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We know that the fossil record is incomplete. But how incomplete? Here we very coarsely estimate the completeness of the mammalian record in the Miocene, assuming that the duration of a mammalian species is about 1 Myr and the species diversity has stayed constant and is structurally comparable to the taxonomic diversity today. The overall completeness under these assumptions appears to be around 4%, but there are large differences across taxonomic groups. We find that the fossil record of proboscideans and perissodactyls as we know it for the Miocene must be close to complete, while we might know less than 15% of the species of artiodactyl or carnivore fossil species and only about 1% of primate species of the Miocene. The record of small mammals appears much less complete than that of large mammals.
Intraspecific aggression, or agonism, is a widespread intrasexual selective behavior important to understanding animal behavioral ecology and reproductive systems. Such behavior can be studied either by direct observation or inferred from wound/scar frequency in extant species but is difficult to document in extinct taxa, limiting understanding of its evolution. Among extant archosaurs, crocodylians display extensive intrasexual aggression, whereas birds show extreme visual/vocal intersexual display. The evolutionary origin of this behavioral divergence, and pattern in non-avian dinosaurs, is unknown. Here we document the morphology, frequency, and ontogeny of intraspecific facial bite lesions (324 lesions) in a large sample of tyrannosaurids (202 specimens, 528 elements) to infer patterns of intraspecific aggression in non-avian theropods. Facial scars are consistent in position and orientation across tyrannosaurid species, suggesting bites were inflicted due to repeated/postured behavior. Facial scars are absent in young tyrannosaurids, first appear in immature animals (∼50% adult skull length), are present in ∼60% of the adult-sized specimens, and show aggressor:victim size isometry. The ontogenetic distribution of bite scars suggests agonistic behavior is associated with the onset of sexual maturity, and scar presence in approximately half the specimens may relate to a sexual pattern. Considered in a phylogenetic context, intraspecific bite marks are consistent and widely distributed in fossil and extant crocodyliforms and non-maniraptoriform theropods, suggesting a potential plesiomorphic behavior in archosaurs. Their absence in maniraptoriform theropods, including birds, may reflect a transition from boney cranial ornamentation and crocodylian-like intrasexual aggression to avian-like intersexual display with the evolution of pennaceous feathers.
The Ordovician–Silurian (∼485–419 Ma) was a time of considerable evolutionary upheaval, encompassing both great evolutionary diversification and one of the first major mass extinctions. The Ordovician diversification coincided with global climatic cooling and paleocontinental collision, the ecological impacts of which were mediated by region-specific processes including substrate changes, biotic invasions, and tectonic movements. From the Sandbian–Katian (∼453 Ma) onward, an extensive carbonate shelf developed in the eastern Baltic paleobasin in response to a tectonic shift to tropical latitudes and an increase in the abundance of calcareous macroorganisms. We quantify the contributions of environmental differentiation and temporal turnover to regional diversity through the Ordovician and Silurian, using brachiopod occurrences from the more shallow-water facies belts of the eastern Baltic paleobasin, an epicontinental sea on the Baltica paleocontinent. The results are consistent with carbonate shelf development as a driver of Ordovician regional diversification, both by enhancing broadscale differentiation between shallow- and deep-marine environments and by generating heterogeneous carbonate environments that allowed increasing numbers of brachiopod genera to coexist. However, temporal turnover also contributed significantly to apparent regional diversity, particularly in the Middle–Late Ordovician.
The effects of overdispersion and zero inflation (e.g., poor model fits) can result in misinterpretation in studies using count data. These effects have not been evaluated in paleoecological studies of predation and are further complicated by preservational bias and time averaging. We develop a hierarchical Bayesian framework to account for uncertainty from overdispersion and zero inflation in estimates of specimen and predation trace counts. We demonstrate its application using published data on drilling predators and their prey in time-averaged death assemblages from the Great Barrier Reef, Australia.
Our results indicate that estimates of predation frequencies are underestimated when zero inflation is not considered, and this effect is likely compounded by removal of individuals and predation traces via preservational bias. Time averaging likely reduces zero inflation via accumulation of rare taxa and events; however, it increases the uncertainty in comparisons between assemblages by introducing variability in sampling effort. That is, there is an analytical cost with time-averaged count data, manifesting as broader confidence regions. Ecological inferences in paleoecology can be strengthened by accounting for the uncertainty inherent to paleoecological count data and the sampling processes by which they are generated.
Constraining patterns of growth using directly observable and quantifiable characteristics can reveal a wealth of information regarding the biology of the Ediacara biota—the oldest macroscopic, complex community-forming organisms in the fossil record. However, these rely on individuals captured at an instant in time at various growth stages, and so different interpretations can be derived from the same material. Here we leverage newly discovered and well-preserved Dickinsonia costataSprigg, 1947 from South Australia, combined with hundreds of previously described specimens, to test competing hypotheses for the location of module addition. We find considerable variation in the relationship between the total number of modules and body size that cannot be explained solely by expansion and contraction of individuals. Patterns derived assuming new modules differentiated at the anterior result in numerous examples in which the oldest module(s) must decrease in size with overall growth, potentially falsifying this hypothesis. Observed polarity as well as the consistent posterior location of defects and indentations support module formation at this end in D. costata. Regardless, changes in repeated units with growth share similarities with those regulated by morphogen gradients in metazoans today, suggesting that these genetic pathways were operating in Ediacaran animals.
The Rhynchonellida is a major group of brachiopods that survived the “big five” mass extinctions and flourished after the Permian/Triassic (P/Tr) crisis. However, phylogenetic and character evolution in the Rhynchonellida across the P/Tr transition is poorly understood. In view of the widespread homoplasy across this order, we employ a tip-dated Bayesian analysis to reconstruct phylogenetic relationships for late Permian–Triassic rhynchonellides. The same data were also analyzed using three other methods: undated Bayesian, equal-weighting, and implied-weighting parsimony. Compared with trees generated by other methods, those constructed by tip-dating best account for the homoplasy in this group and are closer to previous assumptions on the evolution of this order. Based on the analyses of multiple trees, the major increase in lineage richness occurred in the Early and early Middle Triassic. Also, richness in the Anisian almost reached the highest level seen in the Triassic. According to fossil records, a pronounced reduction in shell size and in the development of ornamentation occurred after the P/Tr extinction, which is largely due to the loss of large and highly sculptured genera and the diversification of small-sized and weakly ornamented genera. Ancestral-state estimation of shell size and development of ornamentation, coupled with comparisons of other characters, indicate that the Early–Middle Triassic mature “small-sized taxa” may have characters displayed by juveniles of their ancestors. This suggests that for these genera, paedomorphosis was possibly a strategy to survive and diversify in the harsh environment after the P/Tr extinction.
Studies in extant populations have shown that plasticity in developmental trajectories can contribute to the origin of novel traits and species divergence via the expression of previously cryptic variation in response to environmental change. Finding evidence for plasticity-led evolution in the fossil record remains challenging due to the poor preservation of developmental stages in many organisms. Planktic foraminifera are ideally suited for addressing this knowledge gap, because adult organisms in species in which development has been studied retain information about all the ontogenetic stages they have undergone. Here we map changes in the developmental trajectories of 68 specimens in the Globorotalia plesiotumida–tumida lineage of planktic foraminifera from the late Miocene until Recent using high-resolution computer tomography techniques. Our unique dataset shows that the transition from the ancestral G. plesiotumida to the descendant G. tumida is preceded by an increased variability in total cumulative volume—an important indicator of reproductive success in this taxon. We also find that the transition interval is marked by a distinct shift in developmental trajectory, which supports a rapid lineage division rather than gradual change. We suggest that high levels of plasticity—particularly in the early stages of development—have contributed to divergence in the ancestral morphology when subjected to a global cooling trend in the late Miocene. The large variation in developmental trajectories that we uncover within our samples emphasizes the need for high-throughput approaches in studies of ontogenetic change in the fossil record.
With 14 species recorded, the Miocene La Venta bat fauna is the most diverse bat paleocommunity in South America. It includes the oldest plant-visiting bat in the New World and some of the earliest representatives of the extant families Phyllostomidae, Thyropteridae, and Noctilionidae. La Venta's Notonycteris magdalenensis is an extinct member of the subfamily Phyllostominae, a group of modern Neotropical animalivorous bats, and is commonly included in studies of the evolution of Neotropical bats, but aspects of its biology remain unclear. In this study, we used multivariate dental topography analysis (DTA) to reconstruct the diet of N. magdalenensis by quantitatively comparing measures of molar complexity with those of 25 modern noctilionoid species representing all major dietary habits in bats. We found clear differences in molar complexity between dietary guilds, indicating that DTA is potentially an informative tool to study bat ecomorphology. Our results suggest N. magdalenensis was probably an omnivore or insectivore, rather than a carnivore like its modern relatives Chrotopterus auritus and Vampyrum spectrum. Also, we reconstructed the body mass of N. magdalenensis to be ∼95 g, larger than most insectivorous bats, but smaller than the largest carnivorous bat (V. spectrum). Our results confirm that N. magdalenensis was not a specialized carnivore. It remains to be demonstrated that the specialized carnivory ecological niche was occupied by the same lineage of phyllostomines from at least the middle Miocene. Combining our diet and body-mass reconstructions, we suggest that N. magdalenensis exhibits morphological pre-adaptations crucial for the evolution of specialized carnivory.
Paleobiological and paleoecological interpretations rely on constraining the temporal resolution of the fossil record. The taphonomic clock, that is, a correlation between the alteration of skeletal material and its age, is an approach for quantifying time-averaging scales. We test the taphonomic clock hypothesis for marine demersal and pelagic fish otoliths from a 10–40 m depth transect on the Mediterranean siliciclastic Israeli shelf by radiocarbon dating and taphonomic scoring. Otolith ages span the last ∼8000 yr, with considerable variation in median and range along the transect. Severely altered otoliths, contrary to pristine otoliths, are likely to be older than 1000 yr. For pelagic fish otoliths, at 30 m depth, taphonomic degradation correlates positively with postmortem age. In contrast, no correlation occurs for demersal fishes at 10 and 30 m depth, mostly because of the paucity of very young pristine (<150 yr) otoliths, possibly due to a drop in production over the last few centuries. Contrary to molluscan and brachiopod shells, young otoliths at these depths are little affected and do not show a broad spectrum of taphonomic damage, because those that derive from predation are excreted in calcium- and phosphate-rich feces forming an insoluble crystallic matrix that increases their preservation potential. At 40 m depth, all dated otoliths are very young but rather damaged because of locally chemically aggressive sediments, thus showing no correlation between taphonomic grade and postmortem age. Our results show that local conditions and the target species population dynamics must be considered when testing the taphonomic clock hypothesis.
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