Understanding the patterns, causes, and consequences of biotic interchange—the movement of species between neighboring biotas—is crucial for evaluating the effects of human-introduced species in the modern biosphere. Since at least the early Miocene, the tropical and subtropical western Atlantic has comprised two biogeographic provinces, the Gatunian (including the Caribbean) and the Caloosahatchian (North Carolina to Florida and the Yucatán peninsula). Although these adjacent provinces are not separated by a land barrier, exchange of species between them has been limited and intermittent. A synthesis of taxonomic, phylogenetic, stratigraphic, and biogeographic data on six gastropod and two bivalve groups reveals a dramatic shift in the pattern of interchange between these provinces. About 31% of early Miocene Caloosahatchian subgenus- and species-group-level taxa invaded the Gatunian Province by the late Miocene, but no taxa extended their ranges in the opposite direction. Beginning in the early Pliocene and continuing into the early Pleistocene, 40 taxa (roughly one-third of Gatunian diversity) invaded the Caloosahatchian Province from the Caribbean, whereas only four taxa extended their range from Florida into the Caribbean.

Comparisons between the ranked percentage of Gatunian invaders in Florida and the magnitude of regional extinction there for each of four middle Pliocene to early Pleistocene intervals reveal no consistent relation between invasion and prior or concurrent extinction. During the Pliocene, invaders not only compensated for extinctions, but also accounted for almost all the observed increase in standing diversity in Florida. Only after the large extinction event at the end of the Pliocene did invaders from the Gatunian Province not fully compensate for the loss of species.

Although the Miocene interaction between the Gatunian and Caloosahatchian biotas involved two fully tropical entities, the Plio–Pleistocene interaction exemplifies a general pattern in which tropical species often spread to higher latitudes during warm intervals, but warm-temperate or subtropical species rarely become established in the Tropics. Some evidence indicates that tropical Caribbean molluscs are exposed and adapted to more intense competition and predation than their subtropical counterparts in Florida, implying a role for individual-level biotic interactions in determining the predominant direction of interchange. Intensification of north-flowing currents in the western Atlantic may also contribute to the nearly one-way movement of taxa from the Caribbean to Florida during the Pliocene and early Pleistocene. The changing pattern of interchange from the Miocene to the Pliocene further reflects a change in the geography of species richness, with the richer province serving as the chief donor and the province with lower diversity acting as the main recipient of invaders. Diversity, ocean circulation, and the competitive environment thus account for the observed switch in the predominant direction of invasion in the western Atlantic during the Neogene.

The fact that almost 90% of Gatunian immigrants to Florida differentiated taxonomically there indicates that invasion is intermittent. Long-term consequences of this and many other cases of interchange between provinces include enrichment of the regional and global species pool and the spread of adaptations reflecting intense competition and predation.

The sizes and shapes of marine organisms often vary systematically across latitude and water depth, but the environmental factors that mediate these gradients in morphology remain incompletely understood. A key challenge is isolating the individual contributions of many, often correlated, environmental variables of potential biological significance. Benthic foraminifera, a diverse group of rhizarian protists that inhabit nearly all marine environments, provide an unparalleled opportunity to test statistically among the various potential controls on size and volume–to–surface area ratio. Here, we use 7035 occurrences of 541 species of Rotallid foraminifera across 946 localities spanning more than 60 degrees of latitude and 1600 m of water depth around the North American continental margin to assess the relative influences of temperature, oxygen availability, carbonate saturation, and particulate organic carbon flux on their test volume and volume–to–surface area ratio. For the North American data set as a whole, the best model includes temperature and dissolved oxygen concentration as predictors. This model also applies to data from the Pacific continental margin in isolation, but only temperature is included in the best model for the Atlantic. Because these findings are consistent with predictions from the first principles of cell physiology, we interpret these statistical associations as the expressions of physiological selective pressures on test size and shape from the physical environment. Regarding existing records of temporal variation in foraminiferal test size across geological time in light of these findings suggests that the importance of temperature variation on the evolution of test volume and volume–to–surface area ratio may be underappreciated. In particular, warming may have played as important a role as reduced oxygen availability in causing test size reduction during past episodes of environmental crisis and is expected to inflict metabolic stress on benthic foraminifera over the next century due to anthropogenic climate change.

Growth increments in bivalve shells have been used in geophysics and paleoecology. However, little study has been given to the variability of increment patterns within and between shells. The variability of increment patterns is examined visually and quantitatively, with the help of a serial correlation computer program. Analyses of increment patterns of the bivalves Chione fluctifraga and Protothaca staminea (Veneridae) indicate that: 1) quantitative results complement the visual observations; 2) increments are not consistently detected during successive measurements; 3) a striking similarity exists between right and left valves of the same specimen; and 4) different specimens collected from the same natural environment do not show any one-to-one increment relationship—however, increment width trends may be present. The variation of increment patterns seen between specimens suggests caution in the use of bivalve increments in paleoecology.

The Mesozoic-Cenozoic coral Order Scleractinia has been suggested to have originated or evolved (1) by direct descent from the Paleozoic Order Rugosa or (2) by the development of a skeleton in members of one of the anemone groups that probably have existed throughout Phanerozoic time. In spite of much work on the subject, advocates of the direct descent hypothesis have failed to find convincing evidence of this relationship. Critical points are:

(1) Rugosan septal insertion is serial; Scleractinian insertion is cyclic; no intermediate stages have been demonstrated. Apparent intermediates are Scleractinia having bilateral cyclic insertion or teratological Rugosa.

(2) There is convincing evidence that the skeletons of many Rugosa were calcitic and none are known to be or to have been aragonitic. In contrast, the skeletons of all living Scleractinia are aragonitic and there is evidence that fossil Scleractinia were aragonitic also. The mineralogic difference is almost certainly due to intrinsic biologic factors.

(3) No early Triassic corals of either group are known. This fact is not compelling (by itself) but is important in connection with points 1 and 2, because, given direct descent, both changes took place during this only stage in the history of the two groups in which there are no known corals.

Theoretical morphology is the scientific field in which researchers model organism growth and form. The field is developed well in studies on skeletons, especially shells. Researchers have contributed echinoid skeleton models to the field, but these have yet to be recognized collectively. We present herein the first comprehensive review for echinoid skeleton models in theoretical morphology. We apply a phylogenetic systematic analysis to those models, use the resulting consensus cladogram to classify and interrelate the models in an analogy in which they are likened to fossil specimens in a biostratigraphic record, and utilize the biostratigraphic metaphor to define trends within theoretical morphology as it applies to echinoid skeleton models.

Recent morphometric analysis revealed a juvenile (meraspid) axial growth gradient in the trunk of the ~429 Myr old trilobite Aulacopleura koninckii that resulted from growth control based on positional specification, as is common among extant organisms. Here we explore axial growth gradients in the more anterior body region, the cephalon, and in the cephalon and trunk during subsequent development in the holaspid period. We detected an axial growth gradient in the cephalon in the meraspid period, flatter and opposite in direction to that of the trunk, which also persisted during the holaspid period. We also found an holaspid trunk growth gradient, with a different distribution of growth rates among segments than that of the meraspid period. These newly observed growth gradients are compatible with the mechanism of growth control inferred for the meraspid trunk. Thus, the same kind of growth control may have operated in both body regions and during the whole ontogeny of A. koninckii. This study, along with others on the same species that preceded it, show that morphometric analysis of appropriate data sets can address questions of high interest for evolutionary developmental biology using data from fossils. By revealing developmental features at deep nodes of the phylogenetic tree, these studies will elucidate both how developmental processes evolved and how they themselves affected the evolution of organismal body patterning.

Most of the mammalian diversity is known only from fossils, and only a few of these fossils are well preserved or abundant. This undersampling poses serious problems for understanding mammalian phenotypic evolution under a quantitative genetics framework, since this framework requires estimation of a group’s additive genetic variance–covariance matrix (G matrix), which is impossible, and estimating a phenotypic variance–covariance matrix (P matrix) requires larger sample sizes than what is often available for extinct species. One alternative is to use G or P matrices from extant taxa as surrogates for the extinct ones. Although there are reasons to believe this approach is usually safe, it has not been fully explored. By thoroughly determining the extant and some extinct Xenarthra (Mammalia) cranium P matrices, this study aims to explore the feasibility of using extant G or P matrices as surrogates for the extinct ones and to provide guidelines regarding the reliability of this strategy and the necessary sample sizes. Variance–covariance and correlation P matrices for 35 cranium traits from 16 xenarthran genera (12 extant and 4 extinct) were estimated and compared between genera. Results show xenarthran P-matrix structures are usually very similar if sample sizes are reasonable. This study and others developed with extant therian mammals suggest, in general, that using extant G or P matrices as an approximation to extinct ones is a valid approach. Nevertheless, the accuracy of this approach depends on sample size, selected traits, and the type of matrix being considered.

A key question in studies of mass extinctions is whether the extinction was a sudden or gradual event. This question may be addressed by examining the locations of fossil occurrences in a stratigraphic section. However, the fossil record can be consistent with both sudden and gradual extinctions. Rather than being limited to rejecting or not rejecting a particular scenario, ideally we should estimate the range of extinction scenarios that is consistent with the fossil record. In other words, rather than testing the simplified distinction of “sudden versus gradual,” we should be asking, “How gradual?”

In this paper we answer the question “How gradual could the extinction have been?” by developing a confidence interval for the duration of a mass extinction. We define the duration of the extinction as the time or stratigraphic thickness between the first and last taxon to go extinct, which we denote by Δ. For example, we would like to be able to say with 90% confidence that the extinction took place over a duration of 0.3 to 1.1 million years, or 24 to 57 meters of stratigraphic thickness. Our method does not deny the possibility of a truly simultaneous extinction; rather, in this framework, a simultaneous extinction is one whose value of Δ is equal to zero years or meters.

We present an algorithm to derive such estimates and show that it produces valid confidence intervals. We illustrate its use with data from Late Permian ostracodes from Meishan, China, and Late Cretaceous ammonites from Seymour Island, Antarctica.

Paleobiological phylogenies often contain contrasting biological information to phylogenies based on extant species. Available software is primarily based on a present-day view of the world, however. Here, we present freeware to visualize phylogenies that is more suitable for the needs of paleobiologists, providing flexibility in how stratigraphic uncertainty, the geological timescale, and ancestor-descendant relationships are depicted.

The seemingly aberrant coiling of heteromorphic ammonoids suggests that they underwent more significant changes in hydrostatic properties throughout ontogeny than their planispiral counterparts. Such changes may have been responses to different selective pressures at different life stages. The hydrostatic properties of three species of Didymoceras (D. stevensoni, D. nebrascense, and D. cheyennense) were investigated by creating virtual 3D models at several stages during growth. These models were used to compute the conditions for neutral buoyancy, hydrostatic stability, orientation during life, and thrust angles (efficiency of directional movement). These properties suggest that Didymoceras and similar heteromorphs lived low-energy lifestyles with the ability to hover above the seafloor. The resultant static orientations yielded a downward-facing aperture in the hatchling and a horizontally facing aperture throughout most of the juvenile stage, before terminating in an upward direction at maturity. Relatively high hydrostatic stabilities would not have permitted the orientation of Didymoceras to be considerably modified with active locomotion. During the helical phase, Didymoceras would have been poorly suited for horizontal movement, yet equipped to pirouette about the vertical axis. Two stages throughout growth, however, would have enhanced lateral mobility: a juvenile stage just after the formation of the first bend in the shell and the terminal stage after completion of the U-shaped hook. These two more mobile phases in ontogeny may have improved juvenile dispersal potential and mate acquisition during adulthood, respectively. In general, life orientation and hydrostatic stability change more wildly for these aberrantly coiled ammonoids than their planispiral counterparts.

“There are still many dinosaur people who will say that, ‘Yes, there was an impact, but there were hardly any dinosaurs left by then.’ It's possible, but my first class prejudice is not to accept coincidence on that scale” (S. Gould quoted in Glenn 1994: p. 266, referring to the K/T mass extinction).

Variations in the orientation and cross-sectional shape of filamentous microfossils provide quantitative measures for characterizing them and probing their native mechanical structure. Here, we determine the tangent correlation length, which is the characteristic length scale for the variation in direction of a sinuous curve, for both a suite of Precambrian filamentous microfossils and six strains of modern filamentous cyanobacteria, all with diameters of a few microns. Among 1.9-2-Ga microfossils, Gunflintia grandis, Gunflintia minuta and Eomycetopsis filiformis possess, respectively, correlation lengths of 360 ± 40 μm, 670 ± 40 μm and 700 ±100 μm in two dimensions. Hundreds of times larger than the filament diameters, these values lie in the same range as the cyanobacteria Geitlerinema and Pseudanabaena, but are smaller than several strains of Oscillatoria. In contrast, the 2-Ga microfossil trichome Halythrix, is found to have a short correlation length of 29 ± 4 μm in two dimensions. Micron-wide pyritic replacement filaments observed in 3.23-Ga volcanogenic deposits also display a modest correlation length of 100 ± 15 μm in two dimensions. Sequences of species in two genera of our modern cyanobacteria possess tangent correlation lengths that rise as a power of the filament diameter D—D3.3 ± 1 for Oscillatoria and D5.1 ± 1 for Geitlerinema. These results can be compared with power-law scaling of D3 for hollow tubes and D4 for solid cylinders that is expected from continuum mechanics. Extrapolating the observed scaling behavior to smaller filament diameters, the measured correlation length of the pyrite filaments is consistent with modern Geitlerinema whereas that of Halythrix lies not far from modern Oscillatoria, suggesting that there may be structural similarities among these genera.

Although Darwin was not the first to conceive directional selection as a mechanism of phenotypic change, it is his ideas that were received, and that have shaped population biology to this day. A significant change in his theoretical orientation occurred in the mid-1850s. About then he abandoned environmental selection in favor of competitive selection, and adopted relative adaptation with all its consequences as an alternative. These ideas changed his thinking fundamentally and shaped his argument throughout the writing of his great book. It is still these ideas that predominate today.

Here I examine Darwin's ideas in relation to his principle of divergence, sexual selection, and the nature and origin of species. Finally I suggest that had he not misunderstood the function of sexual communication he might well have understood the nature of species and provided a more penetrating resolution to Herschell's “mystery of mysteries,” with which he opened his book.

Whether or not climate plays a causal role in mammal body-size evolution is one of the longest-standing debates in ecology. Bergmann's Rule, the longest-standing modeladdressing this topic, posits that geographic body-mass patterns are driven by temperature, whereas subsequent research has suggested that other ecological variables, particularly precipitation and seasonality, may be the major drivers of body-size evolution. While paleoecological data provide a unique and crucial perspective on this debate, paleontological tests of Bergmann's rule and its corollaries have been scarce. We present a study of body-size evolution in three ecologically distinct families of mammal (equids, canids, and sciurids) during the Oligo-Miocene of the northwest United States, an ideal natural laboratory for such studies because of its rich fossil and paleoclimatic records. Body-size trends are different in all three groups, and in no case is a significant relationship observed between body size and any climatic variable, counter to what has been observed in modern ecosystems. We suggest that for most of the Cenozoic, at least in the Northwest, body mass has not been driven by any one climatic factor but instead has been the product of complex interactions between organisms and their environments, though the nature of these interactions varies from taxon to taxon. The relationship that exists between climate and body size in many groups of modern mammals, therefore, is the exception to the rule and may be the product of an exceptionally cool and volatile global climate. As anthropogenic global warming continues and ushers in climatic conditions more comparable to earlier intervals of the Cenozoic than to the modern day, models of corresponding biotic variables such as body size may lose predictive power if they do not incorporate paleoecological data.

The late Paleozoic ice age (LPIA) had a profound effect on the biota. Despite much 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 show an 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 medium-scale ecological changes even during intervals of relative macroevolutionary quiescence.

Biophysical modeling and morphologic data from the fossil record were used to investigate the functional significance of changes in thallus morphology during the early evolutionary history of dasycladalean algae (dasyclads), a clade of benthic marine macroalgae. Modeling results indicate that the addition of cylindrical appendages (laterals) to an upright main axis, a key morphological innovation in the evolution of dasyclad thallus form, can provide for large gains in light interception efficiency, near-maximum gains in this regard being achieved when the ratio of total lateral surface area to main axis surface area is 4 or greater. Among the 13 early Paleozoic study taxa, all but one was found to exceed this value. Modeling of surface area to volume ratios for early Paleozoic dasyclads indicates that laterals for these forms conveyed only modest gains in this regard and, therefore, likely played little role in improving nutrient uptake. For survivorship, it appears that increasing thallus complexity by developing laterals conveyed important benefits by imparting both compartmentalization and redundancy, thereby increasing the likelihood that lateral-bearing forms would survive attacks by early mesograzers. Trends and patterns in the fossil record support such a survival-enhancing role for laterals and are consistent with the initial evolution of these structures as an early manifestation of an evolutionary arms race between macroalgae and herbivores initiated near the Proterozoic/Phanerozoic boundary.

Placodontia were a group of marine reptiles that lived in shallow nearshore environments during the Triassic. Based on tooth morphology it has been inferred that they were durophagous, but tooth morphology differs among species: placodontoid placodonts have teeth described as hemispherical, and the teeth of more highly nested taxa within the cyamodontoid placodonts have been described as flat. In contrast, the sister taxon to the placodonts, Palatodonta bleekeri, like many other marine reptiles, has tall pointed teeth for eating soft-bodied prey. The goals of this paper are to quantify these different tooth morphologies and compare tooth shape among taxa and with a functionally “optimal” tooth. To quantify tooth morphology we measured the radius of curvature (RoC) of the occlusal surface by fitting spheres to 3D surface scans or computed microtomographic scans. Large RoCs correspond to flatter teeth, while teeth with smaller RoCs are pointier; positive RoCs have convex occlusal surfaces, and a negative RoC indicates that the occlusal surface of the tooth is concave. We found the placodontoid taxa have teeth with smaller RoCs than more highly nested taxa, and palatine teeth tend to be flatter and closer to the optimal morphology than maxillary teeth. Within one well-nested clade, the placochelyids, the rearmost palatine teeth have a more complex morphology than the predicted optimal tooth, with an overall concave occlusal surface with a small, medial cusp. These findings are in keeping with the hypothesis that placodonts were specialized durophagous predators with teeth modified to break hard prey items while resisting tooth failure.

Morphological gradients in graptoloid colonies are explained by the production and diffusion of a morphogen from the sicula distalwards. Size of the thecae is inversely related to the amount of morphogen present. The graduate decrease of morphogen is given by a set of algebraic equations, and its effect on zooid growth is described by a modified Michaelis-Menten relationship. Changes in size of thecae computed on the basis of these equations fit the changes observed in graptolite colonies. Sets of differential equations are also given to describe the rate of morphogen diffusion and some other processes possibly involved in the development of uniaxiate graptoloid colonies. The suggested basic model of these colonies can be completed by a number of additional assumptions; however, computer experiments reveal that such assumptions do not affect the main properties of the model, namely the appearance of graded series of thecae. New evidence for regeneration of the sicular portion of the broken rhabdosome is presented, providing arguments that morphogen was produced by the siculozooid as a single bolus of secretion. Some local exclusions from the regular size gradient are discussed, and tentative explanations are suggested.

The Permian/Triassic mass extinction marks a permanent phylogenetic shift in the composition of the sessile benthos, from one largely dominated by articulate brachiopods to one dominated by mollusks. Widespread evidence of oceanic hypoxia and anoxia at this time provides a possible selective kill mechanism that could help explain the large taxonomic losses in brachiopods compared to the morphologically and ecologically similar bivalve molluscs. Our study compared the oxygen consumption of an articulate brachiopod, Terebratalia transversa, with that of two pteriomorph bivalves, Glycymeris septentrionalis and Mytilus trossulus, under normoxia and hypoxia, as well as their tolerance to anoxia, to gain insight into the relative metabolic characteristics of each group. We found no significant difference in the oxygen consumption of the three species when normalized to the same dry-tissue mass. However, when calculated for animals of the same external linear dimensions, bivalve oxygen consumption was two to three times greater than that of brachiopods. Our results also showed no significant decrease in the oxygen consumption of the three species until measured at a partial pressure of oxygen ∼10% of normoxic values. Finally, T. transversa and M. trossulus showed no significant difference in their tolerance to complete anoxia, but both showed a much lower tolerance than another bivalve, Acila castrensis. Findings from this study suggest that oxygen limitation is unlikely to account for the observed selective extinction of brachiopods during the Permian/Triassic mass extinction. Results may provide valuable information for assessing hypotheses put forth to explain why articulate brachiopods continue to remain a relatively minor group in marine environments.

If your undergraduate experience in stratigraphy was anything like mine, it was overwhelmingly dull. It was all about nomenclature and classification and endless lists of formation names. There were apparently no questions to be asked and the goal of stratigraphy was to devise boxes into which to place all types of geologic data. Notwithstanding the important work of correlation, the perception was that it was the sedimentologists who did the real science, science that focused on the processes of sediment accumulation and their controlling factors. Since then, stratigraphy has undergone a conceptual revolution, and it is necessary to examine how the field has changed and what those advances mean for paleobiology.