Distinguishing the differential roles of hardpart-input rates and burial rates in the formation of shell beds is important in paleobiologic and sedimentologic studies, because high shelliness can reflect either high population density of shell producers or lack of sediment. The modeling in this paper shows that differences in the relative importance of burial rates and hardpart-input rates lead to distinct patterns with respect to the degree of shelliness and taphonomic alteration in shell beds. Our approach substantially complements other models because it allows computation of both shelliness and assemblage-level alteration. To estimate shelliness, we dissected hardpart-input rates into dead-shell production and shell destruction rates. To estimate assemblage-level alteration, we computed an alteration rate that describes how rapidly shells accrue postmortem damage. Under decreasing burial rates but constant hardpart-input rates, a positive correlation between alteration and shelliness is expected (Kidwell's R-sediment model). In contrast, under decreased destruction rates and/or increased dead-shell production rates and constant burial rates (Kidwell's R-hardpart model), a negative correlation between shelliness and alteration is expected. The contrasting predictions thus provide a theoretical basis for distinguishing whether high shell density in shell beds reflects passive shell accumulation due to a lack of sediment dilution or whether it instead reflects high shell input from a life assemblage. This approach should be applicable for any fossil assemblages that vary in shell density and assemblage-level alteration. An example from the Lower Jurassic of Morocco, which has shell-rich samples less altered than shell-poor samples, suggests that the higher shelliness correlates with higher community-level abundance and lower proportion of juveniles of the main shell producer, supporting the driving role of hardpart-input rates in the origin of the shell-rich samples in this case. This is of significance in paleoecologic analyses because variations in shelliness can directly reflect fluctuations in population density of shell producers.

Ecological investigations were conducted on rodent faunas in the Sonoran Desert of the southwestern United States and the Monte Desert of northwestern Argentina. Both study areas are physiognomically quite similar and the rodents are only distantly related. Such conditions are ideal for an assessment of possible evolutionary convergence. Multivariate analyses of morphoecological characters support the subjective interpretation that ecological equivalents between the deserts differ in their degree of similarity. The data indicate that the Monte rodent fauna is composed of two distinct groups: a) the caviomorph rodents, which are highly desert adapted; and b) the murids, which range widely beyond the desert's boundaries and are probably not as highly desert adapted, as a group, as are the caviomorphs. When the Monte and Sonoran faunas are compared with a North American coniferous forest rodent fauna, data show that both desert groups have converged toward one another. The different degrees of desert adaptation and different distributional patterns indicate that the Monte Desert was colonized in two different periods (one in the early Tertiary and one in the late Pliocene or early Pleistocene) and can be interpreted as supporting theories regarding South America's colonization by mammals which were first proposed by Simpson (1951).

Statistical inference about the upper and lower endpoints of the stratigraphic ranges of fossil taxa can be based on the pattern of finds. Strauss and Sadler (1989) described a test and confidence interval for a common upper or lower endpoint in two or more taxa. This approach is conservative, in the sense that it provides only an upper bound on the significance level. This paper describes and illustrates a test and confidence interval for which the significance level is known.

I examine the late Pleistocene megafaunal extinctions by testing the only extinction model with strong a priori predictions, the blitzkrieg model (Martin 1973; Mosimann and Martin 1975). I first test an assumption of the blitzkrieg and other extinction models that the megafaunal extinctions occurred in the terminal Wisconsin (12-10 Ka). This assumption has been disputed by Grayson (1989, 1991), but I find that both a reanalysis of Grayson's data and an analysis of new reliable data support a terminal Wisconsin extinction.

The blitzkrieg model predicts that the ranges of megafauna in North America were constricted as the semicircular front of hunters moved southeastward; hence the extinctions should be time-transgressive from northwest to southeast. I test this prediction in three separate analyses that examine (1) the location of terminal sites for each taxon relative to all their other late Wisconsin fossil sites, (2) the location of terminal sites for each taxon relative to all their other reliably dated late Wisconsin fossil sites, and (3) the spatio-temporal pattern of all the reliably dated terminal Wisconsin sites without regard to taxonomy. The geographic distribution of the megafaunal remains does not support the blitzkrieg hypothesis in any of the three analyses. Moreover, all of the patterns in the data are in a direction opposite to that predicted by the blitzkrieg hypothesis. I examine how these conclusions affect both climatic and predation models, particularly in relation to the testability of other extinction hypotheses.

Bones of mammals exhibit progressive stages of weathering during their time of subaerial exposure. Consequently, the study of bone weathering in fossil assemblages may help to assess the period represented by an accumulation of bones. Stages of bone decomposition due to subaerial weathering have been identified in assemblages of fossil macromammals from Olduvai Gorge, Tanzania. A modern bone assemblage collected by spotted hyenas is used to devise a method for recognizing attritional accumulations of bones from weathering characteristics. This method, which involves study of long bone diaphyses, is applied to Plio-Pleistocene faunal assemblages from Olduvai, 1.70–1.85 ma old. Previous work indicates that early hominids had an important role in the collection of fauna at five of the six sites studied. It is shown that animal bones were accumulated at each site over a period of probably 5–10 yr or more. The length of this period, along with other taphonomic evidence, suggests that the processes of bone aggregation at these sites differed from those at the short-term campsites of modern, tropical hunter-gatherers.

Sublethal injuries recorded in the shells of five Upper Mississippian ammonoid species from the Imo Formation of northwest Arkansas are manifested as repaired shell breaks, which can be categorized as minor, moderate, massive, deep-acute, or as perforations. Overall, 15% of the ammonoids exhibit some form of repaired break. The injuries are distributed as follows: Anthracoceras discus 9%; Fayettevillea bransoni 13%; Fayettevillea friscoense 21%; Rhadinites miseri 24%; Richardsonites mapesi 38%. These figures are substantially lower than in living Nautilus, in which more than one-half of adult specimens exhibit repaired shell breaks. The different frequencies of injuries may reflect species-selective predation, differential abilities to sustain and to repair shell damage, or they may be due to interspecific differences in physical parameters such as shell thickness and body chamber length. The most likely possible perpetrators of the injuries include sharks, other fishes, and cephalopods.

Neutral theory in ecology is based on the symmetry assumption that ecologically similar species in a community can be treated as demographically equivalent on a per capita basis—equivalent in birth and death rates, in rates of dispersal, and even in the probability of speciating. Although only a first approximation, the symmetry assumption allows the development of a quantitative neutral theory of relative species abundance and dynamic null hypotheses for the assembly of communities in ecological time and for phylogeny and phylogeography in evolutionary time. Although Steve Gould was not a neutralist, he made use of ideas of symmetry and of null models in his science, both of which are fundamental to neutral theory in ecology. Here I give a brief overview of the current status of neural theory in ecology and phylogeny and, where relevant, connect these newer ideas to Gould's work. In particular, I focus on modes of speciation under neutrality, particularly peripheral isolate speciation, and their implications for relative species abundance and species life spans. Gould was one of the pioneers in the study of neutral models of phylogeny, but the modern theory suggests that at least some of the conclusions from these early neutral models were premature. Modern neutral theory is a remarkably rich source of new ideas to test in ecology and paleobiology, the potential of which has only begun to be realized.

The continuous fossil record of therocephalian therapsids (Eutheriodontia) across the Permo-Triassic boundary and their differential survivorship of the end-Permian extinction offer an exceptional deep-time perspective on vertebrate life-history evolution during episodes of large-scale ecological perturbation. To examine potential impacts of the extinction on body size evolution (e.g., “Lilliput” effects) and growth patterns, we investigated cranial sizes and limb bone histology in the therocephalian Moschorhinus kitchingi both before and after the end-Permian extinction, facilitated by analysis of thin-sections of 23 limb bones from an ontogenetic sample of ten individuals across the Permo-Triassic boundary. In general, early subadult Moschorhinus displayed propodial cortices with extensive woven- and parallel-fibered bone (PFB) with dense radial and reticular vascularization and a moderately thickened bone wall with few growth marks. The outer cortex of propodials and epipodials showed a transition to PFB and lamellar bone with longitudinally oriented canals in individuals interpreted as late subadults or adults (>80% largest size). Most elements displayed several (3+) growth marks, though growth marks were more faithfully recorded in the epipodials of Permian individuals. Pearson product-moment correlation tests were performed to examine the relationship between size and robusticity on growth proxies (% cortical vascularity, mean primary osteon diameter), but variation in histomorphology could not be explained by size alone. Variation in body size may be affected by differences in juvenile growth rate and duration, which are highly variable in environmentally stressed extant reptile species. Geologic stage was a more consistent predictor of cortical vascularity. We suggest that Permian and Triassic Moschorhinus exhibited differential rates of early skeletal growth, corroborating the hypothesis that increased environmental variability in the earliest Triassic was associated with rapid growth to a minimum body size requirement and, consequently, shortened developmental times.

The most common methods of estimating the relative abundance of species in a fossil assemblage are all maximum likelihood estimates. They differ from one another in their inherent assumptions made about the effects of fragmentation and differential preservation in the assemblage. For many fossil assemblages, relative abundance is best estimated by the relative frequency of specimens or relative frequency of elements. Monte Carlo simulations suggest, however, that in most other circumstances estimates based on frequency of elements divided by the number of elements in a complete individual provide greater accuracy than estimates based on minimum number of individuals. This relation results from an interaction between random sampling error and a variety of biases inherent in the two estimates.

Marine planktonic microfossils have provided some of the best examples of evolutionary rates and patterns on multi-million-year time scales, including many instances of gradual evolution. Lineage splitting as a result of speciation has also been claimed, but all such studies have used subjective visual species discrimination, and interpretation has often been complicated by relatively small sample sizes and oceanographic complexity at the study sites. Here we analyze measurements on a collection of 10,200 individual tests of the Eocene planktonic foraminifer Turborotalia in 51 stratigraphically ordered samples from a site within the oceanographically stable tropical North Pacific gyre. We use novel multivariate statistical clustering methods to test the hypothesis that a single evolutionary species was present from 45 Ma to its extinction ca. 34 Ma. After identification of a set of biologically relevant traits, the protocol we apply does not require a prior assignment of individuals to species. We find that for most of the record, contemporaneous specimens form one morphological cluster, which we interpret as an evolving species that shows quasi-continuous but non-directional gradual evolutionary change (anagenesis). However, in the upper Eocene from ca. 36 to ca. 34 Ma there are two clusters that persistently occupy distinct areas of morphospace, from which we infer that speciation (cladogenesis) must have occurred.

We analyze relationships among a range of ecological and biological traits—geographic range size, body size, life mode, larval type, and feeding type—in order to identify those traits that are associated significantly with species duration in New Zealand Cenozoic marine molluscs, during a time of background extinction. Using log-linear modeling, we find that bivalves have only a small number of simple, two-way associations between the studied traits and duration. In contrast, gastropods display more complex interactions involving three-way associations between traits, a pattern that suggests greater macroecological complexity of gastropods. This is not an artifact caused by the larger number of gastropods than bivalves in our data set. We used stratified randomized resampling of families to test for associations between traits that might result from shared inheritance rather than ecological trait interactions; we found no evidence of phylogenetic effects in any associations examined. The relationships revealed by our study should serve to constrain the range of possible biological mechanisms that underlie these relationships. As previously observed, two-way associations are present between large geographic range and increased duration, and between large geographic range and large body size, in both bivalves and gastropods. In gastropods, planktotrophic larval type is associated with large range size through a three-way interaction that also involves duration; there is no direct association of larval type and geographic range. Gastropods also display two-way associations between duration and life mode, and duration and feeding type. We note that in gastropods, an infaunal life mode is associated with large range size, whereas in bivalves infaunality is associated with reduced range size.

Paleozoic and post-Paleozoic marine faunas are strikingly different in composition. Paleozoic marine gastropods may be divided into archaic and modern groups based on taxonomic composition, ecological role, and morphology. Paleozoic assemblages were dominated by pleurotomariids (Eotomariidae and Phymatopleuridae), the Pseudozygopleuridae, and, to a lesser extent, the Euomphalidae, while Triassic assemblages were dominated by the Trochiina, Amberleyacea, and new groups of Loxonematoidea and Pleurotomariina. Several new groups of caenogastropods appeared as well. Yet the importance of the end-Permian mass extinction in generating these changes has been questioned. As part of a study of the diversity history of upper Paleozoic and Triassic gastropods, to test the extent to which taxonomic and morphologic trends established in the late Paleozoic are continued after the extinction, and to determine the patterns of selectivity operating during the extinction, I assembled generic and morphologic diversity data for 396 genera in 75 families from the Famennian through the Norian stages. Within this interval, gastropod genera underwent an adaptive radiation during the Visean and Namurian, largely of pleurotomariids, a subsequent period of dynamic stability through the Leonardian, a broad-based decline during the end-Permian mass extinction, and a two-phase post-extinction rebound during the Triassic. The patterns of generic diversity within superfamily-level clades were analyzed using Q-mode factor analysis and detrended correspondence analysis.

The results demonstrate that taxonomic affinity, previous clade history, generic age, and gross morphology did not determine survival probability of genera during the end-Permian extinction, with the exception of the bellerophontids, nor did increasing diversity within clades or expansion of particular morphologies prior to the extinction facilitate survival during the extinction or success after it. The pleurotomariids diversified during the Lower Permian, but were heavily hit by the extinction. Similarly, trochiform and turriculate morphologies, among those which Vermeij (1987) has identified as having increased predation resistance, were expanding in the late Paleozoic, but suffered similar extinction rates to other nondiversifying clades. Survival was a consequence of broad geographic and environmental distribution, as was the case during background periods.

Calliostoma ligatum shells inhabited by hermit crabs were weaker than shells inhabited by snails collected at the same locality. When shells of C. ligatum were loaded repetitively to 80% of their predicted failure load, hermit crab-inhabited shells showed an immediate drop in shell strength followed by a progressive further loss of shell strength over the next 18 days. Snailinhabited C. ligatum shells exhibited a decrease in strength after 9 days, but returned to initial values within 18 days of loading.

Hermit crabs thus bear shells significantly weaker than they were when borne by the gastropods that produced them. Reported similarities in vulnerability of gastropods and hermit crabs to shell-crushing predators may be artifacts of the metric (critical size) used to compare vulnerabilities. Hermit crabs probably were a significant factor in the diversification of durophagous predators in the Mesozoic, supplying a prey base identical in size and shape to gastropods but with significantly lower resistance to crushing. The unpredictability of strength in hermit-crab inhabited shells may maintain the apparently inefficient indiscriminate attacks common among durophagous predators.

In modern ecosystems, regions of topographic heterogeneity, when compared with nearby topographically homogeneous regions, support high species densities of mammals and other groups. This biogeographic pattern could be explained by either greater diversification rates or greater accommodation of species in topographically complex regions. In this context, we assess the hypothesis that changes in landscape history have stimulated diversification in mammals. Landscape history includes tectonic and climatic processes that influence topographic complexity at regional scales. We evaluated the influence of changes in topographic complexity and climate on origination and extinction rates of rodents, the most diverse clade of mammals.

We compared the Neogene records of rodent diversity for three regions in North America. The Columbia Basin of the Pacific Northwest (Region 1) and the northern Rocky Mountains (Region 2) were tectonically active over much of the Cenozoic and are characterized by high topographic complexity today. The northern Great Plains (Region 3) have been tectonically quiescent, with low relief, throughout the Cenozoic. These three regions have distinctive geologic histories and substantial fossil records. All three regions showed significant changes in diversification and faunal composition over the Neogene. In the montane regions, originations and extinctions peaked at the onset and close, respectively, of the Miocene Climatic Optimum (17–14 Ma), with significant changes in faunal composition accompanying these episodes of diversification. In the Great Plains, rodents showed considerable turnover but infrequent diversification. Peak Neogene diversity in the Great Plains occurred during cooling after the Miocene Climatic Optimum. These histories suggest that climatic changes interacting with increasing topographic complexity intensify macroevolutionary processes. In addition, close tracking of diversity and fossil productivity with the stratigraphic record suggests either large-scale sampling biases or the mutual response of diversity and depositional processes to changes in landscape history.

For a survivorship curve to show a meaningful pattern, it is essential that a suitably homogeneous group is selected for analysis. If taxa have originated at different times in the geological record and have consequently experienced different extinction probabilities during the times that they existed, they do not constitute such a homogeneous group. A correction factor for variations in real-time extinction rates must be employed, as is described here. By application of the “corrected survivorship score,” it is shown that the Paleogene planktonic foraminifera exhibit a very strong survivorship pattern, namely an extinction probability that progressively increases with taxonomic longevity. That is to say, when extinction occurred, it was preferentially selective of older taxa. This is manifested as convexity in the survivorship curves. However, the pattern is degraded by variations in the real-time extinction rate, which causes straightening of the survivorship curves if they are calculated in the usual uncorrected way. If the law of constant extinction is to be tested for a given group, it is necessary either that stochastically constant real-time extinction rates are demonstrated or that variations in the extinction rate are corrected for in the manner described. As a separate issue, it is also essential that biologically meaningful taxa are analyzed. The convexity of planktonic foraminiferal survivorship curves is probably an artifact of the way evolving lineages have been subdivided in the application of typological taxonomy. Consequently, we are still a long way from being able to use survivorship analysis of planktonic foraminiferal data for adequately testing evolutionary models such as the Red Queen's hypothesis.

Studies of crinoid morphology have been pivotal in understanding the constraints on the range of morphology within a clade as well as the patterns of disparity throughout the Phanerozoic. Newly discovered and described faunas and recent study of early Paleozoic crinoid diversity provide an ideal opportunity to reanalyze Ordovician through Early Silurian crinoid disparity with more complete taxonomic coverage and finer stratigraphic resolution. Using the coarse stratigraphic binning of Foote (1999), the updated morphologic data set has a similar disparity pattern to those previously reported for the early Paleozoic. However, with the more resolved stratigraphic binning used by Peters and Ausich (2008), a significant difference exists between the original and current data sets. Both data sets have a pronounced disparity high during the late Middle Ordovician. However, the updated disparity curve has a much higher initial disparity during the Early Ordovician and a pronounced rise in disparity during the Silurian recovery. Examination of differential sampling, proportions of the crinoid orders through time, and methods of coding characters indicate these factors have little effect on the pattern of crinoid disparity. The Silurian morphospace expansion occurs primarily within disparids and coincides with the origination of the myelodactylids. These findings corroborate the rapid expansion of morphospace during the Ordovician. However, crinoid disparity did not remain static and, although less frequent than during the initial radiation, new body plans evolved following the Ordovician Extinction (e.g., the myelodactylids). These results are consistent with the hypothesis of ecology constraining the limits on morphologic disparity at the class level.

Mammalian generic, familial, and ordinal diversities correlate significantly with continental area. The area effect is similar in form to that shown for true islands: S = kAz, where S is the diversity, A is the area, and k and z are fitted constants. For mammalian genera and continental area, z equals 0.33, for families, z equals 0.23, and for orders, z equals 0.13.

The area effect permits quantitative modeling of extinction due to biotic competition between previously isolated faunas. The Late Cenozoic extinction of North and South American mammalian faunas following the rise of the Panamanian land bridge is overestimated by seven families. The overestimate may result from assumptions of complete biotic interchange and universal competition. The role of plate tectonics in regulating diversity may be extensively modified by regional environmental conditions.

Ediacaran fronds are key components of terminal-Proterozoic ecosystems. They represent one of the most widespread and common body forms ranging across all major Ediacaran fossil localities and time slices postdating the Gaskiers glaciation, but uncertainty over their phylogenetic affinities has led to uncertainty over issues of homology and functional morphology between and within organisms displaying this ecomorphology. Here we present the first large-scale, multigroup cladistic analysis of Ediacaran organisms, sampling 20 ingroup taxa with previously asserted affinities to the Arboreomorpha, Erniettomorpha, and Rangeomorpha. Using a newly derived morphological character matrix that incorporates multiple axes of potential phylogenetically informative data, including architectural, developmental, and structural qualities, we seek to illuminate the evolutionary history of these organisms. We find strong support for existing classification schema and devise apomorphy-based definitions for each of the three frondose clades examined here. Through a rigorous cladistic framework it is possible to discern the pattern of evolution within and between these clades, including the identification of homoplasies and functional constraints. This work both validates earlier studies of Ediacaran groups and accentuates instances in which previous assumptions of their natural history are uninformative.