Showing papers in "Paleobiology in 2010"

Seasonality, the latitudinal gradient of diversity, and Eocene insects

Abstract: In the modern world, biotic diversity is typically higher in low-latitude tropical regions where there is abundant insolation (light and heat) and low thermal seasonality. Because these factors broadly covary with latitude, separating their possible effects on species diversity is difficult. The Eocene was a much more equable world, however, with low temperature seasonality extending into lower-insolation higher, cooler latitudes, allowing us to test these factors by comparing insect species diversity in (1) modern, temperate, low-insolation, highly seasonal Harvard Forest, Massachusetts, U.S.A., 42°29′N; (2) modern, tropical, high-insolation, low-seasonality La Selva, Costa Rica, 10°26′N, and; (3) Eocene, temperate, low-insolation, yet low-seasonality McAbee, British Columbia, Canada, above 50°N paleolatitude. We found insect diversity at McAbee to be more similar to La Selva than to Harvard Forest, with high species richness of most groups and decreased diversity of ichneumon wasps, indicating that seasonality is key to the latitudinal diversity gradient. Further, midlatitude Eocene woody dicot diversities at McAbee, Republic (Washington, U.S.A.), and Laguna del Hunco (Argentina) are also high, similar to modern tropical samples, higher than at the modern midlatitude Harvard Forest. Modern correlations between latitude, species diversity, and seasonal climates were established some time after the Eocene.

On raptors and rodents: testing the ecological fidelity and spatiotemporal resolution of cave death assemblages

Abstract: Natural accumulations of skeletal remains represent a valuable source of ecological data for paleontologists and neontologists alike. Use of these records requires a quantitative assessment of the degree to which potential biasing factors affect how accurately ecological information from the living community is recorded in the sedimentary record. This has been a major focus in recent years for taphonomists working with marine records, yet terrestrial systems have remained virtually unstudied—particularly communities of small-bodied taxa. Our ability to assess the potential origins and effects of postmortem bias in terrestrial skeletal assemblages (both modern and fossil) has therefore been limited. Predation is a common mechanism by which small-mammal skeletal remains are concentrated; raptors regurgitate the remains of their small-mammal prey in pellets rich in skeletal material, which accumulate below long-term roosting sites, especially in protected areas such as caves and rock shelters. Here I compare small-mammal death assemblages concentrated via owl predation at Two Ledges Chamber, a long-term owl cave roost in northwestern Nevada, with data from modern trapping surveys to evaluate (1) their ecological fidelity to the modern small-mammal community, (2) the effects of temporal variation and time-averaging (over months to centuries) on live-dead agreement, and (3) how spatial averaging affects the landscape-scale picture of the small-mammal community as reconstructed from dead remains. Despite potential obstacles to the recovery of ecological information from skeletal deposits generated via predation, I find high live-dead agreement across all ecological metrics and all temporal comparisons. I also find that the effects of time-averaging (specifically increased species richness of the death assemblage) become significant only at the century scale. Finally, I combine a mixing model approach with a principal coordinates analysis to show that the owls at Two Ledges Chamber sample from all habitats present in the immediate vicinity of the cave, producing a high-fidelity snapshot of the community that is spatially integrated at the local landscape scale.

Completeness metrics and the quality of the sauropodomorph fossil record through geological and historical time

Abstract: Despite increasing concerns about the effect of sampling biases on our reading of the fossil record, few studies have considered the completeness of the fossil remains themselves, and those that have tend to apply non-quantitative measures of preservation quality. Here we outline two new types of metric for quantifying the completeness of the fossil remains of taxa through time, using sauropodomorph dinosaurs as a case study. The “Skeletal Completeness Metric” divides the skeleton up into percentages based on the amount of bone for each region, whereas the “Character Completeness Metric” is based on the number of characters that can be scored for each skeletal element in phylogenetic analyses. For both metrics we calculated the completeness of the most complete individual and of the type specimen. We also calculated how well the taxon as a whole is known from its remains. We then plotted these results against both geological and historical time, and compared curves of the former with fluctuations in sauropodomorph diversity, sea level, and sedimentary rock outcrop area. Completeness through the Mesozoic shows a number of peaks and troughs; the Early Jurassic (Hettangian–Sinemurian) is the interval with highest completeness, whereas the mid-to-Late Cretaceous has completeness levels that are consistently lower than the rest of the Mesozoic. Completeness shows no relationship to rock outcrop area, but it is negatively correlated with sea level during the Jurassic–Early Cretaceous and correlated with diversity in the Cretaceous. Completeness of sauropodomorph type specimens has improved from 1830 to the present, supporting the conclusions of other recent studies. However, when this time interval is partitioned, we find no trend for an increase in completeness from the 1990s onward. Moreover, the 2000s represent one of the poorest decades in terms of average type specimen completeness. These results highlight the need for quantitative methods when assessing fossil record quality through geological time or when drawing conclusions about historical trends in the completeness of taxa. The new metrics may also prove useful as sampling proxies in diversity studies.

Predicting the effects of increasing temporal scale on species composition, diversity, and rank-abundance distributions

Abstract: Paleoecological analyses that test for spatial or temporal variation in diversity must consider not only sampling and preservation bias, but also the effects of temporal scale (ie, time- averaging) The species-time relationship (STR) describes how species diversity increases with the elapsed time of observation, but its consequences for assessing the effects of time-averaging on diversity of fossil assemblages remain poorly explored Here, we use a neutral, dispersal-limited model of metacommunity dynamics, with parameters estimated from living assemblages of 31 molluscan data sets, to model the effects of within-habitat time-averaging on the mean composition and multivariate dispersion of assemblages, on diversity at point (single station) and habitat scales (pooled multiple stations), and on beta diversity We hold sample size constant in STRs to isolate the effects of time-averaging from sampling effects With increasing within-habitat time-averaging, stochastic switching in the identity of species in living (dispersal-limited) assemblages (1) decreases the proportional abundance of abundant species, reducing the steepness of the rank-abundance distribution, and (2) increases the proportional richness of rare, temporally short-lived species that immigrate from the neutral metacommunity with many rare species These two effects together (1) can shift the mean composition away from the non-averaged (dispersal-limited) assemblages toward averaged assemblages that are less limited by dispersal, resembling that of the metacommunity; (2) allow the point and habitat diversity to increase toward metacommunity diversity under a given sample size (ie, the diversity in averaged assemblages is inflated relative to non-averaged assemblages); and (3) reduce beta diversity because species unique to individual stations become shared by other stations when limited by a larger but static species pool Surprisingly, these scale- dependent changes occur at fixed sample sizes and can become significant after only a few decades or centuries of time-averaging, and are accomplished without invoking ecological succession, environmental changes, or selective postmortem preservation Time-averaging results in less inflation of diversity at habitat than at point scales; paleoecological studies should thus analyze data at multiple spatial scales, including that of the habitat where multiple bulk samples have been pooled in order to minimize time-averaging effects The diversity of assemblages that have accumulated over 1000 years at point and habitat scales is expected to be inflated by an average of 21 and 16, respectively This degree of inflation is slightly higher than that observed in molluscan death assemblages at these same spatial scales (18 and 13) Thus, neutral metacommunity models provide useful quantitative constraints on directional but predictable effects of time-averaging They provide minimal estimates for the rate of increase in diversity with time-averaging because they assume no change in environmental conditions and in the composition of the metacommunity within the window of averaging

Quantitative comparisons and models of time-averaging in bivalve and brachiopod shell accumulations

Abstract: The variation in time-averaging between different types of marine skeletal accumulations within a depositional system is not well understood. Here we provide quantitative data on the magnitude of time-averaging and the age structure of the sub-fossil record of two species with divergent physical and ecological characteristics, the brachiopod Bouchardia rosea and the bivalve Semele casali. Material was collected from two sites on a mixed carbonate-siliciclastic shelf off the coast of Brazil where both species are dominant components of the local fauna. Individual shells (n = 178) were dated using amino acid racemization (aspartic acid) calibrated with 24 AMS radiocarbon dates. Shell ages range from modern to 8118 years b.p. for brachiopods, and modern to 4437 years for bivalves. Significant differences in the shape and central tendency of age-frequency distributions are apparent between each sample. Such differences in time-averaging magnitude confirm the assumption that taphonomic processes are...

Comparison of morphometric techniques for shapes with few homologous landmarks based on machine-learning approaches to biological discrimination

Abstract: Biometric analyses are useful tools for the study of organisms, their phylogenetic affiliation, and the pattern and rate of their evolution. Various morphometric techniques have been developed to analyze morphological variation, but methodological choices are often made arbitrarily because quantitative comparisons are lacking or inconclusive. Here we address morphometric quantification of taxa with few unambiguously identifiable landmarks (<15), utilizing ornamented and unornamented gastropod shells. Support vector machines were applied to evaluate classification performances of landmark (LMA), elliptic Fourier (EFA), and semi-landmark analysis (SLM). This evaluation is based on the discrimination of between-group differences relative to within-group variation, and thus allows comparing how the techniques treat different types of biological information. The results suggest that EFA performs slightly better than SLM (and certainly LMA) in discerning a priori identified taxa with unornamented shells, but that SLM is significantly superior to other techniques for ornamented shells. Alignment and homology problems may cause the subtle variations in ornamentation to become blurred as noise in EFA, even though EFA is often cited to be able to deal with complex shapes. Performance of LMA depends entirely on how accurately the structure can be covered with landmarks. Guidelines in choosing a morphometric technique in diverse cases are provided.

A quantitative analysis of environmental associations in sauropod dinosaurs

Abstract: Both the body fossils and trackways of sauropod dinosaurs indicate that they inhabited a range of inland and coastal environments during their 160-Myr evolutionary history. Quantitative paleoecological analyses of a large data set of sauropod occurrences reveal a statistically significant positive association between non-titanosaurs and coastal environments, and between titanosaurs and inland environments. Similarly, "narrow-gauge" trackways are positively associated with coastal environments and "wide-gauge" trackways are associated with inland environments. The statistical support for these associations suggests that this is a genuine ecological signal: non-titanosaur sauropods preferred coastal environments such as carbonate platforms, whereas titanosaurs preferred inland environments such as fluvio-lacustrine systems. These results remain robust when the data set is time sliced and jackknifed in various ways. When the analyses are repeated using the more inclusive groupings of titanosauriforms and Macronaria, the signal is weakened or lost. These results reinforce the hypothesis that "wide-gauge" trackways were produced by titanosaurs. It is commonly assumed that the trackway and body fossil records will give different results, with the former providing a more reliable guide to the habitats occupied by extinct organisms because footprints are produced during life, whereas carcasses can be transported to different environments prior to burial. However, this view is challenged by our observation that separate body fossil and trackway data sets independently support the same conclusions regarding environmental preferences in sauropod dinosaurs. Similarly, analyzing localities and individuals independently results in the same environmental associations. We demonstrate that conclusions about environmental patterns among fossil taxa can be highly sensitive to an investigator's choices regarding analytical protocols. In particular, decisions regarding the taxonomic groupings used for comparison, the time range represented by the data set, and the criteria used to identify the number of localities can all have a marked effect on conclusions regarding the existence and nature of putative environmental associations. We recommend that large data sets be explored for such associations at a variety of different taxonomic and temporal scales.

Carcasses on the coastline: measuring the ecological fidelity of the cetacean stranding record in the eastern North Pacific Ocean

Abstract: To understand how well fossil assemblages represent original communities, paleoecologists seek comparisons between death assemblages and their source communities. These comparisons have traditionally used nearshore, marine invertebrate assemblages for their logistical ease, high abundance, and comparable census data from living communities. For large marine vertebrates, like cetaceans, measuring their diversity in ocean ecosystems is difficult and expensive. Cetaceans, however, often beach or strand themselves along the coast, and archived data on stranded cetaceans have been recorded, in some areas, over several decades. If the stranding record is interpreted as a death assemblage, then the stranding record may represent a viable alternative for measuring diversity in living communities on directly adjacent coastlines. This study assessed the fidelity of the cetacean stranding record in the eastern North Pacific Ocean. The living community in this region has been studied for over 100 years and, recently, extensive and systematic live transect surveys using ship-based observing platforms have produced a valuable source of live diversity data. Over this same period, the U.S. Marine Mammal Stranding Program has collected and archived a record of cetacean strandings along the U.S. Pacific coastline, providing an ideal death assemblage for comparison. Using fidelity metrics commonly used in marine invertebrate taphonomy, I determined that the stranding record samples the living cetacean community with high fidelity, across fine and coarse taxonomic ranks, and at large geographic scales (>1000 km of coastline). The stranding record is also richer than the live surveys, with live-dead ratios between 1.1 and 1.3. The stranding record recovers similar rank-order relative abundances as live surveys, with statistical significance. Also, I applied sample-based rarefaction methods to generate collector's curves for strandings along the U.S. Pacific Coast to better evaluate the spatiotemporal characteristics of the stranding record. Results indicate that saturation (i.e., sampling >95% assemblage) at species, genus, and family levels occurs in less than five years of sampling, with families accumulating faster than species, and larger geographic regions (i.e., longer coastlines) accumulating taxa the most rapidly. The high fidelity of the stranding record, measured both in richness and by ranked relative abundance, implies that ecological structure from living cetacean communities is recorded in the death assemblage, a finding that parallels marine invertebrate assemblages, though at far larger spatial scales. These results have implications for studying cetacean ecology in both modern and ancient environments: first, these results imply that the stranding record, over sufficiently long time intervals, yields a richer assemblage than using line-transect methods, and faithfully records aspects of community structure; and second, these results imply that geochronologically well-constrained fossil cetacean assemblages might preserve ecologically relevant features of community structure, depending on depositional and taphonomic conditions.

An Australian land mammal age biochronological scheme

Abstract: Constrained seriation of a species-locality matrix of the Australian Cenozoic mammal record resolves a preliminary sixfold succession of land mammal ages apparently spanning the late Oligocene to the present. The applied conditions of local chronostratigraphic succession and inferences of relative stage-of-evolution biochronology lead to the expression of a continental geological timescale consisting of, from the base, the Etadunnan, Wipajirian, Camfieldian, Waitean, Tirarian, and Naracoortean land mammal ages. Approximately 99% of the 360 fossil assemblages analyzed are classifiable using this method. Each is characterized by a diagnostic suite of species. An interval of age magnitude may eventually be shown to lie between the Camfieldian and Waitean, but is currently insufficiently represented by fossils to diagnose. Development of a land mammal age framework marks a progressive step in Australian vertebrate biochronology, previously expressed only in terms of local faunas. Overall, however, the record remains poorly calibrated to the Standard Chronostratigraphic Scale. Codifying the empirical record as a land mammal age sequence provides an objective basis for expressing faunal succession without resort to standard chronostratigraphic terms with the attendant (and hitherto commonly taken) risks of miscorrelating poorly dated Australian events to well-dated global events.

Origins of microfossil bonebeds: insights from the Upper Cretaceous Judith River Formation of north-central Montana

Abstract: Microfossil bonebeds are multi-individual accumulations of disarticulated and dissociated vertebrate hardparts dominated by elements in the millimeter to centimeter size range (≥75% of bioclasts ≤5 cm maximum dimension). Modes of accumulation are often difficult to decipher from reports in the literature, although predatory (scatological) and fluvial/hydraulic origins are typically proposed. We studied the sedimentology and taphonomy of 27 microfossil bonebeds in the Campanian Judith River Formation of Montana in order to reconstruct formative histories. Sixteen of the bonebeds examined are hosted by fine-grained facies that accumulated in low-energy aquatic settings (pond/lake microfossil bonebeds). Eleven of the bonebeds are embedded in sandstones that accumulated in ancient fluvial settings (channel-hosted microfossil bonebeds). In lieu of invoking separate pathways to accumulation based on facies distinctions, we present a model that links the accumulation of bioclasts in the two facies. We p...

The geological completeness of paleontological sampling in North America

Abstract: A growing body of work has quantitatively linked many macroevolutionary patterns, including short- and long-term changes in biodiversity, rates of taxonomic extinction and origination, and patterns of extinction selectivity, to temporal variability in the sedimentary rock record. Here we establish a new framework for more rigorously testing alternative hypotheses for these and many other results by documenting the large-scale spatiotemporal intersection of the North American sedimentary rock and fossil records. To do this, we combined 30,387 fossil collections in the spatially explicit Paleobiology Database with a comprehensive macrostratigraphic database consisting of 18,815 sedimentary lithostratigraphic units compiled from 814 geographic regions distributed across the United States and Canada. The geological completeness of paleontological sampling, here defined as the proportion of the available sedimentary rock record that has been documented to have at least one fossil occurrence, irrespective of taxonomy or environment, is measured at four different levels of stratigraphic resolution: (1) lithostratigraphic rock units, (2) hiatus-bound rock packages, (3) regional stratigraphic columns, and (4) sediment coverage area (km2). Mean completeness estimates for 86 Phanerozoic time intervals (approximately stages; median duration 5.3 Myr) range from 0.18 per interval in the case of lithostratigraphic rock units to 0.23 per interval for stratigraphic columns and sediment coverage area. Completeness estimates at all four levels of stratigraphic resolution exhibit similar temporal variation, including a significant long-term increase during the Phanerozoic that is accentuated by an abrupt Campanian–Maastrichtian peak. This Late Cretaceous peak in completeness is approximately five times greater than the least complete Phanerozoic time intervals (Early Cambrian, Early Devonian, late Permian, and Early Cretaceous). Geological completeness in the Cenozoic is, on average, approximately 40% greater than in the Paleozoic. Temporal patterns of geological completeness do not appear to be controlled exclusively by variation in the frequency of subsurface rock units or an increase over time in the proportion of terrestrial rock, but instead may be general features of both the marine and terrestrial fossil records.

Dietary interpretation and paleoecology of herbivores from Pikermi and Samos (late Miocene of Greece)

Abstract: A large sample of the Pikermi and Samos ungulates was examined by microwear analysis using a light stereomicroscope (561 extinct and 809 extant comparative specimens). The results were used to infer the dietary adaptations of individual species and to evaluate the Pikermian Biome ungulate fauna. Many of the bovids have wear consistent with mixed feeding, although a few mesodont taxa apparently enjoyed an exclusive browsing and or grazing diet. The giraffids spanned the entire dietary spectrum of browsing, mixed feeding, and grazing, but most of the three-toed horses (Hippotherium) were hypsodont grazers. The colobine monkey Mesopithecus pentelici displays microwear consistent with a mixed fruit and leaf diet most likely including some hard objects. Similar results were obtained from prior scanning electron microscopy microwear studies at 500 times magnification and from the light microscope method at 35 times magnification for the same species. Results show that diet can differ between species that have very similar gross tooth morphology. Our results also suggest that the Pikermian Biome was most likely a woodland mosaic that provided a diversity of opportunities for species that depended on browsing as well as species that ate grass. The grasses were most likely C3 grasses that would grow in shaded areas of the woodland, glades, and margins of water. The ungulate component of the Pikermi and Samos fauna was more species-rich and more diverse in diet than the ungulates observed in modern African forests, woodlands, or savannas, yet dietarily most similar to the ungulates found in woodland elements of India and to some extent of Africa. It is unlikely that the Pikermi and Samos ungulates inhabited dense forests because we find no evidence for heavy fruit browsing. Conversely, a pure savanna is unlikely because many mixed feeders are present as well as browsers. Extant woodland African species are morphologically and trophically very similar to the African savanna species. Therefore the evolution of grazing and of hypsodont morphology for Africa may have evolved within the Plio-Pleistocene woodlands of Africa. Our results show that major dietary and morphologic ungulate evolution may take place within woodlands rather than as a consequence of species moving into savannas both during the late Miocene of Pikermi and Samos and during the Pleistocene-Recent of Central Africa.

A physiologically explicit morphospace for tracheid-based water transport in modern and extinct seed plants

Abstract: We present a morphometric analysis of water transport cells within a physiologically explicit three-dimensional space. Previous work has shown that cell length, diameter, and pit resistance govern the hydraulic resistance of individual conducting cells; thus, we use these three parameters as axes for our morphospace. We compare living and extinct plants within this space to investigate how patterns of plant conductivity have changed over evolutionary time. Extinct coniferophytes fall within the range of living conifers, despite differences in tracheid-level anatomy. Living cycads, Ginkgo biloba, the Miocene fossil Ginkgo beckii, and extinct cycadeoids overlap with both conifers and vesselless angiosperms. Three Paleozoic seed plants, however, occur in a portion of the morphospace that no living seed plant occupies. Lyginopteris, Callistophyton, and, especially, Medullosa evolved tracheids with high conductivities similar to those of some vessel-bearing angiosperms. Such fossils indicate that extinct seed plants evolved a structural and functional diversity of xylem architectures broader, in some ways, than the range observable in living seed plants.

Global occurrence trajectories of microfossils: environmental volatility and the rise and fall of individual species

Abstract: Species arise and establish themselves over the geologic time scale. This process is manifested as a change in the relative frequency of occurrences of a given species in the global pool of species. Our main goal here is to model this rise and the eventual decline of microfossil species using a mixed-effects model where groups each have a characteristic occurrence trajectory (main effects) and each species belonging to those groups is allowed to deviate from the given group trajectory (random effects). Our model can be described as a “hat” with logistic forms in the periods of increase and decline. Using the estimated timings of rises and falls, we find that the lengths of the periods of rise are about as long as the lengths of the periods when species are above 50% of their estimated maximal occurrences. These latter periods are here termed periods of dominance, which are in turn about the same length as the species' periods of fall. The peak rates of the rises of microfossils are in general faster than their peak rates of falls. These quantified observations may have broad macroevolutionary and macroecological implications. Further, we hypothesize that species that have experienced and survived high levels of environmental volatility (specifically, periods of greater than average variation in temperature and productivity) during their formative periods should have longer periods of dominance. This is because subsequent environmental variations should not drive them to decline with ease. We find that higher estimated environmental volatility early in the life of a species positively correlates with lengths of periods of dominance, given that a species survives the initial stress of the environmental fluctuations. However, we find no evidence that the steepness of the rise of a species is affected by environmental volatility in the early phases of its life.

Integrating phylogeny, molecular clocks, and the fossil record in the evolution of coralline algae (Corallinales and Sporolithales, Rhodophyta)

Abstract: When assessing the timing of branching events in a phylogeny, the most important tools currently recognized are a reliable molecular phylogeny and a continuous, relatively complete fossil record. Coralline algae (Rhodophyta, Corallinales, and Sporolithales) constitute an ideal group for this endeavor because of their excellent fossil record and their consistent phylogenetic reconstructions. We present the evolutionary history of the corallines following a novel, combined approach using their fossil record, molecular phylogeny (based on the 18S rDNA gene sequences of 39 coralline species), and molecular clocks. The order of appearance of the major monophyletic taxa of corallines in the fossil record perfectly matches the sequence of branching events in the phylogeny. We were able to demonstrate the robustness of the node ages in the phylogeny based on molecular clocks by performing an analysis of confidence intervals and maximum temporal ranges of three monophyletic groups of corallines (the families Sporolithaceae and Hapalidiaceae, as well as the subfamily Lithophylloideae). The results demonstrate that their first occurrences are close to their observed appearances, a clear indicator of a very complete stratigraphic record. These chronological data are used to confidently constrain the ages of the remaining branching events in the phylogeny using molecular clocks.

Information landscapes and sensory ecology of the Cambrian Radiation

Abstract: Organisms emit, detect, and respond to a huge array of environmental signals. The distribution of a given signal is dependent, first of all, upon the original spatial distribution of signal sources, the source landscape. The signal sources can be fixed or moving and their output can be stable or ephemeral. Different sources can also occupy the same general spatial location, such as insects living on a host plant. The emitted signals are modified by relevant transport processes, which are often strongly scale and environment dependent. Chemical signals, for example, are propagated by diffusion and turbulence. The resulting complex, three-dimensional, and dynamic distribution of signals in the environment is the signal landscape; it is the environment of potentially available information in which sensory systems function and have evolved. Organisms also differ widely in what signals they can actually detect; the distribution of signals that an organism can potentially respond to is its information landscape. Although increasing the kinds and specificity of signals that can be detected and processed can lead to improved decision making, it almost always comes at an increased cost. The greater the spatial and temporal complexity of the environment, the greater are the costs of incomplete information and the more advantageous is the development of improved information-gathering capabilities. Studies with simulation models suggest how variability in the spatial structure of source and signal landscapes may control patterns of animal movement that could be represented in the trace fossil record. Information landscapes and the corresponding sensory systems should have evolved in concert with major transitions in the history of life. The Ediacaran to Cambrian interval is one of the most intensively studied periods in the history of life, characterized by the profound environmental and biological changes associated with the bilaterian radiation. These include the advent of macroscopic predation, an increase in the size and energy content of organisms, and the transition in seafloors from laminated matgrounds to mixgrounds produced by the development of macroscopic infaunal bioturbation. The overall effect of these transitions was to markedly increase the spatial complexity of the marine environment. We suggest that this increased spatial complexity, in turn, drove the evolution of macroscopic sense organs in mobile bilaterians, leading to their first appearance during the Cambrian. The morphology and distribution of these sense organs should reflect the life habits of the animals that possessed them. Our overall hypothesis was that there was a “Cambrian Information Revolution,” a coevolutionary increase in the information content of the marine environment and in the ability of and necessity for organisms to obtain and process this information. A preliminary analysis of the Maotianshan Shale (Chengjiang) biota indicates that the distribution of eyes and antennae in these animals is consistent with predictions based on their life habit.

Multiple paleoecological controls on the composition of marine fossil assemblages from the Frasnian (Late Devonian) of Virginia, with a comparison of ordination methods

Abstract: Ecological ordination can reveal gradients in the species composition of fossil assemblages that can be correlated with paleoenvironmental gradients. Ordinations of simulated data sets suggest that nonmetric multidimensional scaling (NMDS) generally produces less distorted results than detrended correspondence analysis (DCA). We ordinated 113 brachiopod-dominated samples from the Frasnian (Late Devonian) Brallier, Scherr, and lower Foreknobs Formations of southwest Virginia, which represent a range of siliciclastic marine paleoenvironments. A clear environmental signal in the ordination results was obscured by (apparently) opportunistic species that occurred at high abundance in multiple environments; samples dominated by these species aggregated in ordination space regardless of paleoenvironmental provenance. After the opportunist-dominated samples were removed, NMDS revealed a gradient in species composition that was highly correlated with substrate (grain size); a second, orthogonal gradient likely reflects variation in disturbance intensity or frequency within grain-size regimes. Additional environmental or ecological factors, such as oxygenation, may also be related to the gradients. These two gradients, plus the environmental factors that controlled the occurrence of opportunistic species, explain much of the variation in assemblage composition in the fauna. In general, the composition of fossil assemblages is probably influenced by multiple paleoecological and paleoenvironmental factors, but many of these can be decomposed and analyzed.

Fishes on coral reefs: changing roles over the past 240 million years

Abstract: Key morphological traits reveal changes in functional morphospace occupation of reef fish assemblages over time. We used measurements of key functional attributes (i.e., lower jaw length and orbit diameter) of 208 fossil fish species from five geological periods to create bivariate plots of functional morphological traits through time. These plots were used to examine possible function and ecological characteristics of fossil reef fish assemblages throughout the Mesozoic and Cenozoic. A previously unknown trend of increasing orbit diameter over time became apparent. The Teleostei are the principal drivers of this change. The Eocene appears to mark a dramatic increase in two previously rare feeding modes in fishes: nocturnal feeding and high-precision benthic feeding. Interestingly, members of the Pycnodontiformes had relatively large eyes since the Triassic and appear to be the ecological precursors of their later teleost counterparts and may have been among the earliest nocturnal feeding fishes. Our results highlight potential changes in the roles of fishes on coral reefs through time.

Constraints on clade ages from fossil outgroups

Abstract: This paper presents a method for constraining the age of a clade with the ages of the earliest fossil specimens in that clade's outgroups. Given a sufficiently deep, robust, well-resolved, and stratigraphically consistent cladogram, this method can yield useful age constraints even in the absence of specific information about the fossil preservation and recovery rates of individual taxa. The algorithm is applied to simulated data sets to demonstrate that this method can yield robust constraints of clade ages if there are sufficient fossil outgroups available and if there is a finite chance that additional outgroups may be discovered in the future. Finally, the technique is applied to actual fossil data to explore the origin of modern placental mammals. Using data from recently published cladograms, this method indicates that if all Mesozoic eutherians are regarded as outgroups of Placentalia, then the last common ancestor of modern placental mammals and their Cenozoic allies lived between 65 and 88-98 million years ago, depending on the assumed cladogram and the number of outgroups included in the analysis.

Depth-habitat reorganization of planktonic foraminifera across the Albian/Cenomanian boundary

Abstract: New mid-Cretaceous stable isotope (δ18O and δ13C) records of multiple planktonic foraminiferal species and coexisting coccoliths from Blake Nose (western North Atlantic) document a major depth-ecology reorganization of planktonic foraminifera. Across the Albian/Cenomanian boundary, deep-dwelling Praeglobotruncana stephani and Rotalipora globotruncanoides adapted to living at a shallower depth, while, at the same time, the population of surface-dwelling Paracostellagerina libyca declined. Subsequently, the opportunistic species Hedbergella delrioensis shifted to a deep environment, and the deep-dwelling forms Rotalipora montsalvensis and Rotalipora reicheli first appeared. The primary paleoenvironmental cause of the observed changes in planktonic adaptive strategies is uncertain, yet their coincidence with an earliest Cenomanian cooling trend reported elsewhere implicates the importance of reduced upper-ocean stratification. Although there has been an implicit assumption that the species-specific depth habitats of fossil planktonic foraminifera were invariant through time, planktonic paleoecology is a potential variable. Accordingly, the possibility of evolutionary changes in planktonic foraminiferal depth ecology should be a primary consideration (along with other environmental parameters) in paleoceanographic interpretations of foraminiferal stable isotope data.

The completeness of the continental fossil record and its impact on patterns of diversification

Abstract: Did organisms diversify in different ways on land and in the marine realm over the Phanerozoic, or do the different diversification curves of continental and marine organisms reflect primarily methodological artifacts? To answer this question, a quantitative assessment of the completeness of the global continental fossil record is indispensable. We used comparisons between continental and marine fossil diversity and between past and present-day patterns of continental diversity to assess the absolute and relative completeness of the continental fossil record. Collector's curves of the number of described families over the past 200 years suggest that the global continental fossil record, and even that of European and North American tetrapods, is still highly incomplete. Nevertheless, relative proportions of major continental and marine taxa, patterns of tetrapod endemism, and familial durations suggest that the family-level continental fossil record is reasonably representative. We found that, although continental fossil richness is correlated with the amount of terrestrial clastic sediment available for sampling, the exponential diversification curve of continental metazoans is unlikely to be an artifact of this rock bias. Diversification of the continental fauna appears to have been essentially exponential since the Devonian, with little evidence of major extinction events.

Occupancy, range size, and phylogeny in Eurasian Pliocene to Recent large mammals

Abstract: Temporal patterns in species occupancy and geographic range size are a major topic in evolutionary ecology research. Here we investigate these patterns in Pliocene to Recent large mammal species and genera in Western Eurasia. By using an extensively sampled fossil record including some 700 fossil localities, we found occupancy and range size trajectories over time to be predominantly peaked among both species and genera, meaning that occupancy and range size reached their maxima midway along taxon existence. These metrics are strongly correlated with each other and to body size, after phylogeny is accounted for by using two different phylogenetic topologies for both species and genera. Phylogenetic signal is strong in body size, and weaker but significant in both occupancy and range size mean values among genera, indicating that these variables are heritable. The intensity of phylogenetic signal is much weaker and often not significant at the species level. This suggests that within genera, occupancy and range size are somewhat variable. However, sister taxa inherit geographic position (the center of their geographic distribution). Taken together, the latter two results indicate that sister species occupy similar positions on the earth's surface, and that the expansion of the geographic range during the existence of a given genus is driven by range expansion of one or more of the species it includes, rather than simply being the summation of these species ranges.

Upper Paleolithic Levels XIII-VI (A and B) from the 1937-1938 and 1947-1948 Boston College Excavations and the Levantine Aurignacian at Ksar Akil, Lebanon

Abstract: L’abri sous-roche de Ksar Akil (Liban) presente la plus longue sequence connue pour le Paleolithique superieur du Levant. Les recherches sur ce site ont commence au debut des annees 1920 et ont dure pratiquement 90 ans. Cet article concerne la partie superieure de la sequence stratigraphique qui inclut l’Aurignacien du Levant et il presente la premiere tentative de correlation entre les donnees des campagnes de 1937-1938 et de 1947-1948 conduites par l’universite de Boston. De maniere generale, les divergences portent non pas tant sur la description des assemblages que sur leur interpretation, tout particulierement sur les regroupements de niveaux, la designation des phases et leur affiliation culturelle. Un point important concerne la caracterisation de l’Aurignacien du Levant. Chacun des niveaux etudies dans cet article comprend de nombreuses lames et lamelles, ainsi que l’avait souligne J. Tixier lors de ses fouilles en 1969-1975. Ceci concerne notamment la phase 5 de Ksar Akil, generalement associee a l’Aurignacien « classique » ; cette industrie a ete decrite, dans les autres parties du Levant, comme orientee vers la production d’eclats.

Biotic influences on species duration: interactions between traits in marine molluscs

Abstract: 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.

Climate-averaging of terrestrial faunas: an example from the Plio-Pleistocene of South Africa

Abstract: Fundamental to the interpretation of bone-bearing faunal deposits is an understanding of the taphonomic processes that have modified the once living fossil community. An often neglected source of bias is that of climate-averaging, which occurs when the duration of bone accumulation exceeds the duration of an individual climatic episode. Tropical and subtropical climate change is dominated by precessional cyclicity (,21,000 year cycle), which controls monsoon rainfall intensity and thus plant communities over time. Under a climate-averaging scenario, the paleoecological characteristics of a faunal deposit represent an amalgamation of more than one phase of the precessional cycle. We investigate the degree of climate-averaging in Plio-Pleistocene bone breccias from South Africa by comparing stable isotope measurements of fossil enamel with the evidence from high-resolution speleothem paleoclimate proxies. We conclude that each of the four faunal assemblages studied are climate-averaged, having formed over a time period in excess of one-third of a precessional cycle (,7000 years). This has implications for the reconstruction of hominin paleoenvironments and estimates of Plio-Pleistocene biodiversity. We hypothesize that climate- averaging may be a common feature of tropical terrestrial vertebrate assemblages throughout the Cenozoic and Mesozoic.

The scale of it all: postcanine tooth size, the taxon-level effect, and the universality of Gould's scaling law

Abstract: In a seminal paper in 1975, Gould proposed that postcanine occlusal area (PCOA) should scale metabolically (0.75) with body mass across mammals. By regressing PCOA against skull length in a small sample of large-bodied herbivorous mammals, Gould provided some marginal support for this hypothesis, which he then extrapolated as a universal scaling law for Mammalia. Since then, many studies have sought to confirm this scaling relationship within a single order and have found equivocal support for Gould's assertion. In part, this may be related to the use of proxies for both PCOA and body mass, small sample sizes, or the influence of a “taxon-level effect,” rendering Gould's scaling “universal” problematic. Our goal was to test the universality of Gould's prediction and the impact of the taxon-level effect on regressions of tooth size on body mass in a large extant mammalian sample (683 species spanning 14 orders). We tested for the presence of two types of taxon-level effect that may influence the a...

The living, the dead, and the expected dead: variation in life span yields little bias of proportional abundances in bivalve death assemblages

Abstract: All else being equal, species with short life spans are expected to be overrepresented in time-averaged death assemblages relative to their standing abundance in the living community, but the magnitude of the distortion of proportional abundance and assemblage evenness has received little attention. Here, information from 30 data sets on the living and dead abundances of marine bivalves in local habitats is combined with a global compilation of bivalve life spans to determine whether bias from mortality rate can explain observed differences in species proportional abundances. Although bivalve maximum life spans range from one to 75 years in these data sets, indicating annual mortality rates of 0.97 to 0.09, the “life span bias” (LB) of a species—the difference between its proportional abundance expected dead and that observed alive—is consistently small in magnitude (average change <2%, maximum about 20%) and random in sign relative to observed discordance (OD = difference between that species' proportional abundance observed dead and that observed alive). The aggregate result for 413 living species occurrences is a significantly positive but weak correlation of OD to LB, with only 10% of variation in OD explained. The model performs better among longer-lived species than among shorter-lived species, probably because longer-lived species conform better to the model assumption that species maintain a constant proportional abundance in the living assemblage over time. Among individual data sets, only seven exhibit significant positive correlations between OD and LB. The model also under-predicts the cases where a death assemblage is dominated by a species that is shorter lived than the dominant species in the living assemblage, indicating that some factor(s) other than or in addition to mortality rate is responsible for OD. We can find no evidence of preservational bias linked to life span, for example through body size. This negative outcome reflects a weak biological relationship between life span and living abundance among bivalves in local habitats, contrary to the terrestrial paradigm, and points toward a simpler model of time-averaged death assemblage formation where higher abundances reflect (undersampled) past populations. Contrary to long-held expectations, variation in population turnover among species is not a major source of taphonomic bias in time-averaged death assemblages among bivalves and perhaps among other marine groups: bias must arise largely from other factors.

Species selection and driven mechanisms jointly generate a large-scale morphological trend in monobathrid crinoids

Abstract: All evolution attributable to natural selection, at any level, is due to a causal covariance between fitness and phenotype. Over macroevolutionary time scales, species selection is one of many possible mechanisms for generating large-scale morphological trends. For species selection to sort morphology, a correlation between morphology and taxonomic diversification rate must be present. Other trend mechanisms (driven mechanisms, e.g., a bias in the direction of speciation) produce a systematic change in the mean phenotype over time. All mechanisms can co-occur. Here I demonstrate (1) an inverse correlation between diversification rate and calyx complexity that demonstrates the effect of species selection on morphology. Genera with simple calyces tend to increase in diversity, whereas genera with complex calyces have a net decrease in diversity; and (2) the presence of a driven trend mechanism in monobathrid crinoids where descendant genera tend to be simpler than their ancestors. The separate effects of these two classes of trend mechanisms can be combined by using the Price's Theorem, which partitions the contribution to the overall change in calyx complexity over time accurately among selection and driven mechanisms. Price's Theorem provides significant conceptual and methodological clarification of the contribution of multiple and interacting hierarchical mechanisms in generating large-scale trends.

Notes on Arboricultural and Agricultural Practices in Ancient Iran based on New Pollen Evidence

Abstract: De nouvelles donnees polliniques provenant de deux sites dans les montagnes du Zagros (lac Maharlou, Sud-Ouest l’Iran) et de Sahand (lac Almalou, Nord-Ouest de l’Iran), apportent des indications sur l’emergence de l’arboriculture dans le Sud-Ouest iranien depuis les IIIe et IIe millenaires av. J.-C., et sur les pratiques agricoles dans les hautes terres du Nord-Ouest iranien du Ve au IIIe siecles av. J.-C. Juglans cf. regia fut probablement cultive des 2500 av. J.-C., puis plus extensivement a partir de 1200 av. J.-C. environ. Platanus cf. orientalis fut probablement cultive precocement, vers 1900 av. J.-C., au cours de la periode elamite moyenne (2800-550 av. J.-C.), et il devint plus largement cultive a partir de ca. 1200 av. J.-C, parallelement a la culture de Juglans. Ces innovations agricoles ont pu resulter du developpement des grandes civilisations urbaines dans le Sud-Ouest de l’Iran. L’emergence de l’Empire perse semble etre associee a une revolution agraire a grande echelle sur le plateau iranien. Sous l’Empire achemenide (550-330 av. J.-C), l’arboriculture s’est developpee dans la region du lac Maharlou alors que l’agriculture atteignait les hautes terres autour du lac Almalou, probablement en raison d’une plus grande stabilite socio-economique et du developpement des techniques d’irrigation. Les periodes parthe et sassanide (224-642 de notre ere) ont egalement vu le developpement de l’arboriculture dans la region du lac Maharlou, mais dans une moindre mesure que lors de la periode achemenide. Dans la region du lac Almalou, la periode parthe ne temoigne pas de pratiques agricoles spectaculaires, en raison peut-etre de l’instabilite politique et socio-economique consecutive aux conflits avec l’Empire romain en Armenie. Sous l’Empire sassanide, la stabilite socio-economique et le renouveau des techniques agricoles ont pu toutefois favoriser le developpement a grande echelle de l’arboriculture dans le Nord-Ouest de l’Iran. La periode qui suit l’arrivee de l’Islam est caracterisee par des activites agricoles moins importantes sur le plateau iranien, tres probablement en raison de l’instabilite politique liee aux invasions arabes, turques et mongoles. Le dernier fait agricole significatif enregistre au lac Almalou est l’apparition de Ricinus communis, cultive des le debut de l’Empire safavide (1501-1722 de notre ere).

Houses, Households, and Changing Society in the Late Neolithic and Chalcolithic of the Southern Levant

Abstract: Dans le Sud du Levant, les constructions architecturales temoignent de la complexite des paysages sociaux au Neolithique recent et au Chalcolithique. Plutot qu’un processus evolutif, leurs caracteristiques suggerent une variabilite et une flexibilite considerables tout au long de ces periodes. A differentes epoques et selon les lieux, des bâtiments individuels ont ete isoles du reste de la communaute, dans une cour muree qui servait egalement a proteger leurs provisions et leur betail. Ailleurs, nous trouvons des zones d’habitations apparemment quasi-independantes, partageant une cour enclose par les maisons elles-memes qui renfermaient une grande part de l’espace dedie au stockage, alors que d’autres secteurs d’activite etaient partages. Partout, nous observons egalement des maisons isolees qui ne semblent associees ni a la cour ni a la zone d’habitation. En outre, parmi les maisons coexistantes, nous distinguons d’importantes variations dans leur taille, leur mode de construction, et leur capacite de stockage, suggerant un certain degre de differentiation economique, peut-etre une difference de rang, et probablement une forme de concurrence entre les habitations. Certaines caracteristiques suggerent que les chefs de maisons aient fait des choix strategiques en ce qui concerne la taille de leur demeure, leurs alliances avec d’autres maisonnees, l’accroissement des troupeaux ou l’accumulation des stocks de grains, qu’ils affichent de facon concurrentielle afin de promouvoir l’avantage politique et economique de leur maison. Ce modele pourrait correspondre au concept de C. Levi-Strauss des «societes a maison » .