No full-text available
To read the full-text of this research,
you can request a copy directly from the authors.
Morphological and Mechanical Patterns of Evolution in Triggerfish Fins
ResearchGate has not been able to resolve any citations for this publication.
The genus Chirostoma is virtually restricted to the Mesa Central, the southern part of the Mexican Plateau. The evolution of the 18 species and six subspecies is examined from the standpoint of the major factors underlying their present distribution: the geological history of central México and their trophic relationships as deduced from morphology. An attempt is also made to find evolutionary patterns common to fish species flocks in general. The Mesa Central has a long history of geological instability. Tectonic movements associated with the Laramide Orogeny, mid-Tertiary and Plio-Pleistocene vulcanism and mid-Pleistocene uplift and associated vulcanism have resulted in continuously changing drainage patterns. During the Tertiary and early Pleistocene, the ancestral Río Lerma probably flowed westward, perhaps through a series of lakes, to the Pacific Ocean. The exact course and extent of the ancestral system at this time are unknown. Mid-Pleistocene uplift compartmentalized the drainage. At first the river was able to continue a course through the ancestral Chapala basin probably utilizing the present Ríos Tuxpan, Ameca and perhaps the Río Grande de Santiago drainages as exits. Finally, however, uplift blocked the flow and the present outlet to the Río Grande de Santiago was gained. Lakes, playa lakes, and lacustrine deposits which occur across the Mesa Central are evidence of the size of this late Pleistocene system. Changes in lake levels, presumably reflecting late Pleistocene pluvial periods, are indicated by terraces. Interior basins formerly part of the Lerma system, as indicated by the distribution of Chirostoma, are the Llanos de Puebla, Puebla-Tlaxcala; the Llanos de Apan, Hidalgo; the Valley of México, Federal District-State of México; Lakes Zirahuén, Pátzcuaro and Cuitzeo, Michoacán; Lakes Atotonilco and San Marcos, Jalisco; Lakes Santa Maria and San Pedro Lagunillas, Nayarit; Lake Juanacatlán, Jalisco and Laguna de Santiaguillo, Durango. There appear to have been two centers of evolution for the jordani species group: the Chapala basin and the Uruapan-Lakes Pátzcuaro and Zirahuén region of Michoacán. C. humboldtianum has a distribution which probably antedates the uplift, whereas the range of C. jordani more probably reflects the extent of aquatic connections across the Mesa Central late in the Pleistocene. The species of the arge species group have differentiated throughout most of the Lerma basin. The presence of C. arge in the headwaters of the Río Verde and the Río de la Laja is suggestive of the route utilized by the ancestor of the arge group during its movement onto the Plateau. C. labarcae and C. aculeatum have complementary distributions reflecting their centers of origin in the Chapala basin and the Bajío, respectively. The ranges of other species are too restricted to show any pattern. The coexistence of various numbers of species of Chirostoma is clearly facilitated by differences in body length. Prey are probably selected on the basis of size alone rather than kind. The diphyletic nature of the genus is also clearly revealed in the structure of the atherinid communities. The earlier radiating members of the jordani species group have filled most of the large species niches whereas the later evolving arge species occupy either the small species niches in the lacustrine communities or remain allopatric. C. aculeatum occurs in an area which has been free of the larger species for long periods of time. It is the only member of the arge group which has markedly increased in body length, thus demonstrating that character displacement has occurred. Character displacement has also occurred in the Chapala basin; the morphological differentiation of C. consocium consocium and C. c. reseratum is related to the number of coexisting species rather than the time of isolation. The same responses and interaction governing the evolution and adaptive radiation of Chirostoma seem to have also controlled the evolution of other fish species flocks although a lack of thorough phylogenetic systematic analyses precludes a rigorous discussion.
The potential relationship between ecological versatility and local distribution and abundance for 5 species of triggerfish was examined at Kimbe Bay, Papua New Guinea, The abundances of juvenile and adult Balistapus undulatus, Melichthys vidua, Rhinecanthus verrucosus, Sufflamen bursa and S. chrysopterus were quantified in a range of habitats along a typical coral reef profile, Four of the 5 species displayed distinct and relatively narrow distributions across the reef profile, with the fifth species, B, undulatus, being broadly distributed across all zones and depths, and the most abundant species. For each species, the spatial distribution of juveniles closely matched that of adults and juvenile densities were greater in species with more abundant adults. A detailed description of depth distributions of individuals indicated that shallow species had narrower depth ranges, In terms of microhabitat use, B. unduldtus was the most generalised species, occupying all the microhabitats that were identified. The other 4 species were specialised to varying degrees on different microhabitats. Results from this study provide the first detailed description of patterns of distribution and abundance, habitat use and ecological versatility in triggerfish. Distribution and abundance could partially be explained by differences in the degree to which each species is specialised, both in terms of depth and microhabitat selectivity. Hence, versatility in depth and microhabitat use may play an important role in determining the local distribution and relative abundance of coral reef fishes.
Superimposition methods for comparing configurations of landmarks in two or more specimens are reviewed. These methods show
differences in shape among specimens as residuals after rotation, translation, and scaling them so that they align as well
as possible. A new method is presented that generalizes Siegel and Benson's (1982) resistant-fit theta-rho analysis so that
more than two objects can be compared at the same time. Both least-squares and resistant-fit approaches are generalized to
allow for affine transformations (uniform shape change). The methods are compared, using artificial data and data on 18 landmarks
on the wings of 127 species of North American mosquitoes. Graphical techniques are also presented to help summarize the patterns
of differences in shape among the objects being compared.
Temporal asymmetries in clade histories have often been studied in lower Paleozoic radiations. Post-Paleozoic patterns, however, are less well understood. In this paper, disparity and diversity changes in Mesozoic heart urchins were analyzed at the ordinal level, with contrasts among the sister groups Holasteroida and Spatangoida, their paraphyletic stem group Disasteroida the more inclusive clade, the superorder Atelostomata. A 38-dimensional landmark-based morphospace representing test architecture was used to describe morphological evolution in terms of total variance and total range. Discordances between disparity and diversity were evident and were expressed both as deceleration in morphological diversification in all groups and as disproportionately higher disparity early in the histories of the Atelostomata, Holasteroida Spatangoida. The finding that the early atelostomate disparity peak coincides with the origin of the orders Holasteroida and Spatangoida lends support to the perception of orders as semi-independent entities in the biological hierarchy and as meaningful proxies for morphological distinctness.
A comparison of holasteroid and spatangoid responses to the end-Cretaceous mass extinction revealed morphological selectivity. Paleocene spatangoid survivors showed no change in disparity relative to the Campanian-Maastrichtian sample, suggesting nonselectivity. Holasteroids suffered a pronounced loss in disparity (despite a rather high Late Cretaceous level of disparity), indicating morphological selectivity of extinction.
Partitioning of disparity into plastral and nonplastral components, reflecting different degrees of developmental entrenchment and functionality, suggests that the origin of holasteroids and spatangoids is more consistent with an exploration of the developmental flexibility of nonplastral constructions than with uniform ecospace occupation. Within groups, several patterns were also most consistent with intrinsic controls. For plastral landmarks, there is an apparent increase in developmental modularity and decrease in developmental constraint from disasteroids to holasteroids and spatangoids. For nonplastral landmarks, no substantial change in disparity was observed from disasteroids to holasteroids and spatangoids, suggesting the maintenance of a developmental constraint despite the passage of time and ecological differentiation. More generally, this study suggests that certain topologies of disparity and evolutionary mechanisms potentially characteristic of the lower Paleozoic radiations of higher taxa (e.g., developmental flexibility) need not be confined to any given time period or hierarchical level.
The quantification of disparity is an important aspect of recent macroevolutionary studies, and it is usually motivated by theoretical considerations about the pace of innovation and the filling of morphospace. In practice, varying protocols of data collection and analysis have rendered comparisons among studies difficult. The basic question remains, How sensitive is any given disparity signal to different aspects of sampling and data analysis? Here we explore this issue in the context of the radiation of the echinoid order Spatangoida during the Cretaceous. We compare patterns at the genus and species levels, with time subdivision into subepochs and into stages, and with morphological sampling based on landmarks, traditional morphometrics, and discrete characters. In terms of temporal scale, similarity of disparity pattern accrues despite a change in temporal resolution, and a general deceleration in morphological diversification is apparent. Different morphometric methods also produce similar signals. Both the landmark analysis and the discrete character analysis suggest relatively high early disparity, whereas the analysis based on traditional morphometrics records a much lower value. This difference appears to reflect primarily the measurement of different aspects of overall morphology. Disparity patterns are similar at both the genus and species levels. Moreover, inclusion or exclusion of the sister order Holasteroida and the stem group Disasteroida in the sampled morphospace did not affect proportional changes in spatangoid disparity. Similar results were found for spatangoid subclades vis-à-vis spatangoids as a whole. The relative robustness of these patterns implies that the choice of temporal scale, morphometric scheme, and taxonomic level may not affect broad trends in disparity and the representation of large-scale morphospace structure.
A data matrix is presented of 210 morphological characters (mostly osteological, some external) for 20 extant taxa of the ten Recent families of tetraodontiform fishes and 36 fossil tetraodontiforms. The oldest of these are from the Upper Cretaceous (95 Mya); most are from the Lower to Middle Eocene (50–58 Mya). There are two outgroup taxa (a zeiform and a caproid). A cladistic analysis of this matrix for only the extant taxa produced two equally parsimonious trees that call into question the monophyly of some of the previously accepted major higher-level tetraodontiform clades. Inclusion in the analysis of the large number of available fossil taxa helps to resolve relationships between family level clades. The new phylogenetic hypothesis, together with stratigraphic and biogeographical data, is used to discuss scenarios of the origin and evolution of the major clades of the order. © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139, 565−617.
Covering <0·1% of the ocean’s surface, coral reefs harbour about one-third of all marine fishes or c. 5000 species. Allopatry (geographic isolation) is believed to be the primary mode of speciation, yet few biogeographic barriers exist between reefs, and most reef fishes have a pelagic larval stage capable of extensive dispersal. Under these circumstances, why are there so many species of reef fishes? Since most biogeographic barriers in the oceans are either spatially or temporally permeable on a relatively short time frame, the requirement of isolation during allopatric speciation is hard to satisfy. Evidence from empirical and theoretical studies, the biological characteristics of coral reefs, and a reanalysis of biogeographic barriers indicate that sympatric speciation is possible but not common at small spatial scales and that parapatric speciation is a common (and probably the prevalent) mode of diversification in coral-reef fishes. Regardless of the speciation mode, previous hypotheses of accelerated diversification in the Pleistocene due to sea level fluctuations are not supported by phylogenetic analyses. Recent developments in the area of comparative genomics can fuel a new revolution in the way marine speciation is studied.
Until now, the earliest known members of the triggerfish family Balistidae have been two genera from the Oligocene. Herein
is described the new balistid Gornylistes prodigiosus gen. et sp. nov. from the uppermost Middle Eocene (Kuma Horizon) of the Northern Caucasus (Gorny Luch locality); it is as
thoroughly modern in its bauplan as the taxa of balistids from the Oligocene and more recent periods, and far more advanced
morphologically than the several stem taxa of the balistoid + ostracioid clade known from earlier in the Middle and Lower
Eocene and from the Upper Paleocene.
For most vertebrates, locomotion is a fundamental component of prey capture. Despite this ubiquitous link, few studies have quantified the integration of these complex systems. Several variables related to locomotor performance, including maximum speed, acceleration, deceleration, maneuverability, accuracy, and approach stability, likely influence feeding performance in vertebrates. The relative importance of these measures of performance, however, depends on the ecology of the predator. While factors such as morphology and physiology likely define the limits of these variables, other factors such as motivation of the predator, prey type, and habitat structure can also influence performance. Understanding how these variables relate to feeding under a given suite of ecological conditions is central to understanding predator-prey interactions, and ultimately how locomotion and feeding have co-evolved. The goals of this article are to discuss several variables of locomotor performance related to prey capture, present new data on the relationship between locomotor and feeding morphology in fishes, discuss the evolution of prey capture in cichlid fishes, and outline some future directions for research. While suction feeding is a primary mechanism of prey capture in fishes, swimming is vital for accurately positioning the mouth relative to the prey item. Many fishes decelerate during prey capture using their body and fins, but the pectoral fins have a dominant role in maintaining approach stability. This suggests that fishes employing high-performance suction feeding (relatively small mouth) will have larger pectoral fins to facilitate accurate and stable feeding. I provide new data on the relationship between pectoral fin morphology and maximum gape in centrarchid fishes. For seven species, pectoral fin area was significantly, and negatively, correlated with maximum gape. This example illustrates that the demands from one complex system (feeding) can influence another complex system (locomotion). Future studies that examine the morphological, physiological, and functional evolution of locomotion involved in prey capture by aquatic and terrestrial vertebrates will provide insight into the origin and consequences of diversity.
A common pattern found in phylogeny-based empirical studies of diversification is a decrease in the rate of lineage accumulation toward the present. This early-burst pattern of cladogenesis is often interpreted as a signal of adaptive radiation or density-dependent processes of diversification. However, incomplete taxonomic sampling is also known to artifactually produce patterns of rapid initial diversification. The Monte Carlo constant rates (MCCR) test, based upon Pybus and Harvey's gamma (γ)-statistic, is commonly used to accommodate incomplete sampling, but this test assumes that missing taxa have been randomly pruned from the phylogeny. Here we use simulations to show that preferentially sampling disparate lineages within a clade can produce severely inflated type-I error rates of the MCCR test, especially when taxon sampling drops below 75%. We first propose two corrections for the standard MCCR test, the proportionally deeper splits that assumes missing taxa are more likely to be recently diverged, and the deepest splits only MCCR that assumes that all missing taxa are the youngest lineages in the clade, and assess their statistical properties. We then extend these two tests into a generalized form that allows the degree of nonrandom sampling (NRS)to be controlled by a scaling parameter, α. This generalized test is then applied to two recent studies. This new test allows systematists to account for nonrandom taxonomic sampling when assessing temporal patterns of lineage diversification in empirical trees. Given the dramatic affect NRS can have on the behavior of the MCCR test, we argue that evaluating the sensitivity of this test to NRS should become the norm when investigating patterns of cladogenesis in incompletely sampled phylogenies.
Ecological opportunity--through entry into a new environment, the origin of a key innovation or extinction of antagonists--is widely thought to link ecological population dynamics to evolutionary diversification. The population-level processes arising from ecological opportunity are well documented under the concept of ecological release. However, there is little consensus as to how these processes promote phenotypic diversification, rapid speciation and adaptive radiation. We propose that ecological opportunity could promote adaptive radiation by generating specific changes to the selective regimes acting on natural populations, both by relaxing effective stabilizing selection and by creating conditions that ultimately generate diversifying selection. We assess theoretical and empirical evidence for these effects of ecological opportunity and review emerging phylogenetic approaches that attempt to detect the signature of ecological opportunity across geological time. Finally, we evaluate the evidence for the evolutionary effects of ecological opportunity in the diversification of Caribbean Anolis lizards. Some of the processes that could link ecological opportunity to adaptive radiation are well documented, but others remain unsupported. We suggest that more study is required to characterize the form of natural selection acting on natural populations and to better describe the relationship between ecological opportunity and speciation rates.
Conservation priorities are calculated on the basis of species richness, endemism, and threats. However, areas ranked highly
for these factors may not represent regions of maximal evolutionary potential. The relationship between geography and evolutionary
innovation was analyzed in a dominant complex of Caribbean reef corals, in which morphological and genetic data concur on
species differences. Based on geometric morphometrics of Pleistocene corals and genetically characterized modern colonies,
we found that morphological disparity varies from the center to the edge of the Caribbean, and we show that lineages are static
at well-connected central locations but split or fuse in edge zones where gene flow is limited. Thus, conservation efforts
in corals should focus not only on the centers of diversity but also on peripheral areas of species ranges and population
connectivity.
Studies of diversification patterns often find a slowing in lineage accumulation toward the present. This seemingly pervasive pattern of rate downturns has been taken as evidence for adaptive radiations, density-dependent regulation, and metacommunity species interactions. The significance of rate downturns is evaluated with statistical tests (the gamma statistic and Monte Carlo constant rates (MCCR) test; birth-death likelihood models and Akaike Information Criterion [AIC] scores) that rely on null distributions, which assume that the included species are a random sample of the entire clade. Sampling in real phylogenies, however, often is nonrandom because systematists try to include early-diverging species or representatives of previous intrataxon classifications. We studied the effects of biased sampling, structured sampling, and random sampling by experimentally pruning simulated trees (60 and 150 species) as well as a completely sampled empirical tree (58 species) and then applying the gamma statistic/MCCR test and birth-death likelihood models/AIC scores to assess rate changes. For trees with random species sampling, the true model (i.e., the one fitting the complete phylogenies) could be inferred in most cases. Oversampling deep nodes, however, strongly biases inferences toward downturns, with simulations of structured and biased sampling suggesting that this occurs when sampling percentages drop below 80%. The magnitude of the effect and the sensitivity of diversification rate models is such that a useful rule of thumb may be not to infer rate downturns from real trees unless they have >80% species sampling.
Modern whales are frequently described as an adaptive radiation spurred by either the evolution of various key innovations (such as baleen or echolocation) or ecological opportunity following the demise of archaic whales. Recent analyses of diversification rate shifts on molecular phylogenies raise doubts about this interpretation since they find no evidence of increased speciation rates during the early evolution of modern taxa. However, one of the central predictions of ecological adaptive radiation is rapid phenotypic diversification, and the tempo of phenotypic evolution has yet to be quantified in cetaceans. Using a time-calibrated molecular phylogeny of extant cetaceans and a morphological dataset on size, we find evidence that cetacean lineages partitioned size niches early in the evolutionary history of neocetes and that changes in cetacean size are consistent with shifts in dietary strategy. We conclude that the signature of adaptive radiations may be retained within morphological traits even after equilibrium diversity has been reached and high extinction or fluctuations in net diversification have erased any signature of an early burst of diversification in the structure of the phylogeny.
Many continental sister species are allopatric or parapatric, ecologically similar and long separated, of the order of millions of years. Sympatric, ecologically differentiated, species, are often even older. This raises the question of whether build-up of sympatric diversity generally follows a slow process of divergence in allopatry, initially without much ecological change. I review patterns of speciation among birds belonging to the continental Eurasian Old World leaf warblers (Phylloscopus and Seicercus). I consider speciation to be a three-stage process (range expansions, barriers to gene flow, reproductive isolation) and ask how ecological factors at each stage have contributed to speciation, both among allopatric/parapatric sister species and among those lineages that eventually led to currently sympatric species. I suggest that time is probably the critical factor that leads to reproductive isolation between sympatric species and that a strong connection between ecological divergence and reproductive isolation remains to be established. Besides reproductive isolation, ecological factors can affect range expansions (e.g. habitat tracking) and the formation of barriers (e.g. treeless areas are effective barriers for warblers). Ecological factors may often limit speciation on continents because range expansions are difficult in 'ecologically full' environments.
Major modifications to the pharyngeal jaw apparatus are widely regarded as a recurring evolutionary key innovation that has enabled adaptive radiation in many species-rich clades of percomorph fishes. However one of the central predictions of this hypothesis, that the acquisition of a modified pharyngeal jaw apparatus will be positively correlated with explosive lineage diversification, has never been tested. We applied comparative methods to a new time-calibrated phylogeny of labrid fishes to test whether diversification rates shifted at two scales where major pharyngeal jaw innovations have evolved: across all of Labridae and within the subclade of parrotfishes.
Diversification patterns within early labrids did not reflect rapid initial radiation. Much of modern labrid diversity stems from two recent rapid diversification events; one within julidine fishes and the other with the origin of the most species-rich clade of reef-associated parrotfishes. A secondary pharyngeal jaw innovation was correlated with rapid diversification within the parrotfishes. However diversification rate shifts within parrotfishes are more strongly correlated with the evolution of extreme dichromatism than with pharyngeal jaw modifications.
The temporal lag between pharyngeal jaw modifications and changes in diversification rates casts doubt on the key innovation hypothesis as a simple explanation for much of the richness seen in labrids and scarines. Although the possession of a secondarily modified PJA was correlated with increased diversification rates, this pattern is better explained by the evolution of extreme dichromatism (and other social and behavioral characters relating to sexual selection) within Scarus and Chlorurus. The PJA-innovation hypothesis also fails to explain the most dominant aspect of labrid lineage diversification, the radiation of the julidines. We suggest that pharyngeal jaws might have played a more important role in enabling morphological evolution of the feeding apparatus in labrids and scarines rather than in accelerating lineage diversification.
One of the main explanations for the stunning diversity of teleost fishes (approximately 29,000 species, nearly half of all vertebrates) is that a fish-specific whole-genome duplication event (FSGD) in the ancestor to teleosts triggered their subsequent radiation. However, one critical assumption of this hypothesis, that diversification rates in teleosts increased soon after the acquisition of a duplicated genome, has never been tested.
Here we show that one of three major diversification rate shifts within ray-finned fishes occurred at the base of the teleost radiation, as predicted by the FSGD hypothesis. We also find evidence for two rate increases that are much younger than the inferred age of the FSGD: one in the common ancestor of most ostariophysan fishes, and a second one in the common ancestor of percomorphs. The biodiversity contained within these two clades accounts for more than 88% of living fish species.
Teleosts diversified explosively in their early history and this burst of diversification may have been caused by genome duplication. However, the FSGD itself may be responsible for a little over 10% of living teleost biodiversity. ~88% of species diversity is derived from two relatively recent radiations of freshwater and marine fishes where genome duplication is not suspected. Genome duplications are a common event on the tree of life and have been implicated in the diversification of major clades like flowering plants, vertebrates, and gnathostomes. However our results suggest that the causes of diversification in large clades are likely to be complex and not easily ascribed to a single event, even a dramatic one such as a whole genome duplication.
The uneven distribution of species richness is a fundamental and unexplained pattern of vertebrate biodiversity. Although species richness in groups like mammals, birds, or teleost fishes is often attributed to accelerated cladogenesis, we lack a quantitative conceptual framework for identifying and comparing the exceptional changes of tempo in vertebrate evolutionary history. We develop MEDUSA, a stepwise approach based upon the Akaike information criterion for detecting multiple shifts in birth and death rates on an incompletely resolved phylogeny. We apply MEDUSA incompletely to a diversity tree summarizing both evolutionary relationships and species richness of 44 major clades of jawed vertebrates. We identify 9 major changes in the tempo of gnathostome diversification; the most significant of these lies at the base of a clade that includes most of the coral-reef associated fishes as well as cichlids and perches. Rate increases also underlie several well recognized tetrapod radiations, including most modern birds, lizards and snakes, ostariophysan fishes, and most eutherian mammals. In addition, we find that large sections of the vertebrate tree exhibit nearly equal rates of origination and extinction, providing some of the first evidence from molecular data for the importance of faunal turnover in shaping biodiversity. Together, these results reveal living vertebrate biodiversity to be the product of volatile turnover punctuated by 6 accelerations responsible for >85% of all species as well as 3 slowdowns that have produced "living fossils." In addition, by revealing the timing of the exceptional pulses of vertebrate diversification as well as the clades that experience them, our diversity tree provides a framework for evaluating particular causal hypotheses of vertebrate radiations.
Biologists have long been fascinated by the exceptionally high diversity displayed by some evolutionary groups. Adaptive radiation in such clades is not only spectacular, but is also an extremely complex process influenced by a variety of ecological, genetic, and developmental factors and strongly dependent on historical contingencies. Using modeling approaches, we identify 10 general patterns concerning the temporal, spatial, and genetic/morphological properties of adaptive radiation. Some of these are strongly supported by empirical work, whereas for others, empirical support is more tentative. In almost all cases, more data are needed. Future progress in our understanding of adaptive radiation will be most successful if theoretical and empirical approaches are integrated, as has happened in other areas of evolutionary biology.
Although substantial uncertainty typically surrounds the choice of the best model in most phylogenetic analyses, little is known about how accommodating this uncertainty affects phylogenetic inference. Here we explore the influence of Bayesian model averaging on the phylogenetic inference of the triggerfishes (Family: Balistidae), a charismatic group of reef fishes. We focus on clade support as this area has received little attention and is typically one of the most important outcomes of phylogenetic studies. We present a novel phylogenetic hypothesis for the family Balistidae based on an analysis of two mitochondrial (12S, 16S) and three nuclear genes (TMO-4C4, Rhodopsin, RAG1) sampled from 26 ingroup species. Despite the presence of substantial model uncertainty in almost all partitions of our data, we found model-averaged topologies and clade posteriors to be nearly identical to those conditioned on a single model. Furthermore, statistical comparison of clade posteriors using the Wilcoxon signed-rank test revealed no significant differences. Our results suggest that although current model-selection approaches are likely to lead to overparameterization of the substitution model, the consequences of conditioning on this overparameterized model are likely to be mild. Our phylogenetic results strongly support the monophyly of the triggerfishes but suggest that the genera Balistoides and Pseudobalistes are polyphyletic. Divergence time estimation supports a Miocene origin of the crown group. Despite the presence of several young species-rich subclades, statistical analysis of temporal diversification patterns reveals no significant increase in the rate of cladogenesis across geologic time intervals.
Labriform locomotion, which is powered by oscillating the paired pectoral fins, varies along a continuum from rowing the fins back and forth to flapping the fins up and down. It has generally been assumed (i) that flapping is more mechanically efficient than rowing, a hypothesis confirmed by a recent simulation experiment, and (ii) that flapping should be associated with wing-shaped fins while rowing should be associated with paddle-shaped fins. To determine whether these hypotheses and the results of the simulation experiment are consistent with natural variation, we compared the steady swimming performance (critical swimming speed) of four species of labrid fish (Cirrhilabrus rubripinnis, Pseudocheilinus octotaenia, Gomphosus varius and Halichoeres bivittatus) selected to form two pairs of closely related species that vary in fin shape and in the direction of fin motion. The results were consistent with expectations. Within each pair, the species with the best swimming performance also had (i) a fin shape characterized by a higher aspect ratio, a longer leading edge relative to the trailing edge fin rays and the center of fin area located closer to the fin base, and (ii) a steeper (more dorsoventral) stroke plane.
Geophysical data are currently being interpreted as evidence for a late Pleistocene desiccation of Lake Victoria and its refilling 14,600 years ago. This implies that between 500 and 1000 endemic cichlid fish species must have evolved in 14,600 years, the fastest large-scale species radiation known. A recent review concludes that biological evidence clearly rejects the postulated Pleistocene desiccation of the lake: a 14,600 year history would imply exceptionally high speciation rates across a range of unrelated fish taxa. To test this suggestion, I calculated speciation rates for all 41 phylogenetic lineages of fish in the lake. Except for one cichlid lineage, accepting a 14 600 year history does not require any speciation rates that fall outside the range observed in fishes in other young lakes around the world. The exceptional taxon is a lineage of haplochromine cichlids that is also known for its rapid speciation elsewhere. Moreover, since it is unknown how many founding species it has, it is not certain that its speciation rates are really outside the range observed in fishes in other young lakes. Fish speciation rates are generally faster in younger than in older lakes, and those in Lake Victoria, by far the largest of the young lakes of the world, are no exception. From the speciation rates and from biogeographical observations that Lake Victoria endemics, which lack close relatives within the lake basin, have such relatives in adjacent drainage systems that may have had Holocene connections to Lake Victoria, I conclude that the composition of the fish assemblage does not provide biological evidence against Pleistocene desiccation. It supports a hypothesis of recent colonization from outside the lake basin rather than survival of a diverse assemblage within the basin.
To determine the energetic costs of rigid-body, median or paired-fin (MPF) swimming versus undulatory, body-caudal fin (BCF) swimming, we measured oxygen consumption as a function of swimming speed in two MPF swimming specialists, Schlegel's parrotfish and Picasso triggerfish. The parrotfish swam exclusively with the pectoral fins at prolonged swimming speeds up to 3.2 total lengths per second (L s(-1); 30 min critical swimming speed, U(crit)). At higher speeds, gait transferred to a burst-and-coast BCF swimming mode that resulted in rapid fatigue. The triggerfish swam using undulations of the soft dorsal and anal fins up to 1.5 L s(-1), beyond which BCF undulations were recruited intermittently. BCF swimming was used continuously above 3.5 L s(-1), and was accompanied by synchronous undulations of the dorsal and anal fins. The triggerfish were capable of high, prolonged swimming speeds of up to 4.1 L s(-1) (30 min U(crit)). In both species, the rates of increase in oxygen consumption with swimming speed were higher during BCF swimming than during rigid-body MPF swimming. Our results indicate that, for these species, undulatory swimming is energetically more costly than rigid-body swimming, and therefore support the hypothesis that MPF swimming is more efficient. In addition, use of the BCF gait at higher swimming speed increased the cost of transport in both species beyond that predicted for MPF swimming at the same speeds. This suggests that, unlike for terrestrial locomotion, gait transition in fishes does not occur to reduce energetic costs, but to increase recruitable muscle mass and propulsive surfaces. The appropriate use of the power and exponential functions to model swimming energetics is also discussed.
Shaping Primate Evolution is an edited collection of papers about how biological form is described in primate biology, and the consequences of form for function and behavior. The contributors are highly regarded internationally recognized scholars in the field of quantitative primate evolutionary morphology. Each chapter elaborates upon the analysis of the form-function-behavior triad in a unique and compelling way. This book is distinctive not only in the diversity of the topics discussed, but also in the range of levels of biological organization that are addressed from cellular morphometrics to the evolution of primate ecology. The book is dedicated to Charles E. Oxnard, whose influential pioneering work on innovative metric and analytic techniques has gone hand-in-hand with meticulous comparative functional analyses of primate anatomy. Through the marriage of theory with analytical applications, this volume will be an important reference work for all those interested in primate functional morphology.
Phylogenies reconstructed from gene sequences can be used to investigate the tempo and mode of species diversification. Here we develop and use new statistical methods to infer past patterns of speciation and extinction from molecular phylogenies. Specifically, we test the null hypothesis that per-lineage speciation and extinction rates have remained constant through time. Rejection of this hypothesis may provide evidence for evolutionary events such as adaptive radiations or key adaptations. In contrast to previous approaches, our methods are robust to incomplete taxon sampling and are conservative with respect to extinction. Using simulation we investigate, first, the adverse effects of failing to take incomplete sampling into account and, second, the power and reliability of our tests. When applied to published phylogenies our tests suggest that, in some cases, speciation rates have decreased through time.
Adaptive radiation, which results when a single ancestral species gives rise to many descendants, each adapted to a different part of the environment, is possibly the single most important source of biological diversity in the living world. One of the best-studied examples involves Caribbean Anolis lizards. With about 400 species, Anolis has played an important role in the development of ecological theory and has become a model system exemplifying the integration of ecological, evolutionary, and behavioral studies to understand evolutionary diversification. This major work, written by one of the best-known investigators of Anolis, reviews and synthesizes an immense literature. Jonathan B. Losos illustrates how different scientific approaches to the questions of adaptation and diversification can be integrated and examines evolutionary and ecological questions of interest to a broad range of biologists.
George Gaylord Simpson famously postulated that much of life’s diversity originated as adaptive radiations—more or less simulta- neous divergences of numerous lines from a single ancestral adaptive type. However, identifying adaptive radiations has proven difficult due to a lack of broad-scale comparative datasets. Here, we use phylogenetic comparative data on body size and shape in a diversity of animal clades to test a key model of adaptive radiation, in which initially rapid morphological evolution is followed by relative stasis. We compared the fit of this model to both single selective peak and random walk models. We found little support for the early-burst model of adaptive radiation, whereas both other models, particularly that of selective peaks, were commonly supported. In addition, we found that the net rate of morphological evolution varied inversely with clade age. The youngest clades appear to evolve most rapidly because long-term change typically does not attain the amount of divergence predicted from rates measured over short time scales. Across our entire analysis, the dominant pattern was one of constraints shaping evolution continually through time rather than rapid evolution followed by stasis. We suggest that the classical model of adaptive radiation, where morphological evolution is initially rapid and slows through time, may be rare in comparative data.
The evolutionary pattern of speciation and extinction in any biologic group may be described by a variety of mathematical models. These models provide a framework for describing the history of taxonomic diversity (clade shape) and other aspects of large evolutionary patterns. The simplest model assumes time homogeneity; that is, speciation and extinction probabilities are constant through time and within taxonomic groups. In some cases, the homogeneous model provides a good fit to real world paleontological data, but in other cases the model serves only as a null hypothesis that must be rejected before more complex models can be applied. In cases where the homogeneous model does not fit the data, time-inhomogeneous models can be formulated that specify change, regular or episodic, in speciation and extinction probabilities. An appendix provides a list of the most useful equations based on the homogeneous model.-Author
A major challenge in evolutionary biology lies in explaining patterns of high species numbers found in biodiversity hot spots. Tropical coral reefs underlie most marine hot spots and reef-associated fish faunas represent some of the most diverse assemblages of vertebrates on the planet. Although the standing diversity of modern reef fish clades is usually attributed to their ecological association with corals, untangling temporal patterns of codiversification has traditionally proved difficult. In addition, owing to uncertainty in higher-level relationships among acanthomorph fish, there have been few opportunities to test the assumption that reef-association itself leads to higher rates of diversification compared to other habitats. Here we use relaxed-clock methods in conjunction with statistical measures of species accumulation and phylogenetic comparative methods to clarify the temporal pattern of diversification in reef and nonreef-associated lineages of tetraodontiforms, a morphologically diverse order of teleost fish. We incorporate 11 fossil calibrations distributed across the tetraodontiform tree to infer divergence times and compare results from models of autocorrelated and uncorrelated evolutionary rates. All major tetraodontiform reef crown groups have significantly higher rates of diversification than the order as a whole. None of the nonreef-associated families show this pattern with the exception of the aracanid boxfish. Independent contrasts analysis also reveals a significantly positive relationship between diversification rate and proportion of reef-associated species within each family when aracanids are excluded. Reef association appears to have increased diversification rate within tetraodontiforms. We suggest that both intrinsic factors of reef habitat and extrinsic factors relating to the provincialization and regionalization of the marine biota during the Miocene (about 23-5 MY) played a role in shaping these patterns of diversity
Morphometrics is the statistical study of biological shape and shape change. Its richest data are landmarks, points, such as the bridge of the nose, that have biological names as well as geometric locations. This book is the first systematic survey of morphometric methods for landmark data.
In marine species, high dispersal is often associated with only mild genetic differentiation over large spatial scales. Despite this generalization, there are numerous reasons for the accumulation of genetic differences between,large, semi-isolated marine populations. A suite of well-known evolutionary mechanisms can operate within and between populations to result in genetic divergence, and these mechanisms may well be augmented by newly discovered genetic processes. This variety of mechanisms for genetic divergence is paralleled by great diversity in the types of reproductive isolation shown by recently diverged marine species. Differences in spawning time, mate recognition, environmental tolerance, and gamete compatibility have all been implicated in marine speciation events. There is substantial evidence for rapid evolution of reproductive isolation in strictly allopatric populations (e.g. across the Isthmus of Panama). Evidence for the action of selection in increasing reproductive isolation in sympatric populations is fragmentary. Although a great deal of information is available on population genetics, reproductive isolation, and cryptic or sibling species in marine environments, the influence of particular genetic changes on reproductive isolation is poorly understood for marine (or terrestrial) taxa. For a few systems, like the co-evolution of gamete recognition proteins, changes in a small number of genes may give rise to reproductive isolation. Such studies show how a focus on the physiology, ecology, or sensory biology of reproductive isolation can help uncover the genetic changes associated with speciation and can also help provide a link between the genetics of population divergence and the speciation process.
▪ Abstract The diversity of organismic form has evolved nonuniformly during the history of life. Quantitative morphological studies reveal profound changes in evolutionary rates corresponding with the generation of morphological disparity at low taxonomic diversity during the early radiation of many clades. These studies have also given insight into the relative importance of genomic and ecological factors in macroevolution, the selectivity of extinction, and other issues. Important progress has been made in the development of morphological spaces that can accommodate highly disparate forms, although this area still needs more attention. Other future directions include the relationship between morphological and ecological diversification, geographic patterns in morphological diversity, and the role of morphological disparity as a causal factor in macroevolution.
Elongated-body theory has been fruitfully applied over twenty years to the biofluiddynamic analysis of modes of locomotion of elongated fishes by means of body flexure, with special emphasis on the anguilliform mode using undulatory body movements, and on the carangiform mode where oscillatory movements of only a fish's posterior end(including the caudal fin) exhibit phase lag of posterior movements behind anterior movements just as in an undulation yet not nearly as much as a whole wavelength is apparent at any one time. The extension of elongated-body theory to analyse the locomotion of elongated fishes with elongated median fins (dorsal and/or anal) in modes where the body (together with any caudal fin) remains rigid, being propelled forwards by undulations or oscillations of those median fins, has long been recognized as desirable but is here presented for the first time.
Heterochrony, evolutionary changes in rate or timing of development producing parallelism between ontogeny and phylogeny, is viewed as the most common type of evolutionary change in development. Alternative hypotheses such as heterotopy, evolutionary change in the spatial patterning of development, are rarely entertained. We examine the evidence for heterochrony and heterotopy in the evolution of body shape in two clades of piranhas. One of these is the sole case of heterochrony previously reported in the group; the others were previously interpreted as cases of heterotopy. To compare ontogenies of shape, we computed ontogenetic trajectories of shape by multivariate regression of geometric shape variables (i.e., partial warp scores and shape coordinates) on centroid size. Rates of development relative to developmental age and angles between the trajectories were compared statistically. We found a significant difference in developmental rate between species of Serrasalmus, suggesting that heterochrony is a partial explanation for the evolution of body shape, but we also found a significant difference between their ontogenetic transformations; the direction of the difference between them suggests that heterotopy also plays a role in this group. In Pygocentrus we found no difference in developmental rate among species, but we did find a difference in the ontogenies, suggesting that heterotopy, but not heterochrony, is the developmental basis for shape diversification in this group. The prevalence of heterotopy as a source of evolutionary novelty remains largely unexplored and will not become clear until the search for developmental explanations looks beyond heterochrony.
Over the past 20 years, experimental analyses of the biomechanics of locomotion in fishes have generated a number of key findings
that are relevant to the construction of biomimetic fish robots. In this paper, we present 16 results from recent experimental
research on the mechanics, kinematics, fluid dynamics, and control of fish locomotion that summarize recent work on fish biomechanics.
The findings and principles that have emerged from biomechanical studies of fish locomotion provide important insights into
the functional design of fishes and suggest specific design features relevant to construction of robotic fish-inspired vehicles
that underlie the high locomotor performance exhibited by fishes.
Time-calibrated molecular phylogenies provide a valuable window into the tempo and mode of species diversification, especially for the large number of groups that lack adequate fossil records. Molecular phylogenetic data frequently suggest an initial "explosive speciation" phase, leading to widespread speculation that ecological niche-filling processes might govern the dynamics of species diversification during evolutionary radiations. However, these patterns are difficult to reconcile with the fossil record. The fossil record strongly suggests that extinction rates have been high relative to speciation rates, but such elevated background extinction should erase the signal of early, rapid speciation from molecular phylogenies. For this reason, extinction rates in molecular phylogenies are frequently estimated as zero under the widely used birth-death model. Here, I construct a simple model that combines phylogenetically patterned extinction with pulsed turnover dynamics and constant diversity through time. Using approximate Bayesian methods, I show that heritable extinction can easily explain the phenomenon of explosive early diversification, even when net diversification rates are equal to zero. Several assumptions of the model are more consistent with both the fossil record and neontological data than the standard birth-death model and it may thus represent a viable alternative interpretation of phylogenetic diversification patterns. These results suggest that variation in the absolute rate of lineage turnover through time, in conjunction with phylogenetically nonrandom extinction, may underlie the apparent diversity-dependent speciation observed in molecular phylogenies.
Most studies of adaptive radiations focus on morphological aspects of differentiation, yet behavior is also an important component of evolutionary diversification, often mediating the relationship between animal ecology and morphology. In species within radiations that are convergent in ecology and morphology, we then also expect convergence in behavior. Here, we examined 13 Anolis lizard species to determine whether territorial strategies have evolved convergently with morphology and habitat use. We evaluated two aspects of territoriality: behavioral defense of space via territorial displays, and territory overlap within and between sexes. Controlling for the phylogenetic relationships of the taxa in our study, we found that species similar in perch height and diameter convergently evolved patterns of territory overlap, whereas species similar in habitat visibility (the proportion of space that can be seen from a perch) convergently evolved display behavior. We also found that species with greater display time have more extensive male-male territory overlap. This study provides strong evidence for the role of habitat in the evolution of territoriality and suggests that the social structure of a species ultimately evolves in concert with habitat use and morphology.
I review the concept of optimal vortex formation and examine its relevance to propulsion in biological and bio-inspired systems, ranging from the human heart to underwater vehicles. By using examples from the existing literature and new analyses, I show that optimal vortex formation can potentially serve as a unifying principle to understand the diversity of solutions used to achieve propulsion in nature. Additionally, optimal vortex formation can provide a framework in which to design engineered propulsions systems that are constrained by pressures unrelated to biology. Finally, I analyze the relationship between optimal vortex formation and previously observed constraints on Strouhal frequency during animal locomotion in air and water. It is proposed that the Strouhal frequency constraint is but one consequence of the process of optimal vortex formation and that others remain to be discovered.
Recent application of time-varying birth-death models to molecular phylogenies suggests that a decreasing diversification rate can only be observed if there was a decreasing speciation rate coupled with extremely low or no extinction. However, from a paleontological perspective, zero extinction rates during evolutionary radiations seem unlikely. Here, with a more comprehensive set of computer simulations, we show that substantial extinction can occur without erasing the signal of decreasing diversification rate in a molecular phylogeny. We also find, in agreement with the previous work, that a decrease in diversification rate cannot be observed in a molecular phylogeny with an increasing extinction rate alone. Further, we find that the ability to observe decreasing diversification rates in molecular phylogenies is controlled (in part) by the ratio of the initial speciation rate (Lambda) to the extinction rate (Mu) at equilibrium (the LiMe ratio), and not by their absolute values. Here we show in principle, how estimates of initial speciation rates may be calculated using both the fossil record and the shape of lineage through time plots derived from molecular phylogenies. This is important because the fossil record provides more reliable estimates of equilibrium extinction rates than initial speciation rates.
Balistoid fishes have a unique and reduced pelvic fin structure, which does not exhibit paired structures. The pelvic complex exhibits reductive trends, but its rudimentary structure was retained among balistoids, and its unidirectional and parsimonious reduction in more derived lineages has been hypothesized based on morphology. We investigated the evolution of pelvic complex reduction in balistoids using whole mitochondrial genome (mitogenome) data from 33 species (27 newly determined during the study) that represent the entire morphological diversity of balistoids. Partitioned maximum likelihood and Bayesian analyses were conducted with two datasets that comprised concatenated nucleotide sequences from 13 protein-coding genes (all positions included; third codon positions converted into purine [R] and pyrimidine [Y] [RY-coding]) plus 22 transfer RNA and two ribosomal RNA genes. The resultant trees were well resolved and largely congruent, with most internal branches having high support values. The mitogenomic datasets strongly supported monophylies of both balistids and monacanthids, but rejected previous hypotheses on the intra-relationships in each family. The present tree topology revealed that highly reduced pelvic complexes had multiple origins, and optimization of the traits on the resultant tree strongly suggested the non-unidirectional and independent reduction of pelvic complexes in balistoids. The evolution of balistoid pelvic structure is very different among fishes that exhibit its reductive trends, and this uniqueness in pelvic evolution may be a link to their reproductive behaviors.
Lake Baikal in Eastern Siberia contains a remarkable flock of 29 species of teleost fishes of the suborder Cottoidei (sculpins, bullheads) that are endemic to the lake and its associated rivers and occupy all depth habitats down to over 1500 m. The species are divided into three families, the Cottidae with 7 species, the Abyssocottidae with 20 species, and the Comephoridae with 2 species. Nucleotide sequences of the rod opsin gene from 12 of these species, plus a non-Baikal marine species, have been used to examine the evolutionary relations and the divergence time of the flock. Phylogenetic trees, generated by neighbor-joining and maximum parsimony, indicate that the unique Comephoridae family with its viviparity and unusual appearance is closely related to the Cottidae and Abyssocottidae, whereas the genus Cottocomephorus, at present placed in the Cottidae, was the first to diverge from the ancestral species and forms a separate lineage. The major adaptation to deep water would appear to be of relatively recent origin, and there is evidence that the ancestral species occupied a shallow-water-marine or brackish habitat. Estimates of antiquity obtained from synonymous substitutions place the origin of the species flock at around 4.9 million years ago.