Mesozoic tooth marks on bone surfaces directly link consumers to fossil assemblage formation. Striated tooth marks are believed to form by theropod denticle contact, and attempts have been made to identify theropod consumers by comparing these striations with denticle widths of contemporaneous taxa. The purpose of this study is to test whether ziphodont theropod consumer characteristics can be accurately identified from striated tooth marks on fossil surfaces. We had three major objectives (1) to experimentally produce striated tooth marks and explain how they form; (2) to determine whether body size characteristics are reflected in denticle widths; and (3) to determine whether denticle characters are accurately transcribed onto bone surfaces in the form of striated tooth marks. We conducted controlled feeding trials with the dental analogue Varanus komodoensis (the Komodo monitor). Goat (Capra hircus) carcasses were introduced to captive, isolated individuals. Striated tooth marks were then identified, and striation width, number, and degree of convergence were recorded for each. Denticle widths and tooth/body size characters were taken from photographs and published accounts of both theropod and V. komodoensis skeletal material, and regressions were compared among and between the two groups. Striated marks tend to be regularly striated with a variable degree of branching, and may co-occur with scores. Striation morphology directly reflects contact between the mesial carina and bone surfaces during the rostral reorientation when defleshing. Denticle width is influenced primarily by tooth size, and correlates well with body size, displaying negative allometry in both groups regardless of taxon or position. When compared, striation widths fall within or below the range of denticle widths extrapolated for similar-sized V. komodoensis individuals. Striation width is directly influenced by the orientation of the carina during feeding, and may underestimate but cannot overestimate denticle width. Although body size can theoretically be estimated solely by a striated tooth mark under ideal circumstances, many caveats should be considered. These include the influence of negative allometry across taxa and throughout ontogeny, the existence of theropods with extreme denticle widths, and the potential for striations to underestimate denticle widths. This method may be useful under specific circumstances, especially for establishing a lower limit body size for potential consumers.

Extensive investigation of the close association between biological diversity and environmental temperature has not yet yielded a generally accepted, empirically validated mechanism to explain latitudinal gradients of species diversity, which occur in most taxa. Using the highly resolved late Cenozoic fossil records of four major taxa of marine plankton, we show that their gradients arise as a consequence of asymmetric geographic range expansion rather than latitudinal variation in diversification rate, as commonly believed. Neither per capita speciation nor extinction rates trend significantly with temperature or latitude for these marine plankton. Species of planktonic foraminifera and calcareous nannoplankton that originate in the temperate zone preferentially spread toward and arrive earlier in the tropics to produce a normal gradient with tropical diversity peaks; by contrast, temperate-zone originating species of diatoms and radiolarians preferentially spread toward and arrive earlier in polar regions to produce reversed gradients with high-latitude diversity peaks. Our results suggest that temperature affects latitudinal diversity gradients chiefly by its effect on species’ range limits rather than on probabilities of speciation and extinction. We show that this mechanism also appears to operate in various multicellular taxa, thus providing a widely applicable explanation for the origin of latitudinal diversity gradients.

The fossil record of Phanerozoic brachiopod genera and Late Cenozoic New World mammal genera is examined for evidence of evolutionary equilibria. One necessary (but insufficient) condition is met: within temporal intervals, numbers of originations correlate with numbers of extinctions. Eliminating temporally short-ranging brachiopods, however, reduces the correlation so that it explains only 16% of the variation. More decisive tests of the equilibrium hypothesis appear impossible with available data. Difficulties of temporal and geographic scale, taxonomic level, and ecological consistency must be resolved before equilibrium models can be applied in paleontology for other than inspiration.

The deep-sea Cenozoic planktonic microfossil record has the unique characteristics of continuously well-preserved populations of most species, with virtually unlimited sample size, and therefore constitutes, in principle, a major resource for macroevolutionary research. Antarctic Neogene radiolarians in particular, are diverse, abundant and consistently well-preserved and evolved rapidly. This fauna is, in theory, a near-perfect testing ground for paleodiversity reconstructions. In this study we determined the diversity history of these faunas from a new quantitative, taxonomically complete data set from Neogene and Quaternary sections at several Antarctic sites. The pattern retrieved by our whole-fauna data set shows a significant, largely extinctionless ecological change in faunal composition and decrease in the evenness of species' abundances during the late Miocene, followed 3 Myr later, at around 5 Ma, by a significant drop in diversity. We tentatively associate this ecological event with a synchronous, regional change in the composition of the primary producers, but as yet cannot identify any environmental changes associated with the later extinction. Further, our whole-fauna diversity history was compared to diversity computed from much less complete, biostratigraphically oriented studies of species' occurrences, compiled in the Neptune database and reconstructed by using subsampling methodologies. Comparison of our whole-fauna and subsampling-reconstructed diversity patterns shows that the first-order trends are the same in both, suggesting that, to some degree, such literature compilations can be used to explore diversity history of plankton. However, our results also highlight substantial errors and distortions in the reconstructed diversity which make it poorly suited to more-detailed studies (e.g., for comparison of diversity history with paleoenvironmental history). We conclude that detailed studies of plankton diversity, and particularly those attempting to understand the relation between diversity and paleoceanographic change, should be based on taxonomically comprehensive, quantitative data.

Sutural perimeters of 301 Late Jurassic ammonites scale as the 3/8 power of phragmocone volume. This implies that septal surface grows as the ¾ power of body mass, the exponent of Kleiber's law (1932), one of the best-established empirical laws in biology, which is well known to be the scaling exponent of basal metabolic rate. Sutural complexity, as measured by fractal dimensions, emerges from the relationship between sutural perimeter and phragmocone volume, thus supporting the interpretations of septal folding as a mechanism for the increase in septal surface and as demanded by metabolic and physiologic processes (e.g., respiration or body chamber transport). The implications of these results strongly suggest that ammonite septa were involved in more than a simple structural support.

Torticonic (helically-coiled) ammonoids have been most commonly interpreted as vagile, benthonic forms. Their mode of coiling, however, places the siphuncle in a functionally dorsolateral, rather than ventral position on the shell whorl due to: 1) their probable apex-upward shell orientation during life, and 2) size asymmetry of the lateral sutural saddles due to helical coiling, associated with upward siphuncle displacement. The resultant positioning of the siphuncle produces cameral liquid de-coupling soon after initiation of cameral liquid removal (emptying of liquid from a newly formed chamber within the phragmocone). Since in Recent chambered cephalopods cameral liquid de-coupling increases efficiency of vertical migration, a similar mode of life was indicated for the torticonic ammonites.

Lysorophus, a serpentiform, nearly limbless, aquatic Paleozoic amphibian was described in detail by Sollas in 1920. Although Sollas did not discuss cranial kinesis, his original reconstructions suggest that Lysorophus had an unusually kinetic skull for an amphibian. We have reexamined the skull of Lysorophus and find that kinesis, if present, was slight. The maxillae, premaxillae, and vomers might have moved in limited protraction and retraction, with the premaxillae rotating about their contacts with the nasals. There is a possibility of small medio-lateral movements of the lacrimal and maxilla as a unit at the lacrimal-prefrontal joint. The skull of Lysorophus shows many features convergent with the skulls of burrowing reptiles, such as amphisbaenians. Modifications in the skull of Lysorophus from a more primitive tetrapod condition seem associated with burrowng in a soft substratum rather than with kinesis per se.

We assessed selective extinction patterns in bivalves during a late Neogene mass extinction event observed along the temperate Pacific coast of South America. The analysis of 99 late Neogene and Quaternary fossil sites (recorded from 7°S to 55°S), yielding ∼2800 occurrences and 118 species, revealed an abrupt decline in Lyellian percentages during the late Neogene–Pleistocene, suggesting the existence of a mass extinction that decimated ∼66% of the original assemblage. Using the late Neogene data set (n = 59 species, 1346 occurrences), we tested whether the extinction was nonrandom according to taxonomic structure, life habit, geographic range, and body size. Our results showed that the number of higher taxa that went extinct was not different than expected by random. At first sight, extinction was selective only according to life habit and geographic range. Nevertheless, when phylogenetic effects were accounted for, body size also showed significant selectivity. In general, epifaunal, small-sized (after phylogenetic correction), and short-ranged species tended to have increased probability of extinction. This is verified by strong interactions between the variables herein analyzed, suggesting the existence of nonlinear effects on extinction chances. In the heavily decimated epifaunal forms, survival was not enhanced by widespread ranges or larger body sizes. Conversely, the widespread and large-sized infaunal forms tended to have lower probability of extinction. Overall, the ultimate extinction of late Neogene bivalve species along the Pacific coast of South America seems to have been determined by a complex interplay of ecological and historical (phylogenetic) effects.

Within a ∼60-Myr interval in the Late Triassic to Early Jurassic, a major mass extinction took place at the end of Triassic, and several biotic and environmental events of lesser magnitude have been recognized. Climate warming, ocean acidification, and a biocalcification crisis figure prominently in scenarios for the end-Triassic event and have been also suggested for the early Toarcian. Radiolarians, as the most abundant silica-secreting marine microfossils of the time, provide a control group against marine calcareous taxa in testing selectivity and responses to changing environmental parameters. We analyzed the origination and extinction rates of radiolarians, using data from the Paleobiology Database and employing sampling standardization, the recently developed gap-filler equations and an improved stratigraphic resolution at the substage level. The major end-Triassic event is well-supported by a late Rhaetian peak in extinction rates. Because calcifying and siliceous organisms appear similarly affected, we consider global warming a more likely proximate trigger of the extinctions than ocean acidification. The previously reported smaller events of radiolarian turnover fail to register above background levels in our analyses. The apparent early Norian extinction peak is not significant compared to the long-term trajectory, and is probably a sampling artifact. The Toarcian Oceanic Anoxic Event, previously also thought to have caused a significant radiolarian turnover, did not significantly affect the group. Radiolarian diversity history appears unique and complexly forced, as its trajectory parallels major calcareous fossil groups at some events and deviates at others.

The miniaturization of shells, often observed in Mesozoic brachiopods, is analyzed in two species of the well-known genus Homoeorhynchia. We investigated shell development in the context of recent works that consider growth as a function of size, shape, and age. The miniaturization results from heterochronic disturbances during development. Although these shells exhibit a large morphological plasticity, architectural constraints limit their morphogenesis, so that small adult specimens are not exactly reduced replicas of large ones. Adult morphology depends largely on the pattern of rib formation, with miniaturization arising by two different processes affecting this pattern: predisplacement of the appearance of ribs in H. meridionalis and acceleration in H. cynocephala. The first case seems to be a reversible process without evolutionary consequences, but the second occurs within an evolutionary trend. In both cases, the miniaturization seems to be related to environmental influences. Study of the growth pathway of these species allows recognition of basic characters in the morphological variation of populations.

We have produced computer simulations of multiramous graptoloids with the intention of defining the rules governing branching strategies and colony form. Close matches between such simulations and real graptolites show that complex rhabdosomes may be produced by the permutation of relatively simple sets of rules. Those designs found in nature produce an efficient and regular distribution of zooids through the area included by an essentially planar rhabdosome. Strikingly geometrical arrays of stipes, such as the Goniograptus and yin/yang patterns, closely approach paradigmatic harvesting arrays. For dichotomously branching anisograptids the evolutionary trend in reduction of “primary stipes” can be explained by the production of larger spreading colonies. Multiramous graptoloids fed during vertical movement through the water column. Changes in a single branching decision may produce considerable changes in rhabdosome morphology, but these are not necessarily of high taxonomic importance; this is proved by a specimen which is a morphological combination of two “genera.” Although primarily under genetic control, certain modifications to colony form were probably the result of inhibitory interaction between adjacent stipes.

Determining which biological traits affect taxonomic durations is critical for explaining macroevolutionary patterns. Two approaches are commonly used to investigate the associations between traits and durations and/or extinction and origination rates: analyses of taxonomic occurrence patterns in the fossil record and comparative phylogenetic analyses, predominantly of extant taxa. By capitalizing upon the empirical record of past extinctions, paleontological data avoid some of the limitations of existing methods for inferring extinction and origination rates from molecular phylogenies. However, most paleontological studies of extinction selectivity have ignored phylogenetic relationships because there is a dearth of phylogenetic hypotheses for diverse non-vertebrate higher taxa in the fossil record. This omission inflates the degrees of freedom in statistical analyses and leaves open the possibility that observed associations are indirect, reflecting shared evolutionary history rather than the direct influence of particular traits on durations. Here we investigate global patterns of extinction selectivity in Devonian terebratulide brachiopods and compare the results of taxonomic vs. phylogenetic approaches. Regression models that assume independence among taxa provide support for a positive association between geographic range size and genus duration but do not indicate an association between body size and genus duration. Brownian motion models of trait evolution identify significant similarities in body size, range size, and duration among closely related terebratulide genera. We use phylogenetic regression to account for shared evolutionary history and find support for a significant positive association between range size and duration among terebratulides that is also phylogenetically structured. The estimated range size–duration relationship is moderately weaker in the phylogenetic analysis due to the down-weighting of closely related genera that were both broadly distributed and long lived; however, this change in slope is not statistically significant. These results provide evidence for the phylogenetic conservatism of organismal and emergent traits, yet also the general phylogenetic independence of the relationship between range size and duration.

Reconciliation of paleontological and molecular phylogenetic evidence holds great promise for a better understanding of the temporal succession of cladogenesis and character evolution, especially for taxa with a fragmentary fossil record and uncertain classification. In zoology, studies of this kind have largely been restricted to Bilateria. Hexactinellids (glass sponges) readily lend themselves to test such an approach for early-branching (non-bilaterian) animals: they have a long and rich fossil record, but for certain taxa paleontological evidence is still scarce or ambiguous. Furthermore, there is a lack of consensus for taxonomic interpretations, and discrepancies exist between neontological and paleontological classification systems. Using conservative fossil calibration constraints and the largest molecular phylogenetic data set assembled for this group, we infer divergence times of crown-group Hexactinellida in a Bayesian relaxed molecular clock framework. With some notable exceptions, our results are largely congruent with interpretations of the hexactinellid fossil record, but also indicate long periods of undocumented evolution for several groups. This study illustrates the potential of an integrated molecular/paleobiological approach to reconstructing the evolution of challenging groups of organisms.

The geometric regularity of oblique sculptures in pectinids and some other bivalves was examined both theoretically and empirically. Camptonectes microsculpture occurs exclusively on the external surface of foliated calcite, and is well characterized by an invariably orthogonal relation to growth increments and almost uniform density over the surface. It is genetically unrelated to radial and other sculpturing. According to our SEM observations, its divergent orientation seems to be controlled primarily by the growth of calcite laths. In contrast, the shagreen microsculpture of pectinids is characterized by a quincunx arrangement of scales on the diagonal network. Shagreen microsculpture can be regarded as a variant of diagonal sculpture together with the simple oblique, divaricate, V-shaped, and zigzag patterns that are widespread in other bivalves. The geometric regularity of these variations as well as their mutual relationship can be readily recognized by computer simulations. All of these diagonal sculptures may result from interference between radial and commarginal elements. The formation of diagonal sculptures seems to occur by the periodic activation of regularly arranged sculpture-producing cells on the mantle margin.

A new method of data analysis offers a potentially powerful tool for statistically evaluating hypotheses of rate in temporally-ordered evolutionary phenomena. We present a method for bootstrapping time-ordered data sets to test hypotheses of the equality of rate. This method is applicable to both nonrandom and random generative processes. The method is applied to the data of Malmgren et al. (1983) for the Globorotalia plesiotumida–G. tumida planktonic foraminiferan lineage and the data of Reyment (1982) for the benthonic foraminiferan Afrobolivina afar. G. plesiotumida is recognizable on the basis of independent data as a species distinct from G. tumida, its descendant. Evolutionary change rate during the evolution of G. tumida from G. plesiotumida is shown to be faster than rates within either species. The pattern of variation exhibited by A. afar includes a time interval of more rapid change; this more rapid change is observed post hoc. A bootstrapping model based on post hoc observations reveals the rate in this time interval to be not significantly faster than expected in such post hoc intervals.

Recent ideas about the role of competition in the ecology and evolution of acorn barnacles are based partly on relationships between morphology and the outcome of competition for space. One hypothesis is that present distributions and patterns of evolutionary diversification and decline among acorn barnacles reflect the competitive exclusion and replacement of solid-walled forms by those with tubiferous skeletal structure. An alternate view is that large barnacles generally outcompete smaller ones, independent of differences in skeletal structure, with predation and disturbance favoring the ecological and evolutionary success of small barnacles. Field experiments and observations in the Gulf of California suggested that the small, solid-walled species, Chthamalus anisopoma, competitively exlcudes the larger, tubiferous Tetraclita stalactifera confinis on the lower part of the shore. Greater tolerance to aerial exposure appears to allow Tetraclita to occupy a high intertidal refuge above Chthamalus. A common denominator among several cases of competitive exclusion in acorn barnacles is the greater settlement density of the competitive dominant, not its morphology. Morphological considerations alone are insufficient to predict or explain the outcome of competition between barnacle species.

Reconstructing patterns of macroevolution has become a central endeavor in paleobiology, because it offers insight into evolutionary models shaping the history of life. As the most diverse and abundant animals since the Cambrian period, arthropods provide copious data to elucidate the emergence of body plans in metazoan lineages. However, information provided by fossils on the tempo and mode of this phenomenon has lacked a recent synthesis. Here, I investigate macroevolutionary patterns of morphological evolution in Euarthropoda using a combined extinct and extant dataset optimized for multivariate analyses. Overall ordination patterns between the main morphogroups are consistent with another, independently coded, extant-only dataset providing molecular and morphological rates of evolution. Based on a “deep split” phylogenetic framework, total-group Mandibulata and Arachnomorpha emerge as directional morphoanatomical lineages, with basal fossil morphogroups showing heterogeneously spread-out occupations of the morphospace. In addition to a more homogeneous morphological variation, new morphogroups arose by successive reductions of translation distances; this pattern was interrupted only by terrestrialization events and the origin of pancrustaceans. A displaced optimum type of model is proposed to explain the fast assembly of canalized body plans during the Cambrian, with basal fossil morphogroups fitting intermediate fitness peaks in a moving adaptive landscape. Given time constraints imposed by the paleontological evidence, and owing to the interplay between canalization and modularity, as well as a decoupling between molecular and morphological rates, the rise of euarthropods would support the view that the swiftness of the Cambrian explosion was mostly associated with the buildup of genetic regulatory networks.

The species richness of extant metazoan phyla displays a strong association with the ontogenetic timing of germ-line determination. Taxa with early ontogenetic determination of the germ-line are characterized by low species number relative to taxa with late or variable determination. A test for a similar association in fossil taxa is compromised by the fact that all phyla displaying early determination of the germ-line are small, soft-bodied groups which lack a reliable fossil record. To the extent that this correlation reflects causality, patterns of evolutionary diversification have been decidedly non-random. The fact that ontogenetic timing of germ-line determination defines the extent to which genetic variation arising during the course of ontogeny may be inherited suggests a possible causal foundation for the pattern.

Fish eagles (genus Haliaeetus) potentially have contributed the bones of their prey to many late Cenozoic concentrations of vertebrate microfossils. To evaluate possible biases in the taxa and skeletal elements preserved in assemblages accumulated by fish eagles, which could affect paleoecological and evolutionary interpretations derived from Cenozoic microfaunal localities, a sample of 1883 bones accumulated by modern African fish eagles (H. vocifer) was collected from a roost near Lake Turkana, Kenya. Characteristics of the bone assemblage include (1) taxonomic composition dominated by fish; (2) diverse but small-sized and ecologically restricted animals including inshore fish, aquatic birds, very small nocturnal mammals (which may have been contributed by other raptors), and small reptiles (possible predators on the roost); (3) poor survivorship for specific categories of skeletal elements; (4) considerable bone breakage for all but the smallest animals; and (5) a distinctive pattern of postcranial fragmentation for bird and mammal elements. The fish eagle element profile has some similarities to those of other diurnal raptors, but its cumulative signature can be distinguished from those of other bone accumulators and includes characteristics that would also be manifest in fossil assemblages. Examination of fossil microfaunas from the Plio-Pleistocene localities of Kanapoi (Kenya), Inolelo (Tanzania), and Olduvai Gorge (Tanzania) shows that the patterns of element preservation for Kanapoi and Inolelo do not match those of the Lake Turkana fish eagle roost. Nonetheless, some aspects of the Olduvai Gorge FLKN assemblage show taxonomic and skeletal similarites. In light of the selective choices of prey by fish eagles, the fauna recovered from Olduvai Gorge FLKN might not accurately reflect faunal diversity or patterns of abundance at this locality during the Pleistocene. Because fish eagles of the genus Haliaeetus are found on three continents, have a fossil record extending to the Miocene, and deposit bones in a habitat conducive to preservation, the taxa preserved at many other late Cenozoic microfaunal sites also might be biased. Therefore, taphonomic profiles of late Cenozoic microfaunal assemblages should be compared against the taphonomic profiles typical of fish eagle roosts before faunal characterizations of sites are applied in analyses of paleoecological evolution.