrome Oxidase I). Our sampling spans 148 species and includes 19 of the 20 extant families of anguilliforms and saccopharyngiforms. Maximum likelihood analysis reveals that saccopharyngiform eels are deeply nested within the anguilliforms, and supports the non-monophyly of Congridae and Nettastomatidae, as well as that of Derichthyidae and Chlopsidae. Our analyses suggest that Protanguilla may be the sister group of the Synaphobranchidae, though the recent hypothesis that this species is the sister group to all other anguilliforms cannot be rejected. The molecular phylogeny, time-calibrated using a Bayesian relaxed clock approach and seven fossil calibration points, reveals a Late Cretaceous origin of this expanded anguilliform clade (stem age $116 Ma, crown age $99 Ma). Most major (family level) lineages originated between the end of the Cretaceous and Early Eocene, suggesting that anguilliform radiation may have been facilitated by the recovery of marine ecosystems following the KP extinction.
The shape of the body affects how organisms move, where they live, and how they feed. One body pl... more The shape of the body affects how organisms move, where they live, and how they feed. One body plan that has long engaged the interest of both evolutionary biologists and functional morphologists is axial elongation. There is a growing interest in the correlates and evolution of elongation within different terrestrial and aquatic vertebrate clades. At first glance, Anguilliformes may appear to exhibit a single cylindrical form but there is considerable diversity underlying this seemingly simplified body plan. Here, we explore evolution of the axial skeleton in 54 anguilliform taxa and some close relatives. We describe the diversity of axial elongation as well as investigate how characters such as head length, branchial-arch length, and shape of the pectoral fins correlate with vertebral number to possibly facilitate changes in absolute diameter of the body. Overall, we find that precaudal vertebral numbers and caudal vertebral numbers are evolving independently across elopomorph fishes. We also find that precaudal and caudal vertebral aspect ratios are evolving together across elopomorph fishes. When focusing within Anguilliformes we find striking diversity in the mechanisms of elongation of the body, including almost every trend for axial elongation known within actinopterygian fishes. The three major clades of eels we examined have slightly different mechanisms of elongation. We also find a suite of morphological characters associated with elongation in anguilliform fishes that appears to coincide with a more fossorial lifestyle such as high elongation ratios, a more posteriorly extended-branchial region, and a reduction in the size of the pectoral fins. Lastly, we point out that a diverse range of derived behaviors such as head-and tail-first burrowing, rotational feeding, and knotting around prey are only found in long cylindrical vertebrates.
The effects of prey size on prey-handling behavior for 60 ingestively naive hatchling Elaphe hele... more The effects of prey size on prey-handling behavior for 60 ingestively naive hatchling Elaphe helena were studied in the laboratory. Hatchlings were randomly assigned to one of three diet categories in which prey (Mus musculus) varied by relative mass differences of 20-35%, 40-46%, or 50-59% of an individual snake's own body mass. The effects of prey size on capture position, direction of ingestion, condition of prey at ingestion (dead/alive), feeding duration, and prey-handling tactic were observed and recorded for each feeding episode. Results indicated that prey size significantly affected the prey-handling behavior of hatchling E. helena. In the largest relative mass category, hatchlings captured prey by the anterior end more often than in the smaller two relative mass categories. Prey from the smallest relative mass category were simply seized whereas, in the medium and large categories, pinion and constriction behaviors were observed. Time to subdue and ingest the prey item increased with prey size categories.
... In this study, snakes that exhibited more than one prey restraint behaviour bent their bodies... more ... In this study, snakes that exhibited more than one prey restraint behaviour bent their bodies laterally to wrap around prey. Lateral bending also appears to permit flexibility in which portion of the body can be used to restrain prey: anterior or posterior. ...
The postcranial system is composed of the axial and appendicular skeletons. The axial skeleton, w... more The postcranial system is composed of the axial and appendicular skeletons. The axial skeleton, which consists of serially repeating segments commonly known as vertebrae, protects and provides leverage for movement of the body. Across the vertebral column, much numerical and morphological diversity can be observed, which is associated with axial regionalization. The present article discusses this basic diversity and the early developmental mechanisms that guide vertebral formation and regionalization. An examination of vertebral numbers across the major vertebrate clades finds that actinopterygian and chondrichthyan fishes tend to increase vertebral number in the caudal region whereas Sarcopterygii increase the number of vertebrae in the precaudal region, although exceptions to each trend exist. Given the different regions of axial morphospace that are occupied by these groups, differential developmental processes control the axial patterning of actinopterygian and sarcopterygian species. It is possible that, among a variety of factors, the differential selective regimes for aquatic versus terrestrial locomotion have led to the differential use of axial morphospace in vertebrates.
Vertebrates exhibit tremendous diversity in body shape, though quantifying this variation has bee... more Vertebrates exhibit tremendous diversity in body shape, though quantifying this variation has been challenging. In the past, researchers have used simplified metrics that either describe overall shape but reveal little about its anatomical basis or that characterize only a subset of the morphological features that contribute to shape variation. Here, we present a revised metric of body shape, the vertebrate shape index (VSI), which combines the four primary morphological components that lead to shape diversity in vertebrates: head shape, length of the second major body axis (depth or width), and shape of the precaudal and caudal regions of the vertebral column. We illustrate the usefulness of VSI on a data set of 194 species, primarily representing five major vertebrate clades: Actinopterygii, Lissamphibia, Squamata, Aves, and Mammalia. We quantify VSI diversity within each of these clades and, in the course of doing so, show how measurements of the morphological components of VSI can be obtained from radiographs, articulated skeletons, and cleared and stained specimens. We also demonstrate that head shape, secondary body axis, and vertebral characteristics are important independent contributors to body shape diversity, though their importance varies across vertebrate groups. Finally, we present a functional application of VSI to test a hypothesized relationship between body shape and the degree of axial bending associated with locomotor modes in ray-finned fishes. Altogether, our study highlights the promise VSI holds for identifying the morphological variation underlying body shape diversity as well as the selective factors driving shape evolution.
Animal movement and its muscular control are central topics in functional morphology. As experime... more Animal movement and its muscular control are central topics in functional morphology. As experimentalists we often manipulate stimuli in a controlled setting or compare species to observe the degree of variation in movement and motor control of particular behaviors. Understanding and communicating the biological significance of these sources of variability requires a universal terminology that is presently lacking in the functional morphology literature. We suggest that ʻstereotypyʼ be used to refer to the degree of variability observed in a behavior across trials under a given set of conditions. The ability of an organism to alter its behavior across experimental treatments is referred to as ʻflexibilityʼ. We discuss how there has been a tendency to confound the phenomenon of a behavior exhibiting low variability, which we refer to as stereotyped, with inflexibility, or the inability to alter the behavior in response to a change in stimulus. The degree of stereotypy and flexibility in a behavior need not be correlated, nor need they have a common underlying basis. Coordination, a term used to describe the relationship between different body parts during movement, can be stereotyped and can show flexibility. Stereotypy of coordination can be assessed by the strength of correlations between movements of two body parts. The influence of coordination coherence on behavioral performance has rarely been considered, and could shed light on how taxa differ in their ability to perform behaviors. We suggest definitions of the terms stereotypy, flexibility and coordination, and provide examples of how and when these terms could be used when discussing behavioral changes in functional morphology.
Research on suction-feeding performance has mostly focused on measuring individual underlying com... more Research on suction-feeding performance has mostly focused on measuring individual underlying components such as suction pressure, flow velocity, ram or the effects of suction-induced forces on prey movement during feeding. Although this body of work has advanced our understanding of aquatic feeding, no consensus has yet emerged on how to combine all of these variables to predict prey-capture performance. Here, we treated the aquatic predator-prey encounter as a hydrodynamic interaction between a solid particle (representing the prey) and the unsteady suction flows around it, to integrate the effects of morphology, physiology, skull kinematics, ram and fluid mechanics on suction-feeding performance. We developed the suctioninduced force-field (SIFF) model to study suction-feeding performance in 18 species of centrarchid fishes, and asked what morphological and functional traits underlie the evolution of feeding performance on three types of prey. Performance gradients obtained using SIFF revealed that different trait combinations contribute to the ability to feed on attached, evasive and (strainsensitive) zooplanktonic prey because these prey types impose different challenges on the predator. The low overlap in the importance of different traits in determining performance also indicated that the evolution of suction-feeding ability along different ecological axes is largely unconstrained. SIFF also yielded estimates of feeding ability that performed better than kinematic traits in explaining natural patterns of prey use. When compared with principal components describing variation in the kinematics of suction-feeding events, SIFF output explained significantly more variation in centrarchid diets, suggesting that the inclusion of more mechanistic hydrodynamic models holds promise for gaining insight into the evolution of aquatic feeding performance.
When suction-feeding vertebrates expand their buccal cavity to draw water into their mouth, they ... more When suction-feeding vertebrates expand their buccal cavity to draw water into their mouth, they also exert a hydrodynamic force on their prey. This force is key to strike success, directly countering forces exerted by escaping or clinging prey. While the ability to produce high flow accelerations in front of the mouth is central to the predatorʼs ability to exert high forces on the prey, several mechanisms can contribute to the disparity between the potential and realized performance through their effect on flow and acceleration as experienced by the prey. In the present study, we test how interspecific variation in gape size, mouth displacement speed and the fishʼs ability to locate prey at the optimal position affect variation in the force exerted on attached prey. We directly measured these forces by allowing bluegill sunfish and largemouth bass to strike at ghost shrimp tethered to a load cell that recorded force at 5000 Hz, while synchronously recording strikes with a 500 Hz video. Strike kinematics of largemouth bass were slower than that of bluegill, as were estimated flow speeds and the force exerted on the prey. This difference in force persisted after taking into account the faster suction flows and accelerations of bluegill, and was only accounted for by considering interspecific differences in gape size, mouth displacement speed and fishʼs ability to locate the prey at the optimal position. The contribution to interspecific differences in the force exerted on the prey was estimated to be 42% for flow speed, 25% for strike efficiency, 3% for gape size and 30% for mouth displacement speed. Hence, kinematic diversity results in substantial differences in suction performance, beyond those expected based on the capacity to generate a high flow velocity. This functional complexity, in the form of biomechanically independent mechanisms that are recruited for one function, can potentially mitigate performance trade-offs in suction-feeding fishes.
Convergence is central to the study of evolution because it demonstrates the power of natural sel... more Convergence is central to the study of evolution because it demonstrates the power of natural selection to deterministically shape phenotypic diversity. However, the conditions under which a common morphology repeatedly evolves may be restrictive. Many factors, such as differing genetic and environmental backgrounds and many-to-one mapping of form to function, contribute to variability in responses to selection. Nevertheless, lineages may evolve similar, even if not identical, forms given a shared selective regime, providing opportunities to examine the relative importance of natural selection, constraint, and contingency. Here, we show that following 10 transitions to durophagy (eating hard-shelled prey) in moray eels (Muraenidae), cranial morphology repeatedly evolved toward a novel region of morphological space indicative of enhanced feeding performance on hard prey. Disparity among the resulting 15 durophagous species, however, is greater than disparity among ancestors that fed on large evasive prey, contradicting the pattern expected under convergence. This elevated disparity is a consequence of lineage-specific responses to durophagy, in which independent transitions vary in the suites of traits exhibiting the largest changes. Our results reveal a pattern of imperfect convergence, which suggests shared selection may actually promote diversification because lineages often differ in their phenotypic responses to similar selective demands.
Anguilliformes are an ecologically diverse group of predominantly marine fishes whose members are... more Anguilliformes are an ecologically diverse group of predominantly marine fishes whose members are easily recognized by their extremely elongate bodies, and universal lack of pelvic fins. Recent studies based on mitochondrial loci, including full mitogenomes, have called into question the monophyly of both the Anguilliformes, which appear to be paraphyletic without the inclusion of the Saccopharyngiformes (gulper eels and allies), as well as other more commonly known eel families (e.g., Congridae, Serrivomeridae). However, no study to date has investigated anguilliform interrelationships using nuclear loci. Here we present a new phylogenetic hypothesis for the Anguilliformes based on five markers (the nuclear loci Early Growth Hormone 3, Myosin Heavy Polypeptide 6 and Recombinase Activating Gene 1, as well as the mitochondrial genes Cytochrome b and Cytochrome Oxidase I). Our sampling spans 148 species and includes 19 of the 20 extant families of anguilliforms and saccopharyngiforms. Maximum likelihood analysis reveals that saccopharyngiform eels are deeply nested within the anguilliforms, and supports the non-monophyly of Congridae and Nettastomatidae, as well as that of Derichthyidae and Chlopsidae. Our analyses suggest that Protanguilla may be the sister group of the Synaphobranchidae, though the recent hypothesis that this species is the sister group to all other anguilliforms cannot be rejected. The molecular phylogeny, time-calibrated using a Bayesian relaxed clock approach and seven fossil calibration points, reveals a Late Cretaceous origin of this expanded anguilliform clade (stem age $116 Ma, crown age $99 Ma). Most major (family level) lineages originated between the end of the Cretaceous and Early Eocene, suggesting that anguilliform radiation may have been facilitated by the recovery of marine ecosystems following the KP extinction.
The shape of the body affects how organisms move, where they live, and how they feed. One body pl... more The shape of the body affects how organisms move, where they live, and how they feed. One body plan that has long engaged the interest of both evolutionary biologists and functional morphologists is axial elongation. There is a growing interest in the correlates and evolution of elongation within different terrestrial and aquatic vertebrate clades. At first glance, Anguilliformes may appear to exhibit a single cylindrical form but there is considerable diversity underlying this seemingly simplified body plan. Here, we explore evolution of the axial skeleton in 54 anguilliform taxa and some close relatives. We describe the diversity of axial elongation as well as investigate how characters such as head length, branchial-arch length, and shape of the pectoral fins correlate with vertebral number to possibly facilitate changes in absolute diameter of the body. Overall, we find that precaudal vertebral numbers and caudal vertebral numbers are evolving independently across elopomorph fishes. We also find that precaudal and caudal vertebral aspect ratios are evolving together across elopomorph fishes. When focusing within Anguilliformes we find striking diversity in the mechanisms of elongation of the body, including almost every trend for axial elongation known within actinopterygian fishes. The three major clades of eels we examined have slightly different mechanisms of elongation. We also find a suite of morphological characters associated with elongation in anguilliform fishes that appears to coincide with a more fossorial lifestyle such as high elongation ratios, a more posteriorly extended-branchial region, and a reduction in the size of the pectoral fins. Lastly, we point out that a diverse range of derived behaviors such as head-and tail-first burrowing, rotational feeding, and knotting around prey are only found in long cylindrical vertebrates.
The effects of prey size on prey-handling behavior for 60 ingestively naive hatchling Elaphe hele... more The effects of prey size on prey-handling behavior for 60 ingestively naive hatchling Elaphe helena were studied in the laboratory. Hatchlings were randomly assigned to one of three diet categories in which prey (Mus musculus) varied by relative mass differences of 20-35%, 40-46%, or 50-59% of an individual snake's own body mass. The effects of prey size on capture position, direction of ingestion, condition of prey at ingestion (dead/alive), feeding duration, and prey-handling tactic were observed and recorded for each feeding episode. Results indicated that prey size significantly affected the prey-handling behavior of hatchling E. helena. In the largest relative mass category, hatchlings captured prey by the anterior end more often than in the smaller two relative mass categories. Prey from the smallest relative mass category were simply seized whereas, in the medium and large categories, pinion and constriction behaviors were observed. Time to subdue and ingest the prey item increased with prey size categories.
... In this study, snakes that exhibited more than one prey restraint behaviour bent their bodies... more ... In this study, snakes that exhibited more than one prey restraint behaviour bent their bodies laterally to wrap around prey. Lateral bending also appears to permit flexibility in which portion of the body can be used to restrain prey: anterior or posterior. ...
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Papers by Rita Mehta