Microevolutionary patterns in the common caiman predict macroevolutionary trends across extant crocodilians

Both extinct and extant crocodilians have repeatedly diversified in skull shape along a continuum, from narrow‐snouted to broad‐snouted phenotypes. These patterns occur with striking regularity, although it is currently unknown whether these trends also apply to microevolutionary divergence during population differentiation or the early stages of speciation. Assessing patterns of intraspecific variation within a single taxon can potentially provide insight into the processes of macroevolutionary differentiation. For example, high levels of intraspecific variation along a narrow‐broad axis would be consistent with the view that cranial shapes can show predictable patterns of differentiation on relatively short timescales, and potentially scale up to explain broader macroevolutionary patterns. In the present study, we use geometric morphometric methods to characterize intraspecific cranial shape variation among groups within a single, widely distributed clade, Caiman crocodilus. We show that C. crocodilus skulls vary along a narrow/broad‐snouted continuum, with different subspecies strongly clustered at distinct ends of the continuum. We quantitatively compare these microevolutionary trends with patterns of diversity at macroevolutionary scales (among all extant crocodilians). We find that morphological differences among the subspecies of C. crocodilus parallel the patterns of morphological differentiation across extant crocodilians, with the primary axes of morphological diversity being highly correlated across the two scales. We find intraspecific cranial shape variation within C. crocodilus to span variation characterized by more than half of living species. We show the main axis of intraspecific phenotypic variation to align with the principal direction of macroevolutionary diversification in crocodilian cranial shape, suggesting that mechanisms of microevolutionary divergence within species may also explain broader patterns of diversification at higher taxonomic levels.     

Shape variation in a benthic stream fish across flow regimes

Evolution of fish body shapes in flowing and non-flowing waters have been examined for several species. Flowing water can select for fish body shapes that increase steady swimming efficiency, whereas non-flowing water can favor shapes that increase unsteady swimming efficiency. Benthic stream fishes often use areas near the substrate that exhibit reduced or turbulent flow, thus it is unclear which swimming forms would be favored in such environments, and how shape might change across flow regimes. To test the relationship between fish body shape and flow regime in a benthic stream fish, we used geometric morphometric techniques to characterize lateral body shape in mountain sucker (Catostomus platyrhynchus) across flow rates, using stream gradient as an indicator of stream flow. Mountain suckers from low-flow environments were more streamlined, consistent with steady swimming body shapes, whereas mountain suckers from high flows had deeper bodies, consistent with unsteady swimming body shapes. In addition, smaller individuals tended to have more robust body shapes. These patterns are opposite to those predicted for stream fishes in the mid-water column. The benthic stream environment represents a distinct selective environment for fish shape that does not appear to conform to the simple dichotomy of flowing versus non-flowing water.     

Getting in shape: habitat‐based morphological divergence for two sympatric fishes

Freshwater fishes often show large amounts of body shape variation across divergent habitats and, in most cases, the observed differences have been attributed to the environmental pressures of living in lentic or lotic habitats. Previous studies have suggested a distinct set characters and morphological features for species occupying each habitat under the steady–unsteady swimming performance model. We tested this model and assessed body shape variation using geometric morphometrics for two widespread fishes, Goodea atripinnis (Goodeidae) and Chirostoma jordani (Atherinopsidae), inhabiting lentic and lotic habitats across the Mesa Central of Mexico. These species were previously shown to display little genetic variation across their respective ranges. Our body shape analyses reveal morphometric differentiation along the same axes for both species in each habitat. Both possess a deeper body shape in lentic habitats and a more streamlined body in lotic habitats, although the degree of divergence between habitats was less for C. jordani. Differences in the position of the mouth differed between habitats as well, with both species possessing a more superior mouth in lentic habitats. These recovered patterns are generally consistent with the steady–unsteady swimming model and highlight the significance of environmental forces in driving parallel body shape differences of organisms in divergent habitats. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 152–162.     

Diversity and Evolution of Body Size in Fishes

The diversity of body sizes observed among species of a clade is a combined result of microevolutionary processes (i.e. natural selection and genetic drift) that cause size changes within phylogenetic lineages, and macroevolutionary processes (i.e. speciation and extinction) that affect net rates of diversification among lineages. Here we assess trends of size diversity and evolution in fishes (non-tetrapod craniates), employing paleontological, macroecological, and phylogenetic information. Fishes are well suited to studies of size diversity and evolution, as they are highly diverse, representing more than 50% of all living vertebrate species, and many fish taxa are well represented in the fossil record from throughout the Phanerozoic. Further, the frequency distributions of sizes among fish lineages resemble those of most other animal taxa, in being right-skewed, even on a log scale. Using an approach that measures rates of size evolution (in darwins) within a formal phylogenetic framework, we interpret the shape of size distributions as a balance between the competing forces of diversification, pushing taxa away from ancestral values, and of conservation, drawing taxa closer to a central tendency. Within this context we show how non-directional mechanisms of evolution (i.e. passive diffusion processes) can produce an hitherto unperceived bias to larger size, when size is measured on the conventional log scale. These results demonstrate how the interpretation of macroecological datasets can be enriched from an historical perspective, and document the ways in which macroevolutionary and microevolutionary processes may be decoupled in the production of size diversity.     

Habitat transitions alter the adaptive landscape and shape phenotypic evolution in needlefishes (Belonidae)

Habitat occupancy can have a profound influence on macroevolutionary dynamics, and a switch in major habitat type may alter the evolutionary trajectory of a lineage. In this study, we investigate how evolutionary transitions between marine and freshwater habitats affect macroevolutionary adaptive landscapes, using needlefishes (Belonidae) as a model system. We examined the evolution of body shape and size in marine and freshwater needlefishes and tested for phenotypic change in response to transitions between habitats. Using micro‐computed tomographic (µCT) scanning and geometric morphometrics, we quantified body shape, size, and vertebral counts of 31 belonid species. We then examined the pattern and tempo of body shape and size evolution using phylogenetic comparative methods. Our results show that transitions from marine to freshwater habitats have altered the adaptive landscape for needlefishes and expanded morphospace relative to marine taxa. We provide further evidence that freshwater taxa attain reduced sizes either through dwarfism (as inferred from axial skeletal reduction) or through developmental truncation (as inferred from axial skeletal loss). We propose that transitions to freshwater habitats produce morphological novelty in response to novel prey resources and changes in locomotor demands. We find that repeated invasions of different habitats have prompted predictable changes in morphology.     

Morphological selection in an extreme flow environment: body shape and waterfall-climbing success in the Hawaiian stream fish Sicyopterus stimpsoni

Flow characteristics are a prominent factor determining body shapes in aquatic organisms, and correlations between body shape and ambient flow regimes have been established for many fish species. In this study, we investigated the potential for a brief period of extreme flow to exert selection on the body shape of juvenile climbing Hawaiian gobiid fishes. Because of an amphidromous life history, juvenile gobies that complete an oceanic larval phase return to freshwater habitats, where they become adults. Returning juveniles often must scale waterfalls (typically with the use of a ventral sucker) in order to reach the habitats they will use as adults, thereby exposing these animals to brief periods of extreme velocities of flow. Hydrodynamic theory predicts that bodies with larger suckers and with lower heights that reduce drag would have improved climbing success and, thus, be well suited to meet the demands of the flows in waterfalls. To test the potential for the flow environment of waterfalls to impose selection that could contribute to differences in body shape between islands, we subjected juvenile Sicyopterus stimpsoni to climbing trials up artificial waterfalls (～100 body lengths) and measured differences in body shape between successful and unsuccessful climbers. Waterfalls appear to represent a significant selective barrier to these fishes, as nearly 30% failed our climbing test. However, the effects of selection on morphology were not straightforward, as significant differences in shape between successful and unsuccessful climbers did not always match hydrodynamic predictions. In both selection experiments and in adult fish collected from habitats with different prevailing conditions of flow (the islands of Hawai'i versus Kaua'i), lower head heights were associated with exposure to high-flow regimes, as predicted by hydrodynamic theory. Thus, a premium appears to be placed on the reduction of drag via head morphology throughout the ontogeny of this species. The congruence of phenotypic selection patterns observed in our experiments, with morphological character divergence documented among adult fish from Hawai'i and Kaua'i, suggests that differences in morphology between subpopulations of adult climbing gobies may result, at least in part, from the selective pressures of high-velocity flows encountered by migrating juveniles.     

Environmental and scale-dependent evolutionary trends in the body size of crustaceans

The ecological and physiological significance of body size is well recognized. However, key macroevolutionary questions regarding the dependency of body size trends on the taxonomic scale of analysis and the role of environment in controlling long-term evolution of body size are largely unknown. Here, we evaluate these issues for decapod crustaceans, a group that diversified in the Mesozoic. A compilation of body size data for 792 brachyuran crab and lobster species reveals that their maximum, mean and median body size increased, but no increase in minimum size was observed. This increase is not expressed within lineages, but is rather a product of the appearance and/or diversification of new clades of larger, primarily burrowing to shelter-seeking decapods. This argues against directional selective pressures within lineages. Rather, the trend is a macroevolutionary consequence of species sorting: preferential origination of new decapod clades with intrinsically larger body sizes. Furthermore, body size evolution appears to have been habitat-controlled. In the Cretaceous, reef-associated crabs became markedly smaller than those in other habitats, a pattern that persists today. The long-term increase in body size of crabs and lobsters, coupled with their increased diversity and abundance, suggests that their ecological impact may have increased over evolutionary time.     

Disentangling the functional roles of morphology and motion in the swimming of fish

In fishes the shape of the body and the swimming mode generally are correlated. Slender-bodied fishes such as eels, lampreys, and many sharks tend to swim in the anguilliform mode, in which much of the body undulates at high amplitude. Fishes with broad tails and a narrow caudal peduncle, in contrast, tend to swim in the carangiform mode, in which the tail undulates at high amplitude. Such fishes also tend to have different wake structures. Carangiform swimmers generally produce two staggered vortices per tail beat and a strong downstream jet, while anguilliform swimmers produce a more complex wake, containing at least two pairs of vortices per tail beat and relatively little downstream flow. Are these differences a result of the different swimming modes or of the different body shapes, or both? Disentangling the functional roles requires a multipronged approach, using experiments on live fishes as well as computational simulations and physical models. We present experimental results from swimming eels (anguilliform), bluegill sunfish (carangiform), and rainbow trout (subcarangiform) that demonstrate differences in the wakes and in swimming performance. The swimming of mackerel and lamprey was also simulated computationally with realistic body shapes and both swimming modes: the normal carangiform mackerel and anguilliform lamprey, then an anguilliform mackerel and carangiform lamprey. The gross structure of simulated wakes (single versus double vortex row) depended strongly on Strouhal number, while body shape influenced the complexity of the vortex row, and the swimming mode had the weakest effect. Performance was affected even by small differences in the wakes: both experimental and computational results indicate that anguilliform swimmers are more efficient at lower swimming speeds, while carangiform swimmers are more efficient at high speed. At high Reynolds number, the lamprey-shaped swimmer produced a more complex wake than the mackerel-shaped swimmer, similar to the experimental results. Finally, we show results from a simple physical model of a flapping fin, using fins of different flexural stiffness. When actuated in the same way, fins of different stiffnesses propel themselves at different speeds with different kinematics. Future experimental and computational work will need to consider the mechanisms underlying production of the anguilliform and carangiform swimming modes, because anguilliform swimmers tend to be less stiff, in general, than are carangiform swimmers.     

Ecomorphology of Locomotion in Labrid Fishes

The Labridae is an ecologically diverse group of mostly reef associated marine fishes that swim primarily by oscillating their pectoral fins. To generate locomotor thrust, labrids employ the paired pectoral fins in motions that range from a fore-aft rowing stroke to a dorso-ventral flapping stroke. Species that emphasize one or the other behavior are expected to benefit from alternative fin shapes that maximize performance of their primary swimming behavior. We document the diversity of pectoral fin shape in 143 species of labrids from the Great Barrier Reef and the Caribbean. Pectoral fin aspect ratio ranged among species from 1.12 to 4.48 and showed a distribution with two peaks at about 2.0 and 3.0. Higher aspect ratio fins typically had a relatively long leading edge and were narrower distally. Body mass only explained 3% of the variation in fin aspect ratio in spite of four orders of magnitude range and an expectation that the advantages of high aspect ratio fins and flapping motion are greatest at large body sizes. Aspect ratio was correlated with the angle of attachment of the fin on the body (r = 0.65), indicating that the orientation of the pectoral girdle is rotated in high aspect ratio species to enable them to move their fin in a flapping motion. Field measures of routine swimming speed were made in 43 species from the Great Barrier Reef. Multiple regression revealed that fin aspect ratio explained 52% of the variation in size-corrected swimming speed, but the angle of attachment of the pectoral fin only explained an additional 2%. Labrid locomotor diversity appears to be related to a trade-off between efficiency of fast swimming and maneuverability in slow swimming species. Slow swimmers typically swim closer to the reef while fast swimmers dominate the water column and shallow, high-flow habitats. Planktivory was the most common trophic associate with high aspect ratio fins and fast swimming, apparently evolving six times.     

Rise and fall of †Pycnodontiformes: Diversity,competition and extinction of a successful fish clade

†Pycnodontiformes was a successful lineage of primarily marine fishes that broadly diversified during the Mesozoic. They possessed a wide variety of body shapes and were adapted to a broad range of food sources. Two other neopterygian clades possessing similar ecological adaptations in both body morphology (†Dapediiformes) and dentition (Ginglymodi) also occurred in Mesozoic seas. Although these groups occupied the same marine ecosystems, the role that competitive exclusion and niche partitioning played in their ability to survive alongside each other remains unknown. Using geometric morphometrics on both the lower jaw (as constraint for feeding adaptation) and body shape (as constraint for habitat adaptation), we show that while dapediiforms and ginglymodians occupy similar lower jaw morphospace, pycnodontiforms are completely separate. Separation also occurs between the clades in body shape so that competition reduction between pycnodontiforms and the other two clades would have resulted in niche partitioning. Competition within pycnodontiforms seemingly was reduced further by evolving different feeding strategies as shown by disparate jaw shapes that also indicate high levels of plasticity. Acanthomorpha was a teleostean clade that evolved later in the Mesozoic and which has been regarded as implicated in driving the pycnodontiforms to extinction. Although they share similar body shapes, no coeval acanthomorphs had similar jaw shapes or dentitions for dealing with hard prey like pycnodontiforms do and so their success being a factor in pycnodontiform extinction is unlikely. Sea surface temperature and eustatic variations also had no impact on pycnodontiform diversity patterns according to our results. Conversely, the occurrence and number of available reefs and hardgrounds as habitats through time seems to be the main factor in pycnodontiform success. Decline in such habitats during the Late Cretaceous and Palaeogene might have had deleterious consequences for pycnodontiform diversity. Acanthomorphs occupied the niches of pycnodontiforms during the terminal phase of their existence.     

Predictability of phenotypic differentiation across flow regimes in fishes

Fish inhabit environments greatly varying in intensity of water velocity, and these flow regimes are generally believed to be of major evolutionary significance. To what extent does water flow drive repeatable and predictable phenotypic differentiation? Although many investigators have examined phenotypic variation across flow gradients in fishes, no clear consensus regarding the nature of water velocity's effects on phenotypic diversity has yet emerged. Here, I describe a generalized model that produces testable hypotheses of morphological and locomotor differentiation between flow regimes in fishes. The model combines biomechanical information (describing how fish morphology determines locomotor abilities) with ecological information (describing how locomotor performance influences fitness) to yield predictions of divergent natural selection and phenotypic differentiation between low-flow and high-flow environments. To test the model's predictions of phenotypic differentiation, I synthesized the existing literature and conducted a meta-analysis. Based on results gathered from 80 studies, providing 115 tests of predictions, the model produced some accurate results across both intraspecific and interspecific scales, as differences in body shape, caudal fin shape, and steady-swimming performance strongly matched predictions. These results suggest that water flow drives predictable phenotypic variation in disparate groups of fish based on a common, generalized model, and that microevolutionary processes might often scale up to generate broader, interspecific patterns. However, too few studies have examined differentiation in body stiffness, muscle architecture, or unsteady-swimming performance to draw clear conclusions for those traits. The analysis revealed that, at the intraspecific scale, both genetic divergence and phenotypic plasticity play important roles in phenotypic differentiation across flow regimes, but we do not yet know the relative importance of these two sources of phenotypic variation. Moreover, while major patterns within and between species were predictable, we have little direct evidence regarding the role of water flow in driving speciation or generating broad, macroevolutionary patterns, as too few studies have addressed these topics or conducted analyses within a phylogenetic framework. Thus, flow regime does indeed drive some predictable phenotypic outcomes, but many questions remain unanswered. This study establishes a general model for predicting phenotypic differentiation across flow regimes in fishes, and should help guide future studies in fruitful directions, thereby enhancing our understanding of the predictability of phenotypic variation in nature.     

Coral reefs promote the evolution of morphological diversity and ecological novelty in labrid fishes

Although coral reefs are renowned biodiversity hotspots it is not known whether they also promote the evolution of exceptional ecomorphological diversity. We investigated this question by analysing a large functional morphological dataset of trophic characters within Labridae, a highly diverse group of fishes. Using an analysis that accounts for species relationships, the time available for diversification and model uncertainty we show that coral reef species have evolved functional morphological diversity twice as fast as non-reef species. In addition, coral reef species occupy 68.6% more trophic morphospace than non-reef species. Our results suggest that coral reef habitats promote the evolution of both trophic novelty and morphological diversity within fishes. Thus, the preservation of coral reefs is necessary, not only to safeguard current biological diversity but also to conserve the underlying mechanisms that can produce functional diversity in future.     

Predation drives morphological convergence in the Gambusia panuco species group among lotic and lentic habitats

Fish morphology is often constrained by a trade‐off between optimizing steady vs. unsteady swimming performance due to opposing effects of caudal peduncle size. Lotic environments tend to select for steady swimming performance, leading to smaller caudal peduncles, whereas predators tend to select for unsteady swimming performance, leading to larger caudal peduncles. However, it is unclear which aspect of performance should be optimized across heterogeneous flow and predation environments and how this heterogeneity may affect parallel phenotypic evolution. We investigated this question among four Gambusia species in north‐eastern Mexico, specifically the riverine G. panuco, the spring endemics G. alvarezi and G. hurtadoi, and a fourth species, G. marshi, found in a variety of habitats with varying predation pressure in the Cuatro Ciénegas Basin and Río Salado de Nadadores. We employed a geometric morphometric analysis to examine how body shapes of both male and female fish differ among species and habitats and with piscivore presence. We found that high‐predation and low‐predation species diverged morphologically, with G. marshi exhibiting a variable, intermediate body shape. Within G. marshi, body morphology converged in high‐predation environments regardless of flow velocity, and fish from high‐predation sites had larger relative caudal peduncle areas. However, we found that G. marshi from low‐predation environments diverged in morphology between sub‐basins of Cuatro Ciénegas, indicating other differences among these basins that merit further study. Our results suggest that a morphological trade‐off promotes parallel evolution of body shape in fishes colonizing high‐predation environments and that changing predation pressure can strongly impact morphological evolution in these species.     

Swimming efficiency and the influence of morphology on swimming costs in fishes

Swimming performance is considered a main character determining survival in many aquatic animals. Body morphology highly influences the energetic costs and efficiency of swimming and sets general limits on a species capacity to use habitats and foods. For two cyprinid fishes with different morphological characteristics, carp (Cyprinus carpio L.) and roach (Rutilus rutilus (L.)), optimum swimming speeds (U _mc) as well as total and net costs of transport (COT, NCOT) were determined to evaluate differences in their swimming efficiency. Costs of transport and optimum speeds proved to be allometric functions of fish mass. NCOT was higher but U _mc was lower in carp, indicating a lower swimming efficiency compared to roach. The differences in swimming costs are attributed to the different ecological demands of the species and could partly be explained by their morphological characteristics. Body fineness ratios were used to quantify the influence of body shape on activity costs. This factor proved to be significantly different between the species, indicating a better streamlining in roach with values closer to the optimum body form for efficient swimming. Net swimming costs were directly related to fish morphology.     

Effects of habitat modification on coastal fish assemblages

The purpose of this study was to assess the influence of anthropogenic modification of coastal habitats on fish assemblages in Taiwan, comparing the abundance, species richness and taxonomic composition of fishes on natural v. artificial habitats. While there was no significant variation in the abundance or richness of fishes between natural and artificial habitats, the species composition of fishes in artificial habitats was significantly different from that of natural habitats. Natural reefs were characterized by greater abundance of Stethojulis spp. (Labridae), Abudefduf spp. (Pomacentridae) and Thalassoma spp. (Labridae), whereas anthropogenic habitats were dominated by Parupeneus indicus (Mullidae), Pempheris oualensis (Pempheridae) and Parapriacanthus ransonneti (Pempheridae). In general, it appears that specialist reef-associated species are being replaced with fishes that are much more generalist in their habitat-use. The loss of natural coastal habitats may threaten some species that cannot live in anthropogenically altered habitats, though the overall abundance and diversity of coastal fishes was not significantly different between natural and artificial habitats in Taiwan.     

THE MICRO AND MACRO IN BODY SIZE EVOLUTION

The diversity of body sizes of organisms has traditionally been explained in terms of microevolutionary processes: natural selection owing to differential fitness of individual organisms, or to macroevolutionary processes: species selection owing to the differential proliferation of phylogenetic lineages. Data for terrestrial mammals and birds indicate that even on a logarithmic scale frequency distributions of body mass among species are significantly skewed towards larger sizes. We used simulation models to evaluate the extent to which macro- and microevolutionary processes are sufficient to explain these distributions. Simulations of a purely cladogenetic process with no bias in extinction or speciation rates for different body sizes did not produce skewed log body mass distributions. Simulations that included size-biased extinction rates, especially those that incorporated anagenetic size change within species between speciation and extinction events, regularly produced skewed distributions. We conclude that although cladogenetic processes probably play a significant role in body size evolution, there must also be a significant anagenetic component. The regular variation in the form of mammalian body size distributions among different-sized islands and continents suggests that environmental conditions, operating through both macro- and microevolutionary processes, determine to a large extent the diversification of body sizes within faunas. Macroevolution is not decoupled from microevolution.     

Perspectives on the genetic architecture of divergence in body shape in sticklebacks

The body shape of fishes encompasses a number of morphological traits that are intrinsically linked to functional systems and affect various measures of performance, including swimming, feeding, and avoiding predators. Changes in shape can allow a species to exploit a new ecological niche and can lead to ecological speciation. Body shape results from the integration of morphological, behavioral and physiological traits. It has been well established that functional interdependency among traits plays a large role in constraining the evolution of shape, affecting both the speed and the repeated evolution of particular body shapes. However, it is less clear what role genetic or developmental constraints might play in biasing the rate or direction of the evolution of body shape. Here, we suggest that the threespine stickleback (Gasterosteus aculeatus) is a powerful model system in which to address the extent to which genetic or developmental constraints play a role in the evolution of body shape in fishes. We review the existing data that begins to address these issues in sticklebacks and provide suggestions for future areas of research that will be particularly fruitful for illuminating the mechanisms that contribute to the evolution of body shape in fishes.     

Correlations between habitat use and body shape in a phrynosomatid lizard (Urosaurus ornatus): a population-level analysis

Recent ecomorphological studies have shown that the predicted correlations between morphology and ecology on broad taxonomic levels are often obscured when comparing more closely related groups. Among species, comparisons of lizards often indicate very little support for adaptive radiations into novel habitats. As few population level studies have been performed, we compared body, head and limb shape between four populations of Urosaurus ornatus living in structurally distinct habitats (cliffs, rocks, trees and boulders). Surprisingly, clear correlations between habitat use and body shape among populations were found, most of which were in good accordance with a priori biomechanical predictions (e.g. flat body and head for extreme climbers; long distal hindlimb segments for jumpers and runners; narrow body and long tail for tree dwelling lizards). This indicates that populations of Urosaurus ornatus are seemingly 'adapted' to the habitat they live in. However, quantification of performance and behaviour are needed to determine the adaptive nature of these observations.     

Correlated evolution of body and fin morphology in the cichlid fishes

Body and fin shapes are chief determinants of swimming performance in fishes. Different configurations of body and fin shapes can suit different locomotor specializations. The success of any configuration is dependent upon the hydrodynamic interactions between body and fins. Despite the importance of body–fin interactions for swimming, there are few data indicating whether body and fin configurations evolve in concert, or whether these structures vary independently. The cichlid fishes are a diverse family whose well‐studied phylogenetic relationships make them ideal for the study of macroevolution of ecomorphology. This study measured body, and caudal and median fin morphology from radiographs of 131 cichlid genera, using morphometrics and phylogenetic comparative methods to determine whether these traits exhibit correlated evolution. Partial least squares canonical analysis revealed that body, caudal fin, dorsal fin, and anal fin shapes all exhibited strong correlated evolution consistent with locomotor ecomorphology. Major patterns included the evolution of deep body profiles with long fins, suggestive of maneuvering specialization; and the evolution of narrow, elongate caudal peduncles with concave tails, a combination that characterizes economical cruisers. These results demonstrate that body shape evolution does not occur independently of other traits, but among a suite of other morphological changes that augment locomotor specialization.     

Turbulence: does vorticity affect the structure and shape of body and fin propulsors?

Over the past century, many ideas have been developed on the relationships between water flow and the structure and shape of the body and fins of fishes, largely during swimming in relatively steady flows. However, both swimming by fishes and the habitats they occupy are associated with vorticity, typically concentrated as eddies characteristic of turbulent flow. Deployment of methods to examine flow in detail suggests that vorticity impacts the lives of fishes. First, vorticity near the body and fins can increase thrust and smooth variations in thrust that are a consequence of using oscillating and undulating propulsors to swim. Second, substantial mechanical energy is dissipated in eddies in the wake and adaptations that minimize these losses would be anticipated. We suggest that such mechanisms may be found in varying the length of the propulsive wave, stiffening propulsive surfaces, and shifting to using median and paired fins when swimming at low speeds. Eddies in the flow encountered by fishes may be beneficial, but when eddy radii are of the order of 0.25 of the fish's total length, negative impacts occur due to greater difficulties in controlling stability. The archetypal streamlined "fish" shape reduces destabilizing forces for fishes swimming into eddies.