Rhynchonellida is the stratigraphically oldest and phylogenetically most basal of the extant rhynchonelliform brachiopod orders, yet phylogenetic relationships among rhynchonellides are poorly known. The fourteen named rhynchonellide superfamilies (four of which have extant representatives) were defined primarily on the basis of features of the dorsal cardinalia, particularly crural morphology, but their homology and polarity have not been investigated rigorously. Superfamily monophyly is unclear, as is the evolution of several distinctive rhynchonellide morphological features, such as crura.The purpose of this study is to investigate the phylogenetic relationships among extant rhynchonellide genera using skeletal characters, and to compare the results with the current classification, elucidating the evolution of morphological features in the process. We completed parsimony-based and Bayesian analyses using fifty-eight characters of the interior and exterior of the shell that vary among the nineteen extant genera. Our results are readily interpretable with respect to the classification, and indicate that Hemithiridoidea, Dimerelloidea, and (in some analyses) Pugnacoidea appear to be monophyletic. Species classified in Dimerelloidea and Pugnacoidea, and in certain cases Hemithiridoidea, each form derived subclades that evolve from within a paraphyletic Norelloidea at the base of each subclade. Raduliform crura appear to be the most basal, phylogenetically; five other crural morphologies evolve from the raduliform state. However, morphological characters currently uniting genera in rhynchonellide superfamilies are not clearly diagnostic and exhibit a relatively high degree of homoplasy overall, suggesting that consistency with the classification may be based on a false sense of confidence in rhynchonellide morphology to clearly elucidate evolutionary relationships. Published molecular phylogenetic hypotheses conflict with the morphological topologies, further supporting this possibility.The evolutionary trends among diagnostic characters of Recent rhynchonellides appear to reflect successive juvenilization in adult morphology in several subclades, suggesting that heterochrony may have played an important role in the evolution of the group.
Abstract.-Patterns preserved in the fossil record are of the highest importance in addressing questions about long-term evolutionary processes, yet both the description of pattern and its translation into process can be difficult. With respect to gradual phyletic change, we know that randomly generated sequences may exhibit characteristics of a ''trend''; apparent patterns, therefore, must be interpreted with caution. Furthermore, even when the claim of a gradual trend can be statistically justified, interpretation of the underlying mechanisms may be challenging. Given that we can observe populations changing rapidly over tens or hundreds of years, it is now more difficult to explain instances of geologically gradual (as opposed to punctuated) change.Here we describe morphologic change in two bivalve lineages from the late Miocene Lake Pannon. We evaluate change according to the model-based methods of Hunt. Both lineages exhibit size increases and shape changes over an interval of nearly 4 million years. Size and two shape variables in the conjungens lineage are best fit by a model of directional evolution; remaining shape variables mostly conform to unbiased random walks. Body-size evolution in the diprosopum lineage is also significantly directional but all shape variables are best fit by the unbiased random walk model; the small number of sampling intervals available for this lineage (n 5 6) makes determination of the actual pattern more difficult. Model-fitting results indicate that the parallel trajectories of increasing log shell height over time in the two lineages can be accounted for by an underlying trend shared by both lineages, suggesting that the size increases may be a shared response to the same cause. The pace of phenotypic change, measured as Lynch's D, is slower than the neutral expectation for all size and shape traits.Our examples illustrate well the paradox of gradualism; the sequences exhibit significant directional morphological evolution, but rates of change as measured over the long-term are apparently too slow for directional selection or even drift to be the cause. Viewing long-term phenotypic evolution in terms of populations tracking peaks on adaptive landscapes is useful in this context. Such a view allows for intervals of directional selection (during times of peak movement-resulting in the overall trends we can detect) interspersed with intervals of stasis (during times of peak stability-resulting in overall changes that appear to proceed more slowly than the neutral expectation). The paradox of gradualism thus reduces to (1) peak movements and their drivers, which are not restricted in rate as are population-genetic drivers, and (2) the maintenance of stasis, on which no consensus exists.We can identify no environmental parameter in the central European Neogene that exhibits consistent change across the interval of gradual morphologic change. It may be that in Lake Pannon the long-term persistence of generally ameliorating conditions (plentiful resources and habitat space, few pr...
Independent molecular and morphological phylogenetic analyses have often produced discordant results for certain groups which, for fossil-rich groups, raises the possibility that morphological data might mislead in those groups for which we depend upon morphology the most. Rhynchonellide brachiopods, with more than 500 extinct genera but only 19 extant genera represented today, provide an opportunity to explore the factors that produce contentious phylogenetic signal across datasets, as previous phylogenetic hypotheses generated from molecular sequence data bear little agreement with those constructed using morphological characters. Using a revised matrix of 66 morphological characters, and published ribosomal DNA sequences, we performed a series of combined phylogenetic analyses to identify conflicting phylogenetic signals. We completed a series of parsimony-based and Bayesian analyses, varying the data used, the taxa included, and the models used in the Bayesian analyses. We also performed simulation-based sensitivity analyses to assess whether the small size of the morphological data partition relative to the molecular data influenced the results of the combined analyses. In order to compare and contrast a large number of phylogenetic analyses and their resulting summary trees, we developed a measure for the incongruence between two topologies and simultaneously ignore any differences in phylogenetic resolution. Phylogenetic hypotheses generated using only morphological characters differed among each other, and with previous analyses, whereas molecular-only and combined Bayesian analyses produced extremely similar topologies. Characters historically associated with traditional classification in the Rhynchonellida have very low consistency indices on the topology preferred by the combined Bayesian analyses. Overall, this casts doubt on the use of morphological systematics to resolve relationships among the crown rhynchonellide brachiopods. However, expanding our dataset to a larger number of extinct taxa with intermediate morphologies is necessary to exclude the possibility that the morphology of extant taxa is not dominated by convergence along long branches.
Crura, the calcareous support structures of the lophophore in rhynchonellide brachiopods, have historically been used to justify higher-level rhynchonellide classification and reveal major evolutionary lineages within rhynchonellides. Seventeen crural types have been described and categorized into four groups based on variation in overall structure and cross-sectional shape, but not evaluated in a quantitative or comprehensive manner. Heterochrony has been hypothesized to play a role in the evolutionary transitions among some types, but the structural, developmental, and phylogenetic context for testing these hypotheses has not yet been established. In this study, we use three-dimensional geometric morphometric techniques to quantify morphological disparity among all six crural morphs in Recent adult rhynchonellides, with the goal of delineating more objective criteria for identifying and comparing crural morphs, ultimately to test hypotheses explaining morphological transformations in ontogeny and phylogeny. We imaged the crura of seven Recent rhynchonellide species, using X-ray computed microtomography. We used landmarks and semi-landmarks to define the dimensions and curvature of the crura and the surrounding hinge area. Procrustes-standardized landmark coordinates were analyzed using a principal component analysis to test the discreteness of the individual crural morphs and named groups of morphs, and to identify features that vary most among the crural configurations.Our results demonstrate that microCT imaging techniques provide novel ways to investigate the morphology of small features that may be otherwise impossible to quantify using more conventional imaging techniques. Although we predicted overlap among crural morphs in the 3-D shape space, the principal component analyses suggest that five of the six crural morphs differ distinctly from one another. Some but not all previously designated crural groups appear to exhibit morphological cohesion. This study establishes a quantitative morphological foundation necessary to begin an investigation of the phylogenetic significance of ontogenetic changes in crura, which will allow hypotheses of heterochrony to be tested.
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