Bryozoans are colonial animals and this permits the partitioning of their morphologic variability into components of within colony (i.e. within a single genotype) and between colony (i.e. between genotype) variance. These data have been obtained for four species of the endemic deep-sea genus Euginoma for a series of characters. In 8 comparisons, one component of the total variance dominated at the 5% level. Population (between colony) variance contributed significantly to the total variance in 63% of the comparisons (5 of 8); individual (within colony) variance contributed significantly to the total variance in 37% of the comparisons (3 of 8).

Compared to shallow water species, the surprising feature of the deep-sea data is that the between colony component of variance is as high as it is. Possibly in the more stable, deep-sea environment, the genotypic contribution to the variance of each individual colony is expressed to a greater degree than in the more variable, shallow water regime. If so, then analyses of variability in colonial animals may be an independent means of ascertaining stability gradients in the fossil record.

The most abundant and diverse graptolite assemblages are found in offshore, deep-water black shales—the classical “graptolite fades” (deep-water or isograptid biofacies). The mean duration of Ordovician graptolite species confined to the deep-water facies (here referred to as “group 1” species) is 2.19 Myr, significantly shorter than the mean duration of species in the deep-water facies that are also known in sediments of the shallow-water shelf or platform (“group 2” species) −4.42 Myr, indicating a significantly higher extinction probability (p = <0.001). These figures are based on the precise age ranges of species derived from the time-calibrated composite sequence of 1446 Ordovician to early Devonian graptolites, built by the constrained optimization procedure (CONOP) from 256 measured sections worldwide, and exclude the effects of the Hirnantian mass extinction. The difference between groups cuts across families, morphological types, and pandemic/endemic distributions. An environmental influence is strongly suggested, and although both groups were planktonic, they were unlikely to have shared the same habitat in the water column. The new duration measurements therefore are interpreted as favoring a depth-stratification of graptolite habitats in the water column.

Issues of directionality in the history of life can be framed in terms of six major evolutionary steps, or megatrajectories (cf. Maynard Smith and Szathmáry 1995): (1) evolution from the origin of life to the last common ancestor of extant organisms, (2) the metabolic diversification of bacteria and archaea, (3) evolution of eukaryotic cells, (4) multicellularity, (5) the invasion of the land and (6) technological intelligence. Within each megatrajectory, overall diversification conforms to a pattern of increasing variance bounded by a right wall as well as one on the left. However, the expanding envelope of forms and physiologies also reflects—at least in part—directional evolution within clades. Each megatrajectory has introduced fundamentally new evolutionary entities that garner resources in new ways, resulting in an unambiguously directional pattern of increasing ecological complexity marked by expanding ecospace utilization. The sequential addition of megatrajectories adheres to logical rules of ecosystem function, providing a blueprint for evolution that may have been followed to varying degrees wherever life has arisen.

The stratigraphic distribution of graptoloid species within the upper Middle Ordovician strata of New York State represents a complex pattern of origination, extinction, faunal migration, and fluctuating relative abundances. In particular, the observed patterns of species turnover at graptolite biozone boundaries are apparently strongly correlated with lithofacies, sampling strategies, and the depositional effects of relative sea-level change.

Vertical facies changes that occur within third-order depositional sequences and fourth- or fifth-order parasequences are mirrored by changes in the graptoloid faunal composition. Large-scale faunal turnovers at biozone boundaries tend to occur either at sequence boundaries or at flooding surfaces within sequences (e.g., the base of Highstand System Tracts). The base of the Corynoides americanus and Climacograptus (D.) spiniferus Zones coincide with major onlap events, and the Orthograptus ruedemanni Zone fauna appears just below a Lowstand Systems Tract. Within individual biozones, smaller-scale changes such as the fluctuating relative abundances of graptoloid species coincide with higher-order parasequence cyclicity. These distribution patterns resemble recent computer-generated models for the sequence stratigraphic distribution of hypothetical taxa.

By combining good biogeographic control with a detailed sampling program, we are able to see through the patterns attributable to depositional cyclicity and identify the different components of faunal turnover (migration, speciation, and extinction).

A large body of paleontological literature concerns the importance of ontogeny as a source of morphological variation for evolution; morphologies that appear during one stage of an organism's development are made available for use in another simply by modifying the developmental program. Paleontologists need to know why this occurs, so they can study the process of evolution in extinct animals and so they can discuss the fossil record in terms that are applicable to modern forms. If most cases of heterochrony can be attributed to life-history evolution then the fossil record provides evidence of the nature of selection (in particular the age-specific mortality) that extinct animals experienced. The hypothesis of interest here is that species in which maturity is accelerated will also show generalized morphology and small size, while those with delayed maturity will have more specialized morphology and large size.

Four species of the ostracode genus Cyprideis were studied to determine whether differences in age at maturity are correlated with heterochrony in the expected manner. For each species the changes in size and shape through geological time were evaluated in the statistical context of modern geographic and seasonal variation. Living populations were sampled regularly to detect differences in seasonality and to estimate the duration of development.

Evolution of ontogeny is apparent at the level of species in this group, but it is not simply related to differences in life-history. In comparisons among species, we find evidence of heterochrony where there is no difference in the age at maturity, and a difference in age at maturity where there is no heterochrony. Similarly, three of the four species show the expected positive correlation between size and age at maturity, yet the fourth species is relatively large and matures rapidly. Cyprideis does not support the generalization that life-history evolution causes heterochrony, and casts doubt on the inference of life-history evolution from heterochrony where the data are drawn exclusively from extinct forms.

The unique holotype of the Triassic reptile Sharovipteryx (initially Podopteryx) mirabilis has been reexamined and redescribed, correcting the original account. It is a small lizard-like reptile with an elongate head and densely tubular femur and tibia each longer than the (estimated) intergirdle distance. (Part of the pectoral region has been lost and the remnants remain encased in the matrix.) The matrix retains the impression of portions of the integument, including a patagium that reached from the hindlimbs to the base of the tail. Unlike the original account, the patagium did not extend to the pectoral limbs. Experiments with models indicate that Sharovipteryx could have maintained a shallow glide if the femora were held at a shallow angle to the vertebral column and the tibia and feet extended out at right angles to it, thus stretching the integument. The pectoral limbs (with or without a fringing membrane) might have produced a variable canard; alternatively vertical bending of the tail could have applied drag, each making the glide more stable.

Fibrous ligament and aragonitic shell material have been found preserved in two specimens of radiolitid rudists [cf. Radiolites angeoides (De Lapeirouse)] from the Lower to Middle Coniacian of Tirol, Austria. The invaginated ligament apparently comprised two torn and overgrown non-functional strings, one in each valve, connected medially by a thin strip of active ligament. The original dimensions of the active ligament have been estimated, for shells of 4 to 5 cm commissural diameter, at 0.58 mm2 cross-sectional area and 1.3 mm length, with the shell closed. Using data on the adductor muscle of Recent oysters and the results of compression tests carried out on samples of oyster fibrous ligament, it is estimated that these radiolitid shells opened less than 1 mm in life; their gaping was highly restricted. This is taken to support the proposal that the expanded mantle margins (principally that of the right, attached valve) were responsible for food particle entrapment, rather than the gills as in most other bivalves. It is suggested that a tentaculate mantle flange was extruded through the narrow gap between the valves during feeding. The pair of pronounced radial bands on the posterior flanks of the shell are interpreted as marking pseudofaecal and faecal ejection sites. Finally, it is speculated that slime (possibly poisonous or foul-tasting) on the mantle tentacles inhibited predators.

Kelps and other fleshy macroalgae—dominant reef-inhabiting organisms in cool seas—may have radiated extensively following late Cenozoic polar cooling, thus triggering a chain of evolutionary change in the trophic ecology of nearshore temperate ecosystems. We explore this hypothesis through an analysis of body size in the abalones (Gastropoda; Haliotidae), a widely distributed group in modern oceans that displays a broad range of body sizes and contains fossil representatives from the late Cretaceous (60–75 Ma). Geographic analysis of maximum shell length in living abalones showed that small-bodied species, while most common in the Tropics, have a cosmopolitan distribution, whereas large-bodied species occur exclusively in cold-water ecosystems dominated by kelps and other macroalgae. The phylogeography of body size evolution in extant abalones was assessed by constructing a molecular phylogeny in a mix of large and small species obtained from different regions of the world. This analysis demonstrates that small body size is the plesiomorphic state and largeness has likely arisen at least twice. Finally, we compiled data on shell length from the fossil record to determine how (slowly or suddenly) and when large body size arose in the abalones. These data indicate that large body size appears suddenly at the Miocene/Pliocene boundary. Our findings support the view that fleshy-algal dominated ecosystems radiated rapidly in the coastal oceans with the onset of the most recent glacial age. We conclude with a discussion of the broader implications of this change.

The fossil record provides an important source of data on adaptive radiations, and indeed some of the earliest theoretical insights on the nature of these radiations were made by paleontologists. Here we focus on the diverse Devonian Metacryphaeus group calmoniid trilobites, known from the Malvinokaffric Realm, which have been considered a classic example of an adaptive radiation preserved in the fossil record. We use a geometric morphometric analysis in conjunction with phylogenetic and biogeographic patterns and data on speciation rates. Using ancestral character state reconstruction during speciation events, we quantify patterns of morphological change in order to assess the role ecological and geographical factors may have played in mediating this radiation. We found no significant differences between the amount of morphological change that occurred during speciation events when ancestors and descendants were in the same area as opposed to when they occupied different areas. Further, the magnitude of morphological divergence did not change through time or with cladogenetic rank. These patterns, in conjunction with the fact that the radiation occurs in a geographically heterogeneous region subjected to repeated episodes of sea-level rise and fall, suggest that at the macroevolutionary scale this radiation may have been motivated more by phenomena that facilitated geographic isolation than by competition.

In this paper we analyze the ontogenies of four species of pachypleurosaur (Reptilia: Sauropterygia) occurring in Triassic-age deposits in the Monte San Giorgio region, Switzerland. Preservation of multiple complete specimens representing a growth series from each taxon allows the comparison of ontogenetic trajectories through a space composed of nine variables important in the evolution of the clade. Trajectories are characterized using the multivariate generalization of the allometry equation and then compared through calculations of the angles between allometry vectors. Individual coefficients of vectors are compared after calculation of bootstrapped confidence intervals. Pachypleurosaur ontogeny is found to be allometric and generally conserved, although significant differences between taxa exist. Shape-disassociated allometric changes characterize the transition between Serpianosaurus and Neusticosaurus, while allometric changes within Neusticosaurus are less significant. N. edwardsii is inferred to have arisen through hypermorphosis. Interpretation of whole-body heterochrony in multivariate analysis is discussed.

Primary vascular architecture of members of the Paleozoic Aneurophytales (Progymnosper-mopsida) is described. This architecture is somewhat more complex but fundamentally similar (homologous) to that of members of the Trimerophytina, putative ancestors of aneurophytes. It is suggested that the presence of complex stelar morphology in aneurophytes was epiphenomenal, a passive result of changes in growth and development in a trimerophyte-like ancestor. Specifically, I suggest that the evolutionary transformation in primary vascular architecture from haplostele to ribbed protostele was a direct consequence of changes that affected the vertical spacing and degree of organization of lateral appendages in early vascular plants. This view is in sharp contrast to adaptationist explanations of change in stelar morphology expressed by other authors and provides an example of non-adaptive change in evolution.

Evolutionary trends observed over large clades have the potential to mask underlying trends that occur within their constituent subclades. A recent study of encephalization in the Caniformia (Carnivora, Mammalia) found evidence for an abrupt increase in median log-encephalization quotients (logEQs), indicating higher brain volume relative to body mass, at the end-Miocene, but gradual increase in the variance of logEQs. In this study, new endocranial volume estimates for fossil taxa in the well-sampled caniform subclade Canidae are reported. Using the encephalization data for the Canidae, hypotheses of evolution in encephalization allometries were tested with respect to canid phylogeny. The Akaike Information Criterion and likelihood ratios recovered support for a preferred hypothesis of the evolution of canid encephalization, which proposed two distinct allometric relationships: (1) a plesiomorphic grade of encephalization in the subfamilies Hesperocyoninae and Borophaginae and the paraphyletic canine genus Leptocyon, and (2) an apomorphic grade in the crown radiation of Caninae. This defines a shift in to higher encephalization, but without an associated change in the variance around the allometry. Increased canid encephalization coincides with a reorganization of the brain and the observed trend may reflect the evolution of complex social behavior in this clade.

Biomechanical models illustrate how the principles of physics and physiology determine function in organisms, allowing ecological inferences and functional predictions to be based on morphology. Dynamic lever and linkage models of the mechanisms of the jaw and skull during feeding in fishes predict function from morphology and have been used to compare the feeding biomechanics of diverse fish groups, including fossil taxa, and to test ideas in ecological morphology. Here we perform detailed computational modeling of the four-bar linkage mechanism in the skull and jaw systems of Dunkleosteus terrelli, using software that accepts landmark morphological data to simulate the movements and mechanics of the skull and jaws during prey capture. The linkage system is based on the quadrate and cranio-thoracic joints: Cranial elevation around the cranio-thoracic joint forces the quadrate joint forward, which, coupled with a jaw depressor muscle connecting the jaw to the thoracic shield, causes the jaw to rotate downward during skull expansion. Results show a high speed transmission for jaw opening, producing a rapid expansion phase similar to that in modern fishes that use suction during prey capture. During jaw closing, the model computes jaw and skull rotation and a series of mechanical metrics including effective mechanical advantage of the jaw lever and kinematic transmission of the skull linkage system. Estimates of muscle cross-sectional area based on the largest of five specimens analyzed allow the bite force and strike speed to be estimated. Jaw-closing muscles of Dunkleosteus powered an extraordinarily strong bite, with an estimated maximal bite force of over 6000 N at the jaw tip and more than 7400 N at the rear dental plates, for a large individual (10 m total length). This bite force capability is among the most powerful bites in animals. The combination of rapid gape expansion and powerful bite meant that Dunkleosteus terrelli could both catch elusive prey and penetrate protective armor, allowing this apex predator to potentially eat anything in its ecosystem, including other placoderms.

The geographic distribution of brachiopod genus occurrences over the Phanerozoic shows that secular declines in origination and extinction rates were paralleled by increases in invasion and extirpation rates. Origination and extinction rates declined in two phases, the first from the Cambrian to latest Permian Periods and the second from the latest Permian Period to the present, which were accompanied by concomitant increases in invasion and extirpation rates. In addition to the temporal correlation, an inverse correlation was also weakly evident among time-averaged latitudinal gradients of rates. Compared with faunas at higher latitudes, low-latitude faunas experienced higher origination and extinction rates, and lower invasion and extirpation rates. We suggest that progressive increases in migration ability lowered origination and extinction rates because species that were better equipped to track a preferred habitat, for example, by the ability to disperse larvae over large distances, were less likely to evolve or become extinct in response to local environmental changes. The two phases were separated by the end-Permian mass extinction, which reset to high levels the origination and extinction rates of a taxonomically and ecologically altered global brachiopod fauna. Our data also allow us to quantify the relative contributions of origination, extinction, invasion, and extirpation to regional diversity (quantified as 10° latitudinal zones) more generally. Overall, invasion and extirpation explained slightly more variation in diversity than in situ origination and extinction. The four variables usually occurred in combinations that maintained rather than altered the shape of the latitudinal diversity gradient. For most of the Phanerozoic Eon, the gradient was not the product of continuous renewal, but rather existed as a holdover from a previous interval.

The origins and maintenance of morphological diversity through the Phanerozoic have been documented in several groups of invertebrates. By using shape analysis of the ventral valve, we quantify morphological diversity within the inarticulate brachiopods, which have a species-rich lower Paleozoic history followed by low richness through the rest of the Phanerozoic. Morphological diversity peaked during the Cambrian, and was subsequently maintained despite significant decreases in richness at all taxonomic levels. The loss of morphological diversity, associated with late-Paleozoic decreases in the number of orders and the Permo-Triassic mass extinction, was followed by rapid morphological rediversification without concordant taxonomic diversification. Any limits upon the generation of diverse morphologies were established early in the Cambrian and were maintained and approached through the history of this group.

A computer simulation was developed which estimates the effects of differential preservation on the value of taxonomic diversity in fossil assemblages. Taxonomic diversity was measured with the Shannon-Weaver index, H′ and with evenness, J. Preservation markedly alters quantitative measurements of taxonomic diversity. The degree of alteration can, however, be predicted on the basis of the species-abundance distribution and on the distributions of species of varying preservability. The indices of diversity H′ and J measure different aspects of diversity, but respond to differential preservation in a similar manner. Evenness, J, is recommended for paleoecologic use. The computer model can be used to revise fossil diversity data. A simple form of such a model has been used, and suggests that some previous interpretations of paleoecologic diversity may be in error.

Cases of convergent evolution, particularly within ecomorphological contexts, are instructive in identifying universally adaptive morphological features across clades. Tracing of evolutionary pathways by which ecomorphological convergence takes place can further reveal mechanisms of adaptation, which may be strongly influenced by phylogeny. Ecomorphologies of carnivorous mammals represent some of the most outstanding cases of convergent evolution in the Cenozoic radiation of mammals. This study examined patterns of cranial shape change in the dog (Canidae) and hyena (Hyaenidae) families, in order to compare the evolutionary pathways that led to the independent specialization of bone-cracking hypercarnivores within each clade. Geometric morphometrics analyses of cranial shape in fossil hyaenids and borophagine canids provided evidence for deep-time convergence in morphological pathways toward the independent evolution of derived bone-crackers. Both clades contained stem members with plesiomorphic generalist/omnivore cranial shapes, which evolved into doglike species along parallel pathways of shape change. The evolution of specialized bone-crackers from these doglike forms, however, continued under the constraint of a full cheek dentition and restriction on rostrum length reduction in canids, but not hyaenids. Functionally, phylogenetic constraint may have limited borophagine canids to crack bones principally with their carnassial instead of the third premolar as in hyaenids, but other cranial shape changes associated with durophagy nevertheless evolved in parallel in the two lineages. Size allometry was not a major factor in cranial shape evolution in either lineage, supporting the interpretation of functional demands as drivers for the observed convergence. The comparison between borophagines and hyaenids showed that differential effects of alternative functional “solutions” that arise during morphological evolution may be multiplied with processes of the “macroevolutionary ratchet” already in place to further limit the evolutionary pathways available to specialized lineages.

Edge-drilling is an unusual predation pattern in which a predatory gastropod drills a hole on the commissure between the valves of a bivalve. Although it is faster than wall drilling, it involves the potential risk of amputating the drilling organ. We therefore hypothesize that this risky strategy is advantageous only in environments where predators face high competition or predation pressure while feeding. The high frequency of edge-drilling (EDF, relative to the total number of drilled valves) in a diverse Recent bivalve assemblage from the Red Sea enables us to test this hypothesis, predicting (1) a low EDF in infaunal groups, (2) a high EDF in bivalves with elongated shape, (3) high incidence of edge-drilling in groups showing a high wall-drilling frequency, and (4) high EDF in shallow habitats. We evaluate these predictions based on >15,000 bivalve specimens. Among ecological attributes, we found substrate affinity and predation intensity of a species to be good predictors of edge-drilling incidence. Infaunal taxa with high length/width ratio have a low EDF, in accordance with our predictions. Predation intensity is also a significant predictor of edge-drilling; groups with high predation intensity show higher incidence of edge-drilling, confirming our prediction. Although water depth fails to show any significant effect on EDF, this analysis generally supports the high-risk hypothesis of edge-drilling incidence because shallow depths have considerable microhabitat variability. Classically the drill hole site selection has often been linked to predatory behavior. Our study indicates that prey attributes are also crucial in dictating the behavioral traits of a driller such as site selection. This calls for considering such details of the prey to fully understand predation in modern and fossil habitats. Moreover, this perspective is important for tackling the longstanding riddle of the limited temporal and spatial distribution of edge-drilling.

Standard techniques of energy exchange analysis are applied to modelling the thermal regimes of several species of sailback pelycosaurs. Of particular interest is the role played by the sail in thermoregulation. Although the sail did increase the rate at which the model could warm up in the morning, its effectiveness fell short of previous estimates. The sail increased the daytime internal temperature by typically 3°C to 6°C. The more massive subjects had the highest temperature increases owing partly to their higher thermal inertia and partly to their disproportionately large sails. The sail had no impact upon the nocturnal minimums in temperature.

Overheating does not appear to have been a significant problem in the Permian environment, particularly in view of the high thermal inertia of the subjects modeled. When overheating occurred, the sail was of limited value for dumping excess heat.

Fossil species of the family Hyaenidae represent a wide range of ecomorphological diversity not observed in living representatives of this carnivoran group. Among them, the cursorial meat-and-bone specialists are of particular interest not only because they were the most cursorial of the hyaenids, but also because they were the only members of this family to spread into the New World. Here we conduct a functional morphological analysis of the cranium of the cursorial meat-and-bone specialist Chasmaporthetes lunensis by using finite element modeling to compare it with the living Crocuta crocuta, a well-known bone-cracking carnivoran. As found with previous finite element studies on hyaenid crania, the cranium of C. lunensis is not differentially adapted for stress dissipation between the bone-cracking and meat-shearing teeth. A smaller occlusal surface on the more slender P3 cusp of C. lunensis allowed this species to use less bite force to crack a comparably-sized bone relative to C. crocuta, but higher muscle masses in the latter probably allow it to process larger food items. We use two indices, the stress slope and the bone-cracking index, to show that C. lunensis has a well-adapted cranium for stress dissipation given its size, but the main stresses placed on its cranium might have been more from subduing prey and less from cracking bones. Throughout the Cenozoic, other carnivores besides hyaenids convergently evolved similar morphologies, including domed frontal regions, suggesting an adaptive value for a repetitive mosaic of features. Our analyses add support to the hypothesis that bone-cracking adaptations are a complex model that has evolved convergently several times across different carnivoran families, and these predictable morphologies may evolve along a common gradient of functionality that is likely to be under strong adaptive control.