Brian L. Sidlauskas

Early bursts of body size and shape evolution are rare in comparative data.

George Gaylord Simpson famously postulated that much of lifes diversity originated as adaptive radiations—more or less simultaneous 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.

Continuous and arrested morphological diversification in sister clades of characiform fishes: a phylomorphospace approach.

Abstract Understanding how and why certain clades diversify greatly in morphology whereas others do not remains a major theme in evolutionary biology. Projecting families of phylogenies into multivariate morphospaces can distinguish two scenarios potentially leading to unequal morphological diversification: unequal magnitude of change per phylogenetic branch, and unequal efficiency in morphological innovation. This approach is demonstrated using a case study of skulls in sister clades within the South American fish superfamily Anostomoidea. Unequal morphological diversification in this system resulted not from the morphologically diverse clade changing more on each phylogenetic branch, but from that clade distributing an equal amount of change more widely through morphospace and innovating continually. Although substantial morphological evolution occurred throughout the less diverse clades history, most of that clades expansion in morphospace occurred in the most basal branches, and more derived portions of that radiation oscillated within previously explored limits. Because simulations revealed that there is a maximum 2.7% probability of generating two clades that differ so greatly in the density of lineages within morphospace under a null Brownian model, the observed difference in pattern likely reflects a difference in the underlying evolutionary process. Clade–specific factors that may have promoted or arrested morphological diversification are discussed.

THE INFLUENCE OF AN INNOVATIVE LOCOMOTOR STRATEGY ON THE PHENOTYPIC DIVERSIFICATION OF TRIGGERFISH (FAMILY: BALISTIDAE)

Innovations in locomotor morphology have been invoked as important drivers of vertebrate diversification, although the influence of novel locomotion strategies on marine fish diversification remains largely unexplored. Using triggerfish as a case study, we determine whether the evolution of the distinctive synchronization of enlarged dorsal and anal fins that triggerfish use to swim may have catalyzed the ecological diversification of the group. By adopting a comparative phylogenetic approach to quantify median fin and body shape integration and to assess the tempo of functional and morphological evolution in locomotor traits, we find that: (1) functional and morphological components of the locomotive system exhibit a strong signal of correlated evolution; (2) triggerfish partitioned locomotor morphological and functional spaces early in their history; and (3) there is no strong evidence that a pulse of lineage diversification accompanied the major episode of phenotypic diversification. Together these findings suggest that the acquisition of a distinctive mode of locomotion drove an early radiation of shape and function in triggerfish, but not an early radiation of species.

Linking big: the continuing promise of evolutionary synthesis.

Synthetic science promises an unparalleled ability to find new meaning in old data, extant results, or previously unconnected methods and concepts, but pursuing synthesis can be a difficult and risky endeavor. Our experience as biologists, informaticians, and educators at the National Evolutionary Synthesis Center has affirmed that synthesis can yield major insights, but also revealed that technological hurdles, prevailing academic culture, and general confusion about the nature of synthesis can hamper its progress. By presenting our view of what synthesis is, why it will continue to drive progress in evolutionary biology, and how to remove barriers to its progress, we provide a map to a future in which all scientists can engage productively in synthetic research.

TESTING FOR UNEQUAL RATES OF MORPHOLOGICAL DIVERSIFICATION IN THE ABSENCE OF A DETAILED PHYLOGENY: A CASE STUDY FROM CHARACIFORM FISHES

Abstract This study develops the random phylogenies rate test (RAPRATE), a likelihood method that simulates morphological evolution along randomly generated phylogenies, and uses it to determine whether a considerable difference in morphological diversity between two sister clades of South American fishes should be taken as evidence of differing rates of morphological change or lineage turnover. Despite identical ages of origin, similar species richness, and sympatric geographic distributions, the morphological and ecological diversity of the superfamily Anostomoidea exceeds that of the Curimatoidea. The test shows with 90% confidence (using variance among species as the measure of morphological diversity) or 99% confidence (using volume of occupied morphospace) that the rate of morphological change per unit time in the Anostomoidea likely exceeded that of the Curimatoidea. Variation in the rate of lineage turnover (speciation and extinction rates) is not found to affect greatly the morphological diversity of simulated clades and is not a likely explanation of the observed difference in morphological diversity in this case study. Though a 17% or greater delay in the onset of diversification in the Curimatoidea remains a possible alternative explanation of unequal morphological diversification, further simulations suggest that two clades drawn from the possible treespace of the Anostomoidea and Curimatoidea will rarely differ so greatly in the onset of diversification. Several uniquely derived morphological and ecological features of the Anostomoidea and Curimatoidea may have accelerated or decelerated their rate of morphological change, including a marked lengthening of the quadrate that may have relaxed structural constraints on the evolution of the anostomoid jaw.

Linked morphological changes during palate evolution in early tetrapods

We examined the shapes and sizes of dermal bones of the palate of selected Palaeozoic tetrapods in order to identify the ancestral states of palatal bone morphologies in the earliest tetrapods, to learn how the composition of the palate varies within and among early tetrapod radiations, and to recognize evolutionary correlations among the size and shapes of skeletal elements in this important group of animals. We find that whereas the palatal bones themselves and their arrangements are usually conserved, considerable correlated evolutionary change occurs in the shapes and sizes of the bones. Some of the changes in the bones are allometrically linked to overall palate size, which varies more than 100‐fold among the taxa in our sample. Often, these allometries were only hinted at in traditional independent contrasts‐based regressions of log transformed data, particularly because many allometries are subtle, their slopes may vary among subclades, and the scatter around some trendlines is high. Rather, the allometries showed up in analyses of size‐standardized palatal bone dimensions investigated using independent contrasts, bivariate phylomorphospace plots, and mirrored character reconstructions on the phylogenetic tree. We find negative allometry for parasphenoid lengths and widths essentially across the entire tree of Palaeozoic tetrapods, but with different trajectories characterizing the two largest clades, the temnospondyls and the lepospondyls. The lengths of several other elements may show positive allometries, either across the entire tree or in just a subclade. One possible positive allometry exists for the ectopterygoid, which appears to shorten allometrically in temnospondyls that evolve small body and palate size, and, as in Doleserpeton can be lost altogether. Both shortening and loss could be by the same developmental change, paedomorphosis, a form of heterochrony. Paedomorphosis might also account for evolution of relatively large parasphenoids in both lepospondyls and diminutive temnospondyls, but does not seem to explain evolution of ectopterygoid loss in lepospondyls. A regularity observed across nearly all taxa in our study set is an inverse correlation between the lengths of the vomer and pterygoid, bones that lie adjacent to one another along the long palatal axis. Further work is needed to learn whether such correlated evolution might be due to adaptation and/or to developmental bias, and particularly to learn how correlations and allometries themselves evolve.

Phylotastic! Making tree-of-life knowledge accessible, reusable and convenient

BackgroundScientists rarely reuse expert knowledge of phylogeny, in spite of years ofeffort to assemble a great “Tree of Life” (ToL). A notableexception involves the use of Phylomatic, which provides tools togenerate custom phylogenies from a large, pre-computed, expert phylogeny ofplant taxa. This suggests great potential for a more generalized systemthat, starting with a query consisting of a list of any known species, wouldrectify non-standard names, identify expert phylogenies containing theimplicated taxa, prune away unneeded parts, and supply branch lengths andannotations, resulting in a custom phylogeny suited to the user’sneeds. Such a system could become a sustainable community resource ifimplemented as a distributed system of loosely coupled parts that interactthrough clearly defined interfaces.ResultsWith the aim of building such a “phylotastic” system,the NESCent Hackathons, Interoperability, Phylogenies (HIP) workinggroup recruited 2 dozen scientist-programmers to a weeklong programminghackathon in June 2012. During the hackathon (and a three-month follow-upperiod), 5 teams produced designs, implementations, documentation,presentations, and tests including: (1) a generalized scheme for integratingcomponents; (2) proof-of-concept pruners and controllers; (3) a meta-API fortaxonomic name resolution services; (4) a system for storing, finding, andretrieving phylogenies using semantic web technologies for data exchange,storage, and querying; (5) an innovative new service, DateLife.org,which synthesizes pre-computed, time-calibrated phylogenies to assign agesto nodes; and (6) demonstration projects. These outcomes are accessible viaa public code repository (GitHub.com), a website(http://www.phylotastic.org), and a server image.ConclusionsApproximately 9 person-months of effort (centered on a software developmenthackathon) resulted in the design and implementation of proof-of-conceptsoftware for 4 core phylotastic components, 3 controllers, and 3 end-userdemonstration tools. While these products have substantial limitations, theysuggest considerable potential for a distributed system that makesphylogenetic knowledge readily accessible in computable form. Widespread useof phylotastic systems will create an electronic marketplace for sharingphylogenetic knowledge that will spur innovation in other areas of the ToLenterprise, such as annotation of sources and methods and third-partymethods of quality assessment.

Molecular phylogenetics of the Neotropical fish family Prochilodontidae (Teleostei: Characiformes).

Migratory detritivores of the characiform family Prochilodontidae occur throughout the freshwaters of much of South America. Prochilodontids often form massive populations and many species achieve substantial body sizes; a combination that makes them one of the most commercially important fish groups on the continent. Their economic significance notwithstanding, prochilodontids have never been the subject of a comprehensive molecular phylogenetic analysis. Using three mitochondrial and three nuclear loci spanning all prochilodontid species, we generated a novel phylogenetic hypothesis for the family. Our results strongly support monophyly of the family and the three included genera. A novel, highly supported placement of Ichthyoelephas sister to the clade containing Prochilodus and Semaprochilodus diverges from a previous morphological hypothesis. Most previously hypothesized interspecific relationships are corroborated and some longstanding polytomies within Prochilodus and Semaprochilodus are resolved. The morphologically similar P. brevis, P. lacustris, P. nigricans and P. rubrotaeniatus are embedded within what is herein designated as the P. nigricans group. Species limits and distributions of these species are problematic and the group clearly merits taxonomic revision.

The first molecular phylogeny of Chilodontidae (Teleostei: Ostariophysi: Characiformes) reveals cryptic biodiversity and taxonomic uncertainty

Chilodontidae is a small family of eight described characiform species popularly known as headstanders. These small to moderately sized fishes are well known to aquarists, who prize their striking spotted pigmentation and unusual behaviors, and to systematists, who have revised both chilodontid genera in recent memory and studied their phylogenetic relationships using a comprehensive morphological dataset. However, no molecular phylogeny for the family has ever been proposed. Here, we reconstruct phylogenetic relationships for all eight known chilodontid species using three mitochondrial and two nuclear loci. Results largely agree with the previous morphological hypothesis, and confirm the monophyly of the family as well as its included genera, Caenotropus and Chilodus. The molecular topology differs slightly from the morphological hypothesis by placing Caenotropus maculosus rather than C. mestomorgmatos as the sister to the remaining three congeners, and by reconstructing the Curimatidae as the closest outgroup family, rather than the Anostomidae. However, the topologies supported by the morphological data were only slightly less likely and could not be rejected via Shimodaira-Hasegawa tests. Within Chilodus, two described species with distinctive pigmentation (C. fritillus and C. zunevei) appear embedded within the broad distributed C. punctatus clade, suggesting the presence of cryptic taxa with polymorphic pigmentation within the present concept of C. punctatus. Future work should combine morphological and molecular data to revisit the taxonomy and systematics of Chilodus and determine species limits within the C. punctatus-group sensu lato.

A morphological supermatrix-based phylogeny for the Neotropical fish superfamily Anostomoidea (Ostariophysi: Characiformes): phylogeny, missing data and homoplasy

Although 11 studies have addressed the systematics of the four families and 281 fish species of the ecomorphologically diverse Anostomoidea, none has proposed a global hypothesis of relationships. We synthesized these studies to yield a supermatrix with 463 morphological characters among 174 ingroup species, and inferred phylogeny with parsimony and Bayesian optimization. We evaluated the applicability of the supermatrix approach to morphological datasets, tested its sensitivity to missing data, determined the impact of homoplastic characters on phylogenetic resolution, and determined the distribution of homologies and homoplasies on the topology. Despite more than 60% missing data, analyses supported the monophyly of all families, and phylogenetic structure degraded only with inclusion of species with high percentages of missing data and in analyses limited to homoplasies. The latter differs modestly from the full matrix indicating phylogenetic signal in homoplastic characters. Character distributions differ across the phylogeny, with a greater prevalence of homologies at deeper nodes and homoplasies nearer the tips than expected by chance. This may suggest early diversification into distinct bauplans with subsequent diversification of faster evolving character systems. The morphological supermatrix approach is powerful and allows integration of classical data with modern methods to examine the evolution of multiple character systems.