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The body armor of ankylosaurians is a unique morphological feature among dinosaurs. While ankylosaurian body armor has been studied for decades, paleohistological analyses have only started to uncover the details of its function. Yet there has been an overall bias toward sampling ankylosaurian remains from the Northern Hemisphere, with limited quantitative studies on the morphological and functional evolution of the osteoderms composing their body armor. Here, we describe new ankylosaurian materials recovered from the Late Cretaceous of Antarctica that, in combination with data compiled from the literature, reveal new insights into the evolution of the ankylosaurian body armor. Based on histological microstructure and phylogenetic results, the new Antarctic material can be assigned to Nodosauridae. This group shares the absence/poor development of their osteodermal basal cortex and highly ordered sets of orthogonal structural fibers in the superficial cortex. Our morphospace analyses indicate that large morphological diversity is observed among both nodosaurids and ankylosaurids, but osteoderms became more functionally specialized in late-diverging nodosaurids. Besides acting as effective protection against predation, osteoderms also exhibit highly ordered structural fibers in nodosaurids, enabling a decrease in cortical bone thickness (as in titanosaurs), which could have been co-opted for secondary functions, such as calcium remobilization for physiological balance. The latter may have played a key role in nodosaurid colonization of high-latitude environments, such as Antarctica and the Arctic Circle.
The spatial distribution of in situ sessile organisms, including those from the fossil record, provides information about life histories, such as possible dispersal and/or settlement mechanisms, and how taxa interact with one another and their local environments. At Nilpena Ediacara National Park (NENP), South Australia, the exquisite preservation and excavation of 33 fossiliferous bedding planes from the Ediacara Member of the Rawnsley Quartzite reveals in situ communities of the Ediacara Biota. Here, the spatial distributions of three relatively common taxa, Tribrachidium, Rugoconites, and Obamus, occurring on excavated surfaces were analyzed using spatial point pattern analysis. Tribrachidium have a variable spatial distribution, implying that settlement or post-settlement conditions/preferences had an effect on populations. Rugoconites display aggregation, possibly related to their reproductive methods in combination with settlement location availability at the time of dispersal and/or settlement. Additionally, post-settlement environmental controls could have affected Rugoconites on other surfaces, resulting in lower populations and densities. Both Tribrachidium and Rugoconites also commonly occur as individuals or in low numbers on a number of beds, thus constraining possible reproductive strategies and environmental/substrate preferences. The distribution of Obamus is consistent with selective settlement, aggregating near conspecifics and on substrates of mature microbial mat. This dispersal process is the first example of substrate-selective dispersal among the Ediacara Biota, thus making Obamus similar to numerous modern sessile invertebrates with similar dispersal and settlement strategies.
La distribución espacial de los organismos sésiles in situ, incluyendo los del registro fósil, brinda información sobre las historias de vida, tal como los posibles mecanismos de dispersión y/o asentamiento, y sobre cómo los taxones interactúan entre sí y entre sus entornos locales. En el Parque Nacional Nilpena Ediacara (NENP), Australia Meridional, la excelente preservación y excavación de 33 planos de lecho fosilífero del Miembro Ediacara de la Cuarcita Rawnsley revela comunidades in situ de la biota ediacárica. En este estudio analizamos las distribuciones espaciales de tres taxones relativamente comunes, Tribrachidium, Rugoconites y Obamus, que se encuentran en superficies excavadas mediante el análisis de patrones de puntos espaciales. Tribrachidium tiene una distribución espacial variable, lo que implica que las condiciones/preferencias durante o después del asentamiento tuvieron un efecto en las poblaciones. Rugoconites muestran agregación, posiblemente relacionado con sus métodos reproductivos en combinación con la disponibilidad de lugares de asentamiento en el momento de la dispersión y/o asentamiento. Además, los controles ambientales posteriormente al asentamiento podrían estar afectando a Rugoconites en otras superficies, lo que resultaría en poblaciones y densidades más bajas. Tanto Tribrachidium como Rugoconites ocurren como individuos en varios lechos, restringiendo las posibles estrategias reproductivas y las preferencias ambientales/de sustrato. La distribución de Obamus es consistente con un asentamiento selectivo, agregando cerca de sus congéneres y sobre sustratos de tapete microbiano maduro. Este proceso de dispersión es el primer ejemplo de dispersión selectiva de sustrato entre la biota ediacárica, lo que hace que Obamus sea similar a numerosos invertebrados sésiles modernos con estrategias similares de dispersión y asentamiento.
Body size is an important trait with implications for energy use and ecology as well as generation time and evolutionary rates. Turritelline gastropods are widely distributed through geologic time and space, making them an excellent group for evaluating macroevolutionary patterns. To evaluate the pattern of body-size change in turritelline gastropods, we compiled a dataset of shell lengths of 316 species of turritelline gastropods spanning the Jurassic to Recent. Type specimens were almost always significantly larger than specimen distributions from the same species. We found that turritelline gastropod size was inversely correlated with latitude, a trend likely driven by the Neogene–Recent diversification of small-bodied Southern Hemisphere taxa. A time series model was applied to distinguish among three possible macroevolutionary patterns: unbiased random walk (no directional trend), biased random walk (directional trend), and stasis (no net change). We determined that turritelline gastropods have experienced stasis in body size throughout their evolutionary history, adding to the growing literature documenting directionless body-size trends in marine invertebrate clades. Stasis of geographically widespread clades may be the result of ecological variability across the environmental range occupied by the group or differential diversification into opposing environments. Turritelline life-history patterns, especially their reproductive strategy that combines a short life span and decline in growth rate around 1 year of age to reallocate energy to reproduction, might circumvent selection for longevity and larger size, while further decrease in minimum size is likely limited by feeding efficiency and anti-predatory defense. The expectation that species or clades should continue to evolve to occupy larger size classes conflicts with the evolutionary advantages of small size, which in turritelline gastropods include high generational turnover and larger population sizes that yield opportunities for genetic variance.
Drilling predation is a common reason for mortality of benthic mollusks but did not become common until the late Mesozoic. The scarcity of drill holes in the early Mesozoic fossil record limits our understanding of the evolution of drilling behavior and its role on shaping early Mesozoic marine communities. Here, we use drilling traces on several bivalve taxa from the Lower Jurassic (Pliensbachian) marine soft-bottom deposits in northern Germany to explore behavioral patterns of the predator (e.g., site selectivity, change in site-selective behavior with age). Although none of the known drilling gastropod groups existed in the Pliensbachian, including the studied localities, the drill-hole morphology suggests that the predator was probably a gastropod. The ecology and identity of the target prey changes from a diverse array of epifaunal to infaunal taxa in older deposits to focus on a single, large, deep infaunal taxon, Gresslya intermedia, in younger deposits, suggesting a potential trend in prey selectivity over time. Spatial point pattern analysis of traces (SPPAT) reveals an aggregated pattern of drill holes on Gresslya, suggesting strong selectivity in drill-hole location. Drilling on a single large infaunal taxon and site selectivity are common patterns also inferred previously from the drilled deep infaunal Eothyasira from the Pliensbachian of southern Germany. In addition to the scarcity of predators, the highly specialized behavior of the early drilling predators, including strong prey selectivity in terms of prey identity and life habit, can partly explain the rarity of the early Mesozoic drill holes.
Conodont elements have high rates of morphological evolution, but the drivers of this disparity are debated. Positive allometric relationships between dimensions of food-processing surfaces and entire P1 elements have been used to argue that these elements performed mechanical digestion. If involved in food processing, the surface of the element should grow at a rate proportional to the increase in energy requirements of the animal. This inference of function relies on the assumption that the energy requirements of the animal grew faster (≅ mass0.75) than the tooth area (≅ mass0.67). We reevaluate this assumption based on metabolic rates across animals and calculate the allometry in platform-bearing P1 elements of Late Triassic co-occurring taxa, Metapolygnathus communisti and Epigondolella rigoi, using 3D models of ontogenetic series. Positive allometry is found in platform and element dimensions in both species, supporting a grasping-tooth hypothesis, based on the assumption that metabolic rate in conodonts scaled with body mass similarly to that in fish and ectotherms. We also calculate the curvature of the P1 platform surface using the Dirichlet normal energy (DNE) as a proxy for diet. DNE values increase with body mass, supporting the assumption that conodont metabolic rates increased faster than mass0.67. We finally find that adults in both taxa differ in their food bases, which supports trophic diversification as an important driver of the remarkable disparity of conodont elements.
The hyoid apparatus is essential for underwater feeding in marine tetrapods, but it is unclear whether this complex has evolved as convergently as other traits, such as dentition or locomotion. Here we compare the ossified hyoid elements in ophthalmosaurid ichthyosaurs and odontocete cetaceans, two groups with an overall similar body shape, to understand whether the hyoid elements show any signs of convergence in the context of feeding. We examined three types of data (size, morphology, and internal bone microstructure) in ophthalmosaurid and odontocete taxa in which these elements are preserved. Our data show that ichthyosaurs never experienced a shift in feeding mode, which might indicate that their hyoid apparatus never adapted to suction feeding. Also, the internal microstructure of the two animal groups differs; where the odontocetes have an overall less compact structure, ophthalmosaurid ichthyosaurs have cancellous inner cones in an outer, more compact sheath. These differences are likely explained as biomechanical adaptations to different feeding modes. Thus, the hyoid changed less and acted more as a constraint for feeding innovation in ichthyosaurs compared with cetaceans, and through a much longer time span (more than 150 Myr).
Foramina of bones are beginning to yield more information about metabolic rates and activity levels of living and extinct species. This study investigates the relationship between estimated blood flow rate to the femur and body mass among cursorial birds extending back to the Late Cretaceous. Data from fossil foramina are compared with those of extant species, revealing similar scaling relationships for all cursorial birds and supporting crown bird–like terrestrial locomotor activity. Because the perfusion rate in long bones of birds is related to the metabolic cost of microfracture repair due to stresses applied during locomotion, as it is in mammals, this study estimates absolute blood flow rates from sizes of nutrient foramina located on the femur shafts. After differences in body mass and locomotor behaviors are accounted for, femoral bone blood flow rates in extinct species are similar to those of extant cursorial birds. Femoral robustness is generally greater in aquatic flightless birds than in terrestrial flightless and ground-dwelling flighted birds, suggesting that the morphology is shaped by life-history demands. Femoral robustness also increases in larger cursorial bird taxa, probably associated with their weight redistribution following evolutionary loss of the tail, which purportedly constrains femur length, aligns it more horizontally, and necessitates increased robustness in larger species.
Melanin pigments are central to colors and patterns in modern vertebrate integuments, which inform upon ecological and behavioral strategies like crypsis, aposematism, and sociosexual selection. Over the last decade, melanin has emerged as a valuable tool for predicting color in exceptionally preserved fossil feathers and subsequent testing of paleobiological hypotheses in long-extinct dinosaurs and birds. Yet much remains to be learned about melanin stability, diagenetic alterations to melanin chemistry, and their implications for “paleocolor reconstruction.” Pressure–temperature maturation experiments with modern feathers offer a way to examine these topics but have mostly been conducted in closed-system capsules or open-system aluminum foil. Both methods have operational limitations and do not consider the filtering effect of porous sediment matrices on thermally labile chemical groups versus stable ones during natural fossilization. We use sediment-encased maturation to resolve this issue and demonstrate replication of organic preservation of melanin highly comparable to compression fossils. Our experiments, coupled with time-of-flight secondary ion mass spectrometry, show predictable volatilization of N/S-bearing molecules and increased melanin cross-linking with elevated temperatures. We also suggest that eumelanin is more stable compared with pheomelanin at higher temperatures, explaining why eumelanic colors (black, dark brown, iridescent) are better preserved in fossils than pheomelanic ones (reddish brown). Furthermore, we propose that proteins preferentially undergo hydrolysis more so than forming N-heterocycles in selectively open systems analogous to natural matrices. Thus, we conclude that melanin pigments and not diagenetically altered protein remnants are the key players in promoting fossilization of soft tissues like feathers.
Paleontologists have long stressed the need to know how sampling the fossil record might influence our knowledge of the evolution of life. Here, we combine fossil occurrences of North American marine invertebrates from the Paleobiology Database with lithologic data from Macrostrat to identify sampling patterns in carbonate and siliciclastic rocks. We aim to quantify temporal trends in sampling effort within and between lithologies, focusing on the proportion of total available volume that has been sampled (sampled fossiliferous proportion, here called κ). Results indicate that the sampled fossiliferous proportion was stable during the Paleozoic, and variable during the post-Paleozoic, but showed no systematic increase through time. Fossiliferous carbonate rocks are proportionally more sampled than siliciclastic rocks, with intervals where the carbonate κ is double the siliciclastic κ. Among possible explanations for the apparent oversampling of fossiliferous carbonate rocks, analyses suggest that barren units, taphonomic dissolution, or data entry errors cannot completely explain sampling patterns. Our results suggest that one of the important drivers might be that paleontologists publish taxonomic descriptions from carbonate rocks more frequently. The higher diversity in carbonate rocks might account for an ease in the description of unknown species and therefore a higher rate of published fossils. Finally, a strong effect in favor of carbonate rocks might distort our perception of diversity through time, even under commonly used standardization methods. Our results also confirm that previous descriptions of an increase in the proportion of sampled fossiliferous rocks over time were driven by the sampling of the nonmarine fossil record.
The nature of phenotypic evolution within lineages is central to many unresolved questions in paleontology and evolutionary biology. Analyses of evolutionary time series of ancestor–descendant populations in the fossil record are likely to make important contributions to many of these debates. However, the limited number of models that have been applied to these types of data may restrict our ability to interpret phenotypic evolution in the fossil record. Using uni- and multivariate models of trait evolution that make different assumptions regarding the dynamics of the adaptive landscape, I evaluate contrasting hypotheses to explain evolution of size in the radiolarian Eucyrtidium calvertense and armor in the stickleback Gasterosteus doryssus. Body-size evolution in E. calvertense is best explained by a model in which the lineage evolves as a consequence of a shift in the adaptive landscape that coincides with the initiation of neosympatry with its sister lineage. Multivariate evolution of armor traits in a stickleback lineage (G. doryssus) shows evidence of adaptation toward independent optima on the adaptive landscape at the same time as traits change in a correlated fashion. The fitted models are available in the R package evoTS, which builds on the paleoTS framework.
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