Phylogenetic reconstructions of two tropical American venerid genera, Chione and Chionopsis (subfamily Chioninae) were attempted at the species-level. The purposes of the analyses were to provide historical reconstructions of origination and extinction events in the two clades, as well as patterns of invasion and diversification. The analyses were based entirely on conchological characters to facilitate the inclusion of a substantial number of fossil taxa, but difficulties were encountered due to the quality of preservation and availability of material. Nevertheless the two genera were established as monophyletic clades, and the reconstructions yielded considerable insight into their histories in tropical America. The analyses suggest that both genera originated in the tropical western Atlantic, Chionopsis by at least the early Oligocene, and Chione in the early Miocene. Various branches of both genera subsequently invaded the tropical eastern Pacific several times prior to Seaway closure, with only one possible reciprocal invasion of the western Atlantic. Pliocene extinction affected both genera more significantly in the western Atlantic relative to the eastern Pacific, and diversity is higher today in the latter region. These conclusions are not entirely consistent with the fossil records of the genera, but this incongruency highlights the need for much more extensive sampling of the eastern Pacific Cenozoic record.

A thorough understanding of how communities respond to extreme changes, such as biotic invasions, is essential to manage ecosystems today. Here we constructed fossil food webs to identify changes in Late Ordovician (Katian) shallow-marine paleocommunity structure and functioning before and after the Richmondian invasion, a well-documented ancient invasion. Food webs were compared using descriptive metrics and cascading extinction on graphs models. Richness at intermediate trophic levels was underrepresented when using only data from the Paleobiology Database relative to museum collections, resulting in a spurious decrease in modeled paleocommunity stability. Therefore, museum collections and field sampling may provide more reliable sources of data for the reconstruction of trophic organization in comparison to online data repositories. The invasion resulted in several changes in ecosystem dynamics. Despite topological similarities between pre- and postinvasion food webs, species loss occurred corresponding to a minor decrease in functional groups. Invaders occupied all of the preinvasion functional guilds, with the exception of four incumbent guilds that were lost and one new guild, corroborating the notion that invaders replace incumbents and fill preexisting niche space. Overall, models exhibited strong resistance to secondary extinction, although the postinvasion community had a lower threshold of collapse and more variable response to perturbation. We interpret these changes in dynamics as a decrease in stability, despite similarities in overall structure. Changes in food web structure and functioning resulting from the invasion suggest that conservation efforts may need to focus on preserving functional diversity if more diverse ecosystems are not inherently more stable.

Organismal metabolic rates reflect the interaction of environmental and physiological factors. Thus, calcifying organisms that record growth history can provide insight into both the ancient environments in which they lived and their own physiology and life history. However, interpreting them requires understanding which environmental factors have the greatest influence on growth rate and the extent to which evolutionary history constrains growth rates across lineages. We integrated satellite measurements of sea-surface temperature and chlorophyll-a concentration with a database of growth coefficients, body sizes, and life spans for 692 populations of living marine bivalves in 195 species, set within the context of a new maximum-likelihood phylogeny of bivalves. We find that environmental predictors overall explain only a small proportion of variation in growth coefficient across all species; temperature is a better predictor of growth coefficient than food supply, and growth coefficient is somewhat more variable at higher summer temperatures. Growth coefficients exhibit moderate phylogenetic signal, and taxonomic membership is a stronger predictor of growth coefficient than any environmental predictor, but phylogenetic inertia cannot fully explain the disjunction between our findings and the extensive body of work demonstrating strong environmental control on growth rates within taxa. Accounting for evolutionary history is critical when considering shells as historical archives. The weak relationship between variation in food supply and variation in growth coefficient in our data set is inconsistent with the hypothesis that the increase in mean body size through the Phanerozoic was driven by increasing productivity enabling faster growth rates.

Phylogenetic reconstructions of two tropical American venerid genera, Chione and Chionopsis (subfamily Chioninae) were attempted at the species-level. The purposes of the analyses were to provide historical reconstructions of origination and extinction events in the two clades, as well as patterns of invasion and diversification. The analyses were based entirely on conchological characters to facilitate the inclusion of a substantial number of fossil taxa, but difficulties were encountered due to the quality of preservation and availability of material. Nevertheless the two genera were established as monophyletic clades, and the reconstructions yielded considerable insight into their histories in tropical America. The analyses suggest that both genera originated in the tropical western Atlantic, Chionopsis by at least the early Oligocene, and Chione in the early Miocene. Various branches of both genera subsequently invaded the tropical eastern Pacific several times prior to Seaway closure, with only one possible reciprocal invasion of the western Atlantic. Pliocene extinction affected both genera more significantly in the western Atlantic relative to the eastern Pacific, and diversity is higher today in the latter region. These conclusions are not entirely consistent with the fossil records of the genera, but this incongruency highlights the need for much more extensive sampling of the eastern Pacific Cenozoic record.

Demonstrating stasis in the fossil record has proven to be problematic with respect to both data collection and analysis. A previous approach is morphometric analysis of lineages sampled temporally and geographically. A hypothesis of stasis is apparently supported if morphological distances between descendent species and ancestral species are no greater than those between geographically distant samples of the descendent species. Evidence presented in this paper conflicts with such interpretations for at least one bivalve lineage, Chione erosa–Chione cancellata, of the late Neogene of tropical America. The direction and magnitude of morphological variance were quantified between two geographically distant groups of C. cancellata from the Recent, and compared to Pleistocene samples of C. cancellata and Pliocene samples of C. erosa. The results indicate that, although the magnitude of intraspecific geographic variation is as great as interspecific temporal variation, the species are morphologically discrete groups. The direction of morphological variance is as important as its magnitude, and interpretations overlooking this point are at best equivocal.

Previous studies have suggested that unbiased random walks may serve as appropriate null hypotheses for the detection of pattern in stratophenetic series. While numerous processes that influence the perceived temporal morphological evolution of a species may yield stratophenetic patterns that conform to the model of a random walk, use of the model as a null hypothesis raises several concerns. First, unbiased random walks are only a subset of a much larger set of random motions, including biased and fractional random walks. Some of these motions could also serve as appropriate null models for stratophenetic patterns. Second, due in part to the fractal nature of random walks, many types of time series begin to resemble random walks statistically as sampling resolution decreases. Therefore, indiscriminate support for unbiased random walks as null hypotheses of stratophenetic pattern leads inevitably to the commitment of Type II errors (incorrect failure to reject a null hypothesis). In this paper we simulate different hypothetical patterns of microevolution using various random walk models and apply the test of the null hypothesis. The frequency of Type II errors increases as stratigraphic completeness decreases, but at a currently unknown rate. Moreover, the test is insensitive to nongradual patterns of anagenesis.

We also demonstrate that a previously published approach is closely related to a standard method of fractal time series analysis and represents a good qualitative test of evolutionary pattern. The statistical variation underlying this method, however, is currently unknown, and further work is required to make it a robust quantitative test.

We used 24 fossil and Recent species to construct character states of both composite and exemplar taxa for phylogenetic analyses of Neogene genera and subgenera of the Corbulidae from tropical America. All characters were conchologic and two matrices, which differed in the manner that commarginal-rib characters were coded, were analyzed using branch and bound searches and maximum parsimony. Character polarity was determined using Corbula sensu strictu as an outgroup. These analyses produced a limited number of robust and well-resolved cladograms that require only one ghost lineage. Such stable results indicate a high level of congruency among characters, and demonstrate that conchologic data sets can yield highly resolved cladograms.

Tropical American corbulids are not monophyletic, and include two major clades. Crown groups within these clades are endemic, and all genera endemic to tropical America first appear in the Miocene. In fact, generic diversity and body size peak in the upper Miocene of the Caribbean/western Atlantic. Range restrictions and extinctions of large-bodied genera from both corbulid clades contribute to a post-Miocene decline in body size in this region. The eastern Pacific does not experience a similar decline in diversity and body size. Diversity and morphologic trends in Caribbean corbulids coincide with regional environmental changes, in particular decreases in seasonality and productivity. Except for the extinction of Bothrocorbula, however, corbulid extinctions apparently predate faunal turnover reported for other molluscs.

Associated with these extinctions, we found evidence of geographic range restriction, but not range shifts, in corbulid genera, indicating that the geologic development of environmental refugia contributed more to survival than eurytopy. Large-bodied genera of the southern Caribbean Gatunian Province (Tenuicorbula, Panamicorbula, and Hexacorbula) became restricted to the eastern Pacific. Range restriction to this high productivity refugium (i.e., paciphilic genera), rather than origination of new taxa, produced several corbulid genera now endemic to the region. Large-bodied genera from the Caloosahatchian Province of the southeastern U.S. (Bothrocorbula and Bicorbula), however, underwent global extinction. These subtropical and warm temperate taxa are presumably more eurytopic than their tropical counterparts, but do not exhibit range shifts in response to Neogene environmental change.

The incorporation of the random walk model into stratophenetic analysis marked a turning point by presenting a potential null model for microevolutionary patterns. Random walks are derived from a family of statistical fractals, and their statistics can be reconstructed using appropriate techniques. This paper lays the foundation for the explicit and uniform description of evolutionary mode in stratophenetic series using random walk null models and the information contained within incompletely preserved time series.

The method relies upon the iterative analysis of subseries of an original stratophenetic series by measuring the presence of deviations from statistical randomness as the lineage evolves. This measure, and its probability of significance (evaluated using a randomization test), forms the dimensions of a descriptive space for microevolutionary modes. Each stratophenetic series can then be viewed as a journey through this space. Computer simulation of various evolutionary modes demonstrates that different modes, for example stasis and gradualism, have differing trajectories and occupy different regions of the microevolutionary space. The method is applied to two published foraminiferal stratophenetic series, the Mio-Pliocene Globorotalia plesiotumida-tumida punctuated transition and an anagenetic trend in the Late Cretaceous Contusotruncana fornicata-contusa lineage. An anagenetic trend is strongly supported in the latter example, whereas transformation of the Globorotalia species seems to result from the fluctuating effectiveness of constraining processes.

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.

A model is developed to explore the potential responses of paleocommunities to disruptions of primary production during times of mass extinction and ecological crisis. Disruptions of primary production are expected to generate bottom-up cascades of secondary extinction, and these are predictable given species richnesses, functional diversity, and trophic link distributions. If, however, consumers are permitted to compensate for the loss of trophic resources by increasing the intensities of their remaining biotic interactions, top-down driven catastrophic increases of secondary extinction emerge from the model. Both bottom-up and top-down effects are themselves controlled by the geometry of the food webs. The general Phanerozoic trends of increasing taxonomic and ecological diversities, as well as the varying strengths of biotic interactions, have led to food webs of increasing complexity. The frequency of catastrophic secondary extinction increases as food web complexity increases, but increased complexity also serves to dampen the magnitude of the secondary extinctions. When intraguild competitive interactions are included in the model, competitively inferior taxa are observed to possess greater probabilities of survival if the guilds are embedded in simple subnetworks of the overall food web. The result is the emergence of post-extinction guilds dominated by those inferior taxa. These results are congruent with empirical observations of “disaster taxa” dominance after some mass extinction events, and provide a mechanism for the reorganization of ecosystems that is observed after those events. The model makes the testable prediction that dominance by disaster taxa, however, should be observed only when bottom-up disruptions have caused ecosystems to collapse catastrophically.

Liromissus, a new genus of chionine bivalve from the Neogene of Venezuela, serves as an example of the link between geographic range, endemism and extinction susceptibility. Unlike other Neogene tropical American chionine genera, Liromissus was not widespread in the tropical Western Atlantic and Eastern Pacific, but was instead restricted to the Atlantic Gatunian region. Late Neogene extinctions in tropical America affected taxa with restricted geographic ranges, and regions with higher degrees of endemism more severely. Accordingly, Liromissus is the only tropical chionine genus to suffer extinction during the Neogene.

The type species of Liromissus, L. quirosensis, was originally described as a member of the genus Lirophora. Morphological considerations preclude this assignment, but also demonstrate that Liromissus possesses all the characters diagnostic of the Chioninae.

One of the most enduring evolutionary metaphors is Van Valen's (1973) Red Queen. According to this metaphor, as one species in a community adapts by becoming better able to acquire and defend resources, species with which it interacts are adversely affected. If those other species do not continuously adapt to compensate for this biotically caused deterioration, they will be driven to extinction. Continuous adaptation of all species in a community prevents any single species from gaining a long-term advantage; this amounts to the Red Queen running in place. We have critically examined the assumptions on which the Red Queen metaphor was founded. We argue that the Red Queen embodies three demonstrably false assumptions: (1) evolutionary adaptation is continuous; (2) organisms are important agents of extinction; and (3) evolution is a zero-sum process in which living things divide up an unchanging quantity of resources. Changes in the selective regime need not always elicit adaptation, because most organisms function adequately under many “suboptimal” conditions and often compensate by demonstrating adaptive flexibility. Likewise, ecosystems are organized in such a way that they tend to be robust and capable of absorbing invasions and extinctions, at least up to a point. With a simple evolutionary game involving three species, we show that Red Queen dynamics (continuous adaptation by all interacting species) apply in only a very small minority of possible outcomes. Importantly, cooperation and facilitation among species enable competitors to increase ecosystem productivity and therefore to enlarge the pool and turnover of resources. The Red Queen reigns only under a few unusual circumstances.

Food webs represent trophic interactions among species in communities. Those interactions both structure and are structured by species richness, ecological diversity, and evolutionary processes. Geological and macroevolutionary timescales are therefore important to the understanding of food web dynamics, and there is a need for the consideration of paleocommunity food webs. The fossil record presents challenges in this regard, but the problem can be approached with combinatoric analysis and network theory. This paper is an introduction to the aspects of those disciplines relevant to the study of paleo-food webs, and explores a probabilistic and numerical approach.

The objective of this project is to understand the long-term relationships, if any, if within-clade bivalve shell morphology to variations in shell growth rate and possible underlying environmental factors. As a preliminary approach to this problem, a study was undertaken to record the history of shell morphology, on the Florida peninsula, of the venerid bivalve Chione cancellata L. (Lower Pliocene - Recent). Samples were obtained from the following formations: Lower Pinecrest (3.5-3.0 mya), Caloosahatchee Fm. (2.5-1.8 mya), Upper Pinecrest Beds (2.4-1.8 mya), Bermont Fm. (1.6-1.1 mya), Anastasia Fm. (0.05 mya), and Recent. The total data set comprised ten samples.

Eleven morphometric measurements were selected on their predicted sensitivity to growth rate-caused variations in valve convexity. Comparisons of sample data sets were performed using principal components and canonical variates analyses. Only left-handed valves were analyzed. The results of both the principal component and canonical variates analyses indicate that a disproportionately large amount of the variance in the composite data set is accounted for by one morphometric dimension. This dimension is essentially a summary of contrast between anterior hinge morphology and posterior adductor muscle morphology. The samples form three major clusters; (1) Bermont Fm., Anastasia Fm. and Recent samples, (2) Caloosahatchee Fm. and U. Pinecrest samples, and (3) L. Pinecrest samples. There is some overlap of extremes between clusters (2) and (3), but otherwise the clusters are distinct. The result is a gradual but significant change of morphology during a 3.5 m.y. time span.

The next step in the project is to seek an explanation for the above observations. The provinciality of the phenomenon will be tested more extensively with the inclusion of Neogene and Recent samples from the Southern Caribbean and the Atlantic Coastal Plain. Also, considering the dynamism of Florida's oceanographic conditions (temperature, productivity and circulation) during the last 3.5 m.y., correlation of oceanographic conditions with C. cancellata's growth rate and morphology will be examined. The proposed method is a systematic sampling of stable oxygen and carbon isotopes from valves within the samples analyzed.