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Evolutionary paleoecologists have proposed many explanations for Phanerozoic trends in ecospace utilization, including escalation, seafood through time, filling of an empty ecospace, and tiering, among others. These hypotheses can be generalized into four models of functional diversification within a life-habit ecospace framework (functional-trait space). The models also incorporate concepts in community assembly, functional diversity, evolutionary diversification, and morphological disparity. The redundancy model produces an ecospace composed of clusters of functionally similar taxa. The partitioning model produces an ecospace that is progressively subdivided by taxa along life-habit gradients. The expansion model produces an ecospace that becomes progressively enlarged by the accumulation of taxa with novel life habits. These models can be caused by a wide range of ecological and evolutionary processes, but they are all caused by particular “driven” mechanisms. A fourth, neutral model also exists, in which ecospace is filled at random by life habits: this model can serve as a passive null model. Each model produces distinct dynamics for functional diversity/disparity statistics when simulated by stochastic simulations of ecospace diversification. In this first of two companion articles, I summarize the theoretical bases of these models, describe their expected statistical dynamics, and discuss their relevance to important paleoecological trends and theories. Although most synoptic interpretations of Phanerozoic ecological history invoke one or more of the driven models, I argue that this conclusion is premature until tests are conducted that provide better statistical support for them over simpler passive models.
Models of functional ecospace diversification within life-habit frameworks (functional-trait spaces) are increasingly used across community ecology, functional ecology, and paleoecology. In general, these models can be represented by four basic processes, three that have driven causes and one that occurs through a passive process. The driven models include redundancy (caused by forms of functional canalization), partitioning (specialization), and expansion (divergent novelty), but they also share important dynamical similarities with the passive neutral model. In this second of two companion articles, Monte Carlo simulations of these models are used to illustrate their basic statistical dynamics across a range of data structures and implementations. Ecospace frameworks with greater numbers of characters (functional traits) and ordered (multistate) character types provide more distinct dynamics and greater ability to distinguish the models, but the general dynamics tend to be congruent across all implementations. Classification-tree methods are proposed as a powerful means to select among multiple candidate models when using multivariate data sets. Well-preserved Late Ordovician (type Cincinnatian) samples from the Kope and Waynesville formations are used to illustrate how these models can be inferred in empirical applications. Initial simulations overestimate the ecological disparity of actual assemblages, confirming that actual life habits are highly constrained. Modifications incorporating more realistic assumptions (such as weighting potential life habits according to actual frequencies and adding a parameter controlling the strength of each model's rules) provide better correspondence to actual assemblages. Samples from both formations are best fit by partitioning (and to lesser extent redundancy) models, consistent with a role for local processes. When aggregated as an entire formation, the Kope Formation pool remains best fit by the partitioning model, whereas the entire Waynesville pool is better fit by the redundancy model, implying greater beta diversity within this unit. The ‘ecospace’ package is provided to implement the simulations and to calculate their dynamics using the R statistical language.
Numerous methods exist for estimating the true stratigraphic range of a fossil taxon based on the stratigraphic positions of its fossil occurrences. Many of these methods require the assumption of uniform fossil recovery potential—that fossils are equally likely to be found at any point within the taxon's true range. This assumption is unrealistic, because factors such as stratigraphic architecture, sampling effort, and the taxon's abundance and geographic range affect recovery potential. Other methods do not make this assumption, but they instead require a priori quantitative knowledge of recovery potential that may be difficult to obtain. We present a new Bayesian method, the Adaptive Beta method, for estimating the true stratigraphic range of a taxon that works for both uniform and non-uniform recovery potential. In contrast to existing methods, we explicitly estimate recovery potential from the positions of the occurrences themselves, so that a priori knowledge of recovery potential is not required. Using simulated datasets, we compare the performance of our method with existing methods. We show that the Adaptive Beta method performs well in that it achieves or nearly achieves nominal coverage probabilities and provides reasonable point estimates of the true extinction in a variety of situations. We demonstrate the method using a dataset of the Cambrian mollusc Anabarella.
Paleoecological studies enhance our understanding of biotic responses to climate change because they consider long timescales not accessible through observational and experimental studies. Using predatory drillholes produced on fossil bivalve shells by carnivorous gastropods, we provide an example of how climate change affected predator—prey interactions. We quantitatively examine temporal changes in fossil molluscan assemblages and predation patterns from the Pleistocene Japan Sea, which experienced drastic environmental changes in relation to glacial—interglacial climate cycles. We found significant changes in predation patterns associated with a decline in the abundance of warm-water molluscan species. Climate-mediated fluctuations in the eustatic sea level and resultant weakening of the Tsushima Warm Current caused a decline in a warm-water shell-drilling predator, which moderated the predation pressure and size relationship between the predators and the bivalve prey. Our results indicate that climate-mediated range shifts of species in present-day and future marine ecosystems can likewise increase altered predator—prey interactions.
We explore the relationships among the geographic ranges of genera, the ranges and positions of their constituent species, and the number of species they contain, considering variation among coeval genera and changes within genera over time. Measuring range size as the maximal distance, or extent, between occurrences within a taxon, we find that the range of the most widespread species is a good predictor of the range of the genus, and that the number of species is a better predictor still. This analysis is complicated by a forced correlation: the range of a genus must be at least as large as that of each of its constituent species. We therefore focus on a second measure of range, the mean squared distance, or dispersion, of occurrences from the geographic centroid, which, by analogy to the analysis of variance, allows the total dispersion of a genus to be compared to the mean within-species dispersion and the dispersion among species centroids. We find that among-species dispersion is the principal determinant of genus dispersion. Within-species dispersion also plays a major role. The role of species richness is relatively small. Our results are not artifacts of temporal variation in the geographic breadth of sampled data. The relationship between changes in genus dispersion and changes in within- and among-species dispersion shows a symmetry, being similar in cases when the genus range is expanding and when it is contracting. We also show that genera with greater dispersion have greater extinction resistance, but that within- and among-species dispersion are not demonstrable predictors of survival once the dispersion of the genus is accounted for. Thus it is the range of the genus, rather than how it is attained, that is most relevant to its fate. Species richness is also a clear predictor of survival, beyond its effects on geographic range.
Lungfishes are known for, and indeed take their name from, their bimodal respiratory abilities. All three extant genera can use their lungs to extract oxygen from the atmosphere, although their reliance upon this capability differs among taxa. Lungs are considered primitive for the Osteichthyes, however the distinctive buccal pump mode of air gulping exhibited by extant lungfishes appears to be a specialization. It is associated with a number of derived skeletal characters (cranial ribs, long parasphenoid stalk, midline gap between palatal tooth plates) that first appeared during the Devonian. These have been described individually, but in no Devonian lungfish has their three-dimensional (3D) spatial relationship been reconstructed and analyzed. Here we present the 3D morphology of Rhinodipterus, a Mid-Late Devonian lungfish from Australia and Europe, based on synchrotron tomography and conventional microtomography scans.
Unlike less crownward contemporaneous lungfishes such as Griphognathus and Chirodipterus, Rhinodipterus has a full set of skeletal buccal pump components that can be directly compared to those of extant lungfishes, suggesting that it made more extensive use of air breathing than other Gogo or Bergisch Gladbach genera. This is interesting in relation to the environmental context as Gogo and Bergisch Gladbach are both marine, contrasting with the frequently hypoxic tropical to subtropical fresh water environments inhabited by modern lungfishes. The evolution of buccal pump-supported lung ventilation was evidently not necessarily associated with a transition to non-marine habitats.
New growth rate estimates for nine species from three genera of New Zealand Crassatellidae (Mollusca: Bivalvia), combined with existing morphometric ontogenetic descriptions, allow identification of heterochronic processes in the evolution of these genera. Both paedomorphosis (progenesis and neoteny) and peramorphosis (hypermorphosis and acceleration) have occurred within the clade. Overall, morphological variability and response to environmental pressure in this nonsiphonate group is restricted by the interplay of anatomical and life habit constraints. Stability in the substrate, predator avoidance, sluggish burrowing speed, and inability to escape by deep burial are suggested as key drivers of, or constraints on, morphological change. Two groups of shell characters are identified: heavy, armored “anchors” and elongate “snorkels,” which combine juvenile and adult traits in shells of different sizes and ages, produced by heterochronic variation in developmental timing. Anchors and snorkels both represent different “solutions” to the problems of life as a nonsiphonate, infaunal bivalve.
Graphoglyptids are deep-marine trace fossils, often found preserved as casts in positive relief on the base of turbidites. Previous analyses of the behavioral evolution of graphoglyptids suggested they were slowly diversifying, becoming optimized, and getting smaller over time until the Late Cretaceous, when a sudden increase in diversification occurred. This current study quantifies the morphology of approximately 400 different graphoglyptid specimens, ranging in age from the Cambrian to the present, in order to evaluate the behavioral evolutionary interpretations made previously. Results from this study indicate that although some general evolutionary patterns can be discerned, they are not as straightforward as previously reported.
Different topological categories of trace fossils represent organisms' responses to evolutionary pressures in unique ways. While burrow widths of meandering traces were becoming smaller over time, as predicted by previous workers, the burrow widths of the network traces were becoming smaller only until the Late Cretaceous, when they started to get larger again. The times of significant evolutionary changes in behavior were not consistent among various topological categories, with some morphological features being affected in the Late Cretaceous and others during the beginning of the Eocene. It is likely that the behavioral evolution of graphoglyptids was influenced by deep-marine global influences linked to climate change, glaciation, and deep-ocean warming. These influences affected each topological group uniquely, suggesting that different species or genera of trace makers were creating each of the topological categories. This is contrary to the hypothesis that all graphoglyptids were created by closely related species.
The late Paleozoic ice age (LPIA) had a profound effect on the biota. Despitemuch research having been focused on paleotropical regions or global-scale analyses, regional ecological changes have seldom been studied in ice-proximal basins. Here, I study the compositional turnover and diversity structure across the main Carboniferous glacial event recorded in western Argentina and the subsequent nonglacial interval. Brachiopod and bivalve data from western Argentina suggest that the transition from glacial to nonglacial climates caused major compositional changes. Turnover, however, was not uniform across the bathymetric gradient, being higher in deep environments. Because extirpation was concentrated in brachiopods, but immigration was similar in both clades, the taxonomic structure of the region was significantly modified. Although regional hierarchical diversity structure and occupancy distributions remained stable, dissecting the analysis in brachiopods and bivalves underscores that both clades had different responses to climate change. Brachiopods, on the one hand, show stability in the diversity structure and a very slight decrease in occupancies of intermediate genera, while bivalves showan important rise in diversity, both at the environment and regional scale, and an increase in genera with intermediate occupancies. The bathymetric diversity gradient was also modified from hump shaped with maximum diversity in the deep subtidal to a linear gradient with maximum values toward the offshore. However, relative compositional differences within environments remained stable, with maximum values at intermediate depths both in glacial and nonglacial intervals. Moreover, local-scale coexistence between brachiopods and bivalves changed in the nonglacial interval, showing significant segregation, which indicates relevant modifications in community assembly dynamics. Results from western Argentina highlight the magnitude of regional-scale ecological changes during the LPIA in ice-proximal regions, suggesting that the waxing and waning of glaciers was able to cause regional taxonomic turnover and mediumscale ecological changes even during intervals of relative macroevolutionary quiescence.