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For decades, theoretical morphological studies of different groups of organisms have been successfully pursued in biological, paleontological, and computational contexts, often with distinct modeling approaches and research questions. A regular influx of new perspectives and varied expertise has contributed to the emergence of a veritable multidisciplinary outlook for theoretical morphology. The broadening of this discipline is reflected in a substantial increase in the number of models, leading to a bewildering diversity that has yet to be scrutinized. In this work, we tackle this issue in a synthetic fashion, with a quantitative meta-analysis that allows an objective comparison of theoretical morphological models treated as entities. By analogy with empirical morphospace analyses of actual organisms, we performed a multivariate ordination of a representative sample of models, producing a metaspace of models in which patterns of similarity and difference are visualized. A phenetic tree was used to characterize the relationships between models. Four major groups have been identified, and their disparity analyzed. We suggest this typology as a useful starting point to identify a core set of fundamental principles and protocols for better interpretation of the plethora of current models and for more efficient construction of models in the future. This in turn can help in diversifying the scope of macroevolutionary, developmental, and bioenvironmental questions in theoretical morphology.
Extinction risk is inversely related to genus age (time since first appearance) in most intervals of the Phanerozoic marine fossil record, in apparent contradiction to the macroevolutionary Red Queen's Hypothesis, which posits that extinction risk is independent of taxon age. Age-dependent increases in the mean species richness and geographic range of genera have been invoked to reconcile this genus-level observation with the presumed prevalence of Red Queen dynamics at the species level. Here we test these explanations with data from the Paleobiology Database. Multiple logistic regression demonstrates that the association of extinction risk with genus age is not adequately explained by species richness or geographic range: there is a residual association between age and extinction risk even when range and richness effects are accounted for. Throughout most of the Phanerozoic the age selectivity gradient is highest among the youngest age cohorts, whereas there is no association between age and extinction risk among older age cohorts. Some of the apparent age selectivity of extinction in the global fauna is attributable to differences in extinction rate among taxonomic groups, but extinction risk declines with genus age even within most taxonomic orders. Notable exceptions to this pattern include the Cambrian–Ordovician, latest Permian, Triassic, and Paleocene intervals. The association of age with extinction risk could reflect sampling heterogeneity or taxonomic practice more than biological reality, but at present it is difficult to evaluate or correct for such biases. Alternatively, the pattern may reflect consistent extinction selectivity on some as-yet unidentified covariate of genus age. Although this latter explanation is not compatible with a Red Queen model if most genus extinctions have resulted from biological interactions, it may be applicable if most genus extinctions have instead been caused by recurrent physical disturbances that repeatedly impose similar selective pressures.
Major shifts in ecological dominance are one of the most conspicuous but poorly understood features of the fossil record. Here we examine one of the most prominent such shifts, the Ordovician shift from trilobite to brachiopod dominance of benthic ecosystems. Using an integrated database of high-resolution paleoecological samples and body size data, we show that while the average local richness and relative abundance of trilobites declined significantly through the Ordovician, the estimated standing biomass of trilobites, and by implication the amount of energy that they used, remained relatively invariant. This is attributable to an increase in the average body size of trilobite species in our data set, and especially to the widespread occurrence of the exceptionally large Middle–Late Ordovician trilobite genus Isotelus. Brachiopods increase in both mean body size and relative abundance throughout the Ordovician, so that estimates of brachiopod biomass and energetic use increase substantially between the Early and Late Ordovician. Although the data set includes a range of depositional environments, similar trends are observed in both shallow subtidal and deep subtidal settings. These results suggest that diversification of the Paleozoic Fauna did not come at the energetic expense of the Cambrian Fauna. The declining relative abundance of trilobites may reflect a combination of numerical dilution and the necessary energetic trade-offs between body size and abundance.
The problem of gradual versus punctuated change within phyletic lineages can be understood in terms of the homogeneity of evolutionary dynamics. Hypotheses of punctuated change imply that the rules governing evolutionary change shift over time such that the normal dynamics of stasis are temporarily suspended, permitting a period of net evolutionary change. Such explanations are members of a larger class of models in which evolutionary dynamics are in some way heterogeneous over time. In this paper, I develop a likelihood-based statistical framework to evaluate the support for this kind of evolutionary model. This approach divides evolutionary sequences into nonoverlapping segments, each of which is fit to a separate evolutionary model. Models with heterogeneous dynamics are generally more complex—they require more parameters to specify— than uniform evolutionary models such as random walks and stasis. The Akaike Information Criterion can be used to judge whether the greater complexity of punctuational models is offset by a sufficient gain in log-likelihood for these models to be preferred.
I use this approach to analyze three case studies for which punctuational explanations have been proposed. In the first, a model of punctuated evolution best accounted for changes in pygidial morphology within a lineage of the trilobite Flexicalymene, but the uniform model of an unbiased random walk remains a plausible alternative. Body size evolution in the radiolarian Pseudocubus vema was neither purely gradual nor completely pulsed. Instead, the best-supported explanation posited a single, pulsed increase, followed later by a shift to an unbiased random walk. Finally, for the much-analyzed claim of “punctuated gradualism” in the foraminifera Globorotalia, the best-supported model implied two periods of stasis separated by a period of elevated but not inherently directional evolution. Although the conclusions supported by these analyses generally refined rather than overturned previous views, the present approach differs from those prior in that all competing interpretations were formalized into explicit statistical models, allowing their relative support to be unambiguously compared.
The hypothesis of limiting similarity, which postulates that morphologically and/or ecologically similar species will differ enough in shape, size, or other variables to minimize competition, has been controversial among ecologists and paleoecologists. Many studies have reported the occurrence of limiting similarity in modern environments or in time-averaged fossil deposits; however, empirical high-resolution time series demonstrating limiting similarity over longer time scales are lacking. We have integrated radiocarbon-calibrated amino acid dating techniques, stable isotope estimates, and morphometric data to test the hypothesis of limiting similarity in late Quaternary land snails from the Canary Islands over a period of 42,500 years. We tested for both ecological character displacement (two closely related species will differ in size in order to minimize competition in sympatry and these differences will be minimized in allopatry) and community-wide character displacement (overdispersion of body size among competitors in a guild). Multiple proxies of body size consistently show that two endemic congeneric pulmonate gastropod species (Theba geminata and T. arinagae) maintained a difference in size from ∼42,500 b.p. through the last occurrence of T. arinagae 14,900 b.p., with a concomitant trend of a decreasing body size. Theba geminata body size did not converge on that of T. arinagae and variation in T. geminata body size did not increase significantly following the extinction of T. arinagae; therefore, ecological character displacement and release did not occur. Community-wide character displacement was found in only one time bin over the last 42,500 years. These results suggest that limiting similarity is a transient ecological phenomenon rather than a long-term evolutionary process. This study not only demonstrates the problems inherent in biological “snapshot” studies and geological studies of time-averaged deposits to test limiting similarity adequately, but it also presents a more adequate research protocol to test the importance of interspecific competition in the history of life.
An early Pliocene fossil locality in the Canadian High Arctic preserves the remains of the extinct beaver Dipoides sp. (Castoridae, Rodentia) in association with an assemblage of fossil beaver-cut wood. The wood assemblage presents a first opportunity to investigate woodcutting behavior and ecological performance in an extinct castorid genus. This study compares woodcutting in Dipoides sp. with that of the modern beaver, Castor canadensis, using evidence from small-diameter cut sticks (i.e., sticks transected by parallel series of cut marks) in combination with behavioral observations of Castor woodcutting. During woodcutting both Castor and Dipoides used their incisors unilaterally; the upper incisor was pressed against the stick while the corresponding lower incisor cut. Cut marks were relatively larger for Castor than Dipoides (scaled to incisor size). Compared with Dipoides, Castor more frequently used a cutting strategy that minimized the number of cuts needed to transect a stick (e.g., clipping as opposed to chip removal). Taken together, the behavioral evidence suggests that ecological cutting performance was lower for Dipoides than Castor.
Sabertooths exhibit one of the most extreme feeding adaptations seen in mammals. The functional consequences of accommodating extremely elongate upper canine teeth are severe, resulting in a well-documented suite of cranial modifications. We used geometric morphometric methods to study the evolution of overall shape in the skulls of extant and extinct feline and machairodontine felids, as well as extinct nimravids. Trends in skull evolution were evaluated by using relative warps analysis and tested for association with body size and canine tooth length. Primitive sabertooths from all lineages exhibit cranial shapes more similar to conical-toothed cats, despite the presence of moderately developed saberteeth. More-derived forms in both nimravids and felids diverge in skull morphospace to form two distinct sabertooth types (dirk-toothed and scimitar-toothed) that differ in canine shape. Skull shape in conical-toothed cats is strongly associated with body size, but not canine length. However, within each sabertooth lineage, skull shape is significantly correlated with canine length, suggesting that gape-related demands drove the evolution of sabertooth skull morphology.