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Encrusting bryozoans provide one of the few systems in the fossil record in which ecological competition can be observed directly at local scales. The macroevolutionary history of diversity of cyclostome and cheilostome bryozoans is consistent with a coupled-logistic model of clade displacement predicated on species within clades interacting competitively. The model matches observed diversity history if the model is perturbed by a mass extinction with a position and magnitude analogous to the Cretaceous/Tertiary boundary event. Although it is difficult to measure all parameters in the model from fossil data, critical factors are intrinsic rates of extinction, which can be measured. Cyclostomes maintained a rather low rate of extinction, and the model solutions predict that they would lose diversity only slowly as competitively superior species of cheilostomes diversified into their environment. Thus, the microecological record of preserved competitive interactions between cyclostome and cheilostome bryozoans and the macroevolutionary record of global diversity are consistent in regard to competition as a significant influence on diversity histories of post-Paleozoic bryozoans.
Members of the neogastropod muricid subfamily Rapaninae are abundant, shallow-water predators whose phylogeny was previously investigated by Kool (1993b), who used mainly anatomical characters. In order to deepen understanding of the evolution of this important clade and to incorporate functional, ecological, and fossil evidence, we performed a phylogenetic analysis based on 34 shell characters in 45 genus-level taxa, including five muricid outgroups. Cladograms based on shell characters alone differed from those founded on anatomical features, and these analyses differed from the phylogenetic reconstruction combining all available morphological evidence. The preferred cladogram incorporates all evidence and reveals a “Thais group” and an “Ergalatax clade” that both emerge from the derived portion of a more primitive, paraphyletic group of other rapanines. The Ocenebrinae, the other four outgroup taxa, and three ergalataxine taxa all lie outside the rapanine clade that includes the remaining ergalataxines as a derived subclade.
We used the phylogenetic results to probe aspects of the ecological history of the Rapaninae. Our data imply that antipredatory shell defenses (elongated aperture, denticles on the inner side of the outer lip, and robust external spines and tubercles) evolved multiple times, mainly in post–early Miocene clades in the Indo–West Pacific region. These results support earlier nonphylogenetic inferences.
We compared known prey types and methods of predation of living rapanines with their distribution on our phylogenetic tree. The plesiomorphic mode of feeding in the Rapaninae is drilling of hard-shelled prey. Feeding by other means and on such soft-bodied prey as sipunculan and polychaete worms evolved several times independently among post–early Miocene rapanines in the Indo–West Pacific. Methods of predation on hard-shelled prey that involve edge-drilling or attack by way of the aperture also evolved independently several times, but did so throughout the geographical range of the subfamily.
Specialization for life on the upper shore occurred in at least eight lineages, all but two of which are confined to the Indo–West Pacific. Ecological diversification of the Rapaninae was therefore most common in the tropical Indo–West Pacific during and after early Miocene time. This diversification occurred in a setting of already high biological diversity and intense competition and predation.
The bulk of Neoproterozoic trace fossils can be interpreted as horizontal creeping trails produced by minute vermiform organisms moving on or just beneath the seafloor or under algal mats. We have investigated the formation of trails by living cnidarians and platyhelminths that creep by cilia on mucus ribbons. These relatively simple metazoans produce trails that are similar in size and morphology to some Neoproterozoic traces, owing to the entrainment of sediment within their mucus trails. Thus a mucociliary locomotory system provides sufficient means to form some types of Neoproterozoic traces. It follows that the body architectures of the Neoproterozoic trace-makers may have been quite simple, though complex bodyplans are, of course, not ruled out. Thus, the use of Neoproterozoic trace fossils to constrain the time of origin of bilaterians or of any crown-group bilaterian taxon remains questionable.
Temporal asymmetries in clade histories have often been studied in lower Paleozoic radiations. Post-Paleozoic patterns, however, are less well understood. In this paper, disparity and diversity changes in Mesozoic heart urchins were analyzed at the ordinal level, with contrasts among the sister groups Holasteroida and Spatangoida, their paraphyletic stem group Disasteroida, and the more inclusive clade, the superorder Atelostomata. A 38-dimensional landmark-based morphospace representing test architecture was used to describe morphological evolution in terms of total variance and total range. Discordances between disparity and diversity were evident and were expressed both as deceleration in morphological diversification in all groups and as disproportionately higher disparity early in the histories of the Atelostomata, Holasteroida, and Spatangoida. The finding that the early atelostomate disparity peak coincides with the origin of the orders Holasteroida and Spatangoida lends support to the perception of orders as semi-independent entities in the biological hierarchy and as meaningful proxies for morphological distinctness.
A comparison of holasteroid and spatangoid responses to the end-Cretaceous mass extinction revealed morphological selectivity. Paleocene spatangoid survivors showed no change in disparity relative to the Campanian-Maastrichtian sample, suggesting nonselectivity. Holasteroids suffered a pronounced loss in disparity (despite a rather high Late Cretaceous level of disparity), indicating morphological selectivity of extinction.
Partitioning of disparity into plastral and nonplastral components, reflecting different degrees of developmental entrenchment and functionality, suggests that the origin of holasteroids and spatangoids is more consistent with an exploration of the developmental flexibility of nonplastral constructions than with uniform ecospace occupation. Within groups, several patterns were also most consistent with intrinsic controls. For plastral landmarks, there is an apparent increase in developmental modularity and decrease in developmental constraint from disasteroids to holasteroids and spatangoids. For nonplastral landmarks, no substantial change in disparity was observed from disasteroids to holasteroids and spatangoids, suggesting the maintenance of a developmental constraint despite the passage of time and ecological differentiation. More generally, this study suggests that certain topologies of disparity and evolutionary mechanisms potentially characteristic of the lower Paleozoic radiations of higher taxa (e.g., developmental flexibility) need not be confined to any given time period or hierarchical level.
Contrary to the geological stereotype of pure-carbonate reef platforms, approximately 50% of shallow shelf area in the Tropics is accumulating siliciclastic and mixed siliciclastic-carbonate sediments. Taphonomic characterization of these settings is thus essential for assessing variation among major facies types within the Tropics, as well as for eventual comparison with higher-latitude settings. Our grab samples and dredge samples of bivalve death assemblages from nine stations in five subtidal habitats in a large marine embayment of Caribbean Panama (Bocas del Toro) provide the first actualistic information on the taphonomic condition of shells in Recent tropical siliciclastic sediments. Focusing on unambiguous damage to bivalve shell interiors, we found that the quality of shell preservation in fine-grained siliciclastics is superb: commonly <10% of specimens are affected by encrustation, boring, edge-rounding, and fine-scale surface alteration via dissolution, microbioerosion, and maceration. Pure-carbonate and mixed siliciclastic-carbonate environments containing hard substrata (patch reefs, Halimeda gravelly sand, mud among patch reefs) contain higher numbers of more severely damaged shells (generally >25%) and also higher diversities of fossilizable encrusters and borers. Disarticulation and fragmentation are pervasive across all environments and are probably related to predation rather than to postmortem processes. As in other shallow subtidal study areas, the taxonomic compositions of death assemblages have not been homogenized by postmortem transport but show high spatial fidelity to the distribution of living species. Assemblages from the five sedimentary environments have distinct taphonomic signatures, but the strongest differences are between the two fine-grained, exclusively soft-sediment siliciclastic environments on the one hand and the three environments containing hard substrata on the other. Experimental tests for rates and agents of damage, still in progress, indicate that the most critical environmental variables are exhumation cycles and burial rate. Bivalve death assemblages from Bocas del Toro demonstrate that damage levels in tropical fine-grained siliciclastic environments are much lower than in closely associated reefs and algal sands, and suggest a less filtered record of biological information.
Bivalve death assemblages from subtidal environments within the tropical Bocas del Toro embayment of Caribbean Panama permit a test of the extent to which levels of damage are determined by the intrinsic nature of shell supply (proportion of epifaunal species, thick shells, calcitic shells, low-organic microstructures), as opposed to the extrinsic postmortem environment that shells experience. Only damage to interior surfaces of shells was used, to ensure that damage was unambiguously postmortem in origin. We find that facies-level differences in patterns of damage (the rank order importance of postmortem encrustation, boring, edge-rounding, fine-scale surface degradation) are overwhelmingly controlled by environmental conditions: in each environment, all subsets of the death assemblage present the same damage profile. The composition of shell supply affects only the intensity of the taphonomic signature (i.e., percentage of shells affected), and only in environments containing hard substrata (patch reefs, Halimeda gravelly sand, mud among patch reefs). In these environments, epifauna, whether aragonitic or calcitic and whether thin or thick, exhibit significantly higher damage than co-occurring infauna, probably due to the initial period of seafloor exposure they typically experience after death. Thick shells (>0.5 mm), regardless of life habit or mineralogy, are damaged more frequently than thin shells, probably because of selective colonization by fouling organisms. Calcitic shells show no consistently greater frequency of damage than aragonitic shells, and high-organic microstructures yield mixed patterns. Taphofacies surveys in such depositional systems could thus be confidently based on any subset of the fauna, including diagenetically residual assemblages of calcitic shells and thick-shelled molds. Further tests are needed to determine whether the higher levels of damage observed on some subsets of shells are a consequence of greater time-averaging (thus lower temporal resolution), greater exposure time, preferential attack (potential bias in relative abundance), or some combination of these. Paleobiologically, however, the implication is that ecological subsets of bivalve assemblages are not isotaphonomic, either in tangible damage or in probable bias, within hard-substrate environments, although they may be within soft-sediment environments. In actualistic studies, targeting broad classes of taxa for comparison across environments maximizes our ability to extrapolate taphonomic guidelines into the fossil record, where life habits, skeletal types, and shallow subtidal habitats have dramatically different patterns of abundance and deployment.
A geometric analysis of fenestrate bryozoan lophophore shape and arrangement is conducted by creating a theoretical morphospace of apertural positioning within the colonial meshwork. Working from the assumption that fenestrate bryozoans needed to form a continuous filtering surface with contact between adjacent lophophores, we show that within the morphospace three regions exist for optimum close-packing of lophophores with circular projections; all other close-packing configurations in the morphospace require the existence of noncircular lophophores.
Examination of the actual distribution of 251 fenestrate colonies within the morphospace reveals that the morphospace regions occupied by fenestellids and polyporids are displaced and have little overlap, but that they are very similar in size and shape, and that the colonies scale similarly. With increasing size, fenestrate meshworks expand laterally faster than the branches widen and the proximodistal spacing of the apertures increases, apparently because the larger zooids require disproportionately more room for their lophophores.
Two of the optimum close-packing regions of the morphospace are occupied by fenestrates. The positioning of the fenestellid region within the morphospace suggests that these biserial bryozoans followed a proximodistal-row placement of the lophophores, and that the lophophores were generally equitentacular, with circular projections. The positioning of the polyporid region within the morphospace suggests that these polyserial bryozoans followed a diagonal-row placement of the lophophores, and that the lophophores were heteromorphic, with medial lophophores on the branch being more equitentacular whereas the laterally placed lophophores were obliquely truncate. The third optimum close-packing region in the morphospace, corresponding to a hypothetical lateral-row placement of the lophophores within the colony, is unoccupied. We suggest that hypothetical fenestrate morphologies in the vacant region of morphospace have branches that would be too narrow to support normally shaped zooids, and that the lateral-row placement of the lophophores would have required the branches of the colony to have been perfectly aligned throughout growth.
Atmospheric carbon dioxide is the raw material for the biosphere. Therefore, changes in the carbon isotopic composition of the atmosphere will influence the terrestrial δ13C signals we interpret. However, reconstructing the atmospheric δ13C value in the geologic past has proven challenging. Land plants sample the isotopic composition of CO2 during photosynthesis. We use a model of carbon isotopic fractionation during C3 photosynthesis, in combination with a meta–data set (519 measurements from 176 species), to show that the δ13C value of atmospheric CO2 can be reconstructed from the isotopic composition of plant tissue. Over a range of pCO2 (198–1300 ppmv), the δ13C value of plant tissue does not vary systematically with atmospheric carbon dioxide concentration. However, environmental factors, such as water stress, can influence the δ13C value of leaf tissue. These factors explained a relatively small portion of variation in the δ13C value of plant tissue in our data set and emerged strongly only when the carbon isotopic composition of the atmosphere was held constant. Members of the Poaceae differed in average δ13C value, but we observed no other differences correlated with plant life form (herbs, trees, shrubs). In contrast, over 90% of the variation the carbon isotopic composition of plant tissue was explained by variation in the δ13C value of the atmosphere under which it was fixed. We use a subset of our data spanning a geologically reasonable range of atmospheric δ13C values (−6.4‰ to −9.6‰) and excluding C3 Poaceae to develop an equation to reconstruct the δ13C value of atmospheric CO2 based on plant values. Reconstructing the δ13C value of atmospheric CO2 in geologic time will facilitate chemostratigraphic correlation in terrestrial sediments, calibrate pCO2 reconstructions based on soil carbonates, and offer a window into the physiology of ancient plants.