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Fossil bone microanalyses reveal the ontogenetic histories of extinct tetrapods, but incomplete fossil records often result in small sample sets lacking statistical strength. In contrast, a histological sample of 50 tibiae of the hadrosaurid dinosaur Maiasaura peeblesorum allows predictions of annual growth and ecological interpretations based on more histologic data than any previous large sample study. Tibia length correlates well (R2 > 0.9) with diaphyseal circumference, cortical area, and bone wall thickness, thereby allowing longitudinal predictions of annual body size increases based on growth mark circumference measurements. With an avian level apposition rate of 86.4 µm/day, Maiasaura achieved over half of asymptotic tibia diaphyseal circumference within its first year. Mortality rate for the first year was 89.9% but a seven year period of peak performance followed, when survivorship (mean mortality rate = 12.7%) was highest. During the third year of life, Maiasaura attained 36% (x = 1260 kg) of asymptotic body mass, growth rate was decelerating (18.2 µm/day), cortical vascular orientation changed, and mortality rate briefly increased. These transitions may indicate onset of sexual maturity and corresponding reallocation of resources to reproduction. Skeletal maturity and senescence occurred after 8 years, at which point the mean mortality rate increased to 44.4%. Compared with Alligator, an extant relative, Maiasaura exhibits rapid cortical increase early in ontogeny, while Alligator cortical growth is much lower and protracted throughout ontogeny. Our life history synthesis of Maiasaura utilizes the largest histological sample size for any extinct tetrapod species thus far, demonstrating how large sample microanalyses strengthen paleobiological interpretations.
Research into the relationship between leaf form and climate over the last century has revealed that, in many species, the sizes and shapes of leaf characters exhibit highly structured and predictable patterns of variation in response to the local climate. Several procedures have been developed that quantify covariation between the relative abundance of plant character states and the states of climate variables as a means of estimating paleoclimate parameters. One of the most widely used of these is the Climate Leaf Analysis Multivariate Program (CLAMP). The consistency, accuracy and reliability with which leaf characters can be identified and assigned to CLAMP character-state categories is critical to the accuracy of all CLAMP analyses. Here we report results of a series of performance tests for an image-based, fully automated at the point of use, leaf character scoring system that can be used to generate CLAMP leaf character state data for: leaf bases (acute, cordate and round), leaf apices (acute, attenuate), leaf shapes (ovate, elliptical and obovate), leaf lobing (unlobed, lobed), and leaf aspect ratios (length/width). This image-based system returned jackknifed identification accuracy ratios of between 87% and 100%. These results demonstrate that automated image-based identification systems have the potential to improve paleoenvironmental inferences via the provision of accurate, consistent and rapid CLAMP leaf-character identifications. More generally, our results provide strong support for the feasibility of using fully automated, image-based morphometric procedures to address the general problem of morphological character-state identification.
Arthropods are known to display a variable number of eye lenses and this number mostly increases during their development. In trilobites, most species possessing schizochroal eyes exhibit a notable intraspecific variation in the number of dorso ventral files of eye lenses that can be age related (i.e., growth) or not (i.e., living environment). Several previous studies have shown that some trilobite groups (e.g. phacopids) tend to have fewer lenses/files in representatives from the deeper habitats than those from shallower habitats. In this study, we analyzed the pattern of variation in the number of dorso ventral files of eye lenses in two Devonian phacopid trilobites from the Prague Basin of the Czech Republic. We quantified their intraspecific variability. To better understand the patterning, we compared more than 120 individuals. Data first reveal evidence of a bimodal distribution of lens/file number without intermediate forms among each of two studied populations of Prokops prokopi (Chlupáč, 1971) and throughout the ontogeny of Pedinopariops insequens (Chlupáč, 1977). Our results indicate that caution must be taken for taxonomical affiliation and biodiversity analyses of taxa in which the intraspecific variability is unclear. Additionally, we investigated possible relations of these bimodalities to the stratigraphical position of studied populations and to the paleoenvironment. In Prokops prokopi, a slightly different age of both populations, together with supposed differences in the local environments can be responsible for observed variability. In Pedinopariops insequens, stress conditions possibly related to the approaching onset of the Basal Choteč Event can be responsible for surprising intrapopulation variability. We speculate that the stress conditions could cause a bimodal selection and possibly also the change of ontogenetic trajectory within this species. Pedinopariops insequens was the only phacopid in the Prague Basin that crosses the Lower/Middle Devonian boundary and survived also the onset of Basal Choteč Event.
Paleobiologists must propose a priori hypotheses of homology when conducting a phylogenetic analysis of extinct taxa. The distributions of such “primary” homologies among species are fundamental to phylogeny reconstruction because they reflect a prior belief in what constitutes comparable organismal elements and are the principal determinants of the outcome of phylogenetic analysis. Problems arise when fossil morphology presents seemingly equivocal hypotheses of homology, herein referred to as antinomies. In groups where homology recognition has been elusive, such as echinoderms, these problems are commonly accompanied by the presence (and persistence) of poor descriptive terminology in taxonomic literature that confounds an understanding of characters and stymy phylogenetic research. This paper combines fossil morphology, phylogenetic systematics, and insights from evolutionary developmental biology to outline a research program in Phylogenetic Paleo-ontogeny. A “paleo” ontogenetic approach to character analysis provides a logical basis for homology recognition and discerning patterns of character evolution in a phylogenetic context. To illustrate the utility of the paleo-ontogenetic approach, I present a reassessment of historically contentious plate homologies for “pan-cladid” crinoids (Cladida, Flexibilia, Articulata). Developmental patterns in living crinoids were combined with the fossil record of pan-cladid morphologies to investigate primary posterior plate homologies. Results suggest the sequence of morphologic transitions unfolding during the ontogeny of extant crinoids are developmental relics of their Paleozoic precursors. Developmental genetic modules controlling posterior plate development in pan-cladid crinoids have likely experienced considerable constraint for over 250 million years and limited morphologic diversity in the complexity of calyx characters. Future phylogenetic analyses of pan-cladids are recommended to consider the presence of a single plate in the posterior region homologous with the radianal, rather than the anal X, as is commonly assumed.
The use of sclerobiosis as a tool for paleoenvironmental and paleoecological research is undermined by a lack of comparable methods for sclerobiont data collection and analysis. We present a new method for mapping sclerobiont distributions across any host, and offer an example of how the method may be used to interpret sclerobiont data in relation to host orientation. This approach can also be used to assess the suitability of beds and fossil material for paleoenvironmental reconstruction.
A sample of 150 encrusted dorsibiconvex atrypide brachiopods were selected from six beds in the Waterways Formation (latest Givetian — Early Frasnian; Alberta, Canada). The dorsal and ventral valves of each brachiopod were photographed. Sclerobiont taxa were mapped onto the photographs, and the maps were used to create stacked images with each of the 25 brachiopod specimens from each bed. Based on the life orientation of dorsibiconvex atrypides, three zones were designated on the host: the post mortem zone, (only available to sclerobionts after death and reorientation of the host); the shaded zone (brachial valve, excluding the post mortem zone); and the exposed zone (ventral valve).
Randomization simulation results indicate that all beds likely exhibit non random encrustation patterns, and corroborate the hypotheses that: (1) much of the encrustation occurred while the hosts were alive, and (2) these beds and fossils have experienced little physical reworking or transport and would be suitable for paleoenvironmental analysis. Mapping sclerobionts across hosts can serve as a unifying method to increase the recognition and use of sclerobiosis in paleontological studies.
Proterozoic strata host evidence of global “Snowball Earth” glaciations, large perturbations to the carbon cycle, proposed changes in the redox state of oceans, the diversification of microscopic eukaryotes, and the rise of metazoans. Over the past half century, the number of fossils described from Proterozoic rocks has increased exponentially. These discoveries have occurred alongside an increased understanding of the Proterozoic Earth system and the geological context of fossil occurrences, including improved age constraints. However, the evaluation of relationships between Proterozoic environmental change and fossil diversity has been hampered by several factors, particularly lithological and taphonomic biases. Here we compile and analyze the current record of eukaryotic fossils in Proterozoic strata to assess the effect of biases and better constrain diversity through time. Our results show that mean within assemblage diversity increases through the Proterozoic Eon due to an increase in high diversity assemblages, and that this trend is robust to various external factors including lithology and paleogeographic location. In addition, assemblage composition changes dramatically through time. Most notably, robust recalcitrant taxa appear in the early Neoproterozoic Era, only to disappear by the beginning of the Ediacaran Period. Within assemblage diversity is significantly lower in the Cryogenian Period than in the preceding and following intervals, but the short duration of the nonglacial interlude and unusual depositional conditions may present additional biases. In general, large scale patterns of diversity are robust while smaller scale patterns are difficult to discern through the lens of lithological, taphonomic, and geographic variability.
A new turnover rate metric is introduced that combines simplicity and precision. Like the related three-timer and gap-filler equations, it involves first identifying a cohort of taxa sampled in the time interval preceding the one of interest (call the intervals i0 and i1). Taxa sampled in i0 and i1 are two-timers (t2); those sampled in i0 and i2 but not i1 are part-timers (p); and taxa sampled only in either i1, i2, or i3 are newly notated here as either s1, s2, or s3. The gap-filler extinction proportion can be reformulated as (s1 - s3)/(t2 p). The method proposed here is to substitute s3 with the second-highest of the three counts when the expected ordering s1≤s2≤s3 is violated. In simulation, this new estimator yields values that are highly correlated with those produced by the gap-filler equation but more precise. In particular, it rarely produces highly negative values even when sample sizes are quite small. It is mildly upwards biased when sampling is extremely poor and turnover rates are extremely low, but it is otherwise highly accurate. Examples of Phanerozoic extinction rates for four major marine invertebrate groups are given to illustrate the method's improved precision. Based on the results, the procedure is recommended for general use.
Detailed quantitative data has previously been collected from plant megafossil assemblages from a Middle Jurassic (Aalenian) plant bed from Hasty Bank, North Yorkshire, UK. We conducted a similar analysis of palynological dispersed sporomorph (spore and pollen) assemblages collected from the same section using the same sampling regime: 67 sporomorph taxa were recorded from 50 samples taken at 10 cm intervals through the plant bed. Basic palynofacies analysis was also undertaken on each sample. Both dispersed sporomorph and plant megafossil assemblages display consistent changes in composition, diversity (richness), and abundance through time. However, the dispersed sporomorph and plant megafossil records provide conflicting evidence for the nature of parent vegetation. Specifically, conifers and ferns are underrepresented in plant megafossil assemblages, bryophytes and lycopsids are represented only in sporomorph assemblages, and sphenophytes, pteridosperms, Caytoniales, Cycadales, Ginkgoales and Bennettitales are comparatively underrepresented in sporomorph assemblages. Combined multivariate analysis (correspondence analysis and nonmetric multidimensional scaling) of sporomorph occurrence/abundance data demonstrates that temporal variation in sporomorph assemblages is the result of depositional change through the plant bed. The reproductive strategies of parent plants are considered to be a principal factor in shaping many of the major abundance and diversity irregularities between dispersed sporomorph and plant megafossil data sets that seemingly reflects different parent vegetation. Preferential occurrence/preservation of sporomorphs and equivalent parent plants is a consequence of a complex array of biological, ecological, geographical, taphonomic, and depositional factors that act inconsistently between and within fossil assemblages, which results in notable discrepancies between data sets.
The earliest Paleocene record of calcareous nannoplankton presents a unique opportunity to understand the evolutionary recovery of life from mass extinction. Nannoplankton were devastated at the Cretaceous/Paleogene boundary and their subsequent recovery can be studied in great detail because of their abundance in sediments, continuous stratigraphic occurrence, and near global distribution. Here we determine when and where new species of nannoplankton originated and how they dispersed following the Cretaceous/Paleogene mass extinction. Initially, we focus our efforts on North Pacific and South Atlantic deep sea sites with orbital age control to compare the precise timing and dynamics of the recovery between the locations. We then broaden our investigation to six sites from different basins and a variety of environments to study global patterns of the initial recovery. Our results show that many taxa in key Paleogene lineages originated in the North Pacific Ocean and that assemblages comprised primarily of new Paleogene taxa were not observed at other sites for several hundred thousand years. Survivors that were adapted to eutrophic post extinction conditions rapidly expanded in Southern Hemisphere sites where they dominated assemblages for most of the initial recovery. We therefore hypothesize that groups of survivors formed regionally incumbent assemblages in the Southern Hemisphere that limited diversification and dispersal of new Paleogene taxa. The end of survivor dominance correlates to the recovery of the biologic pump and subsequent decrease in surface ocean nutrient concentration 300–400 Kyr after the boundary. Only after survivors were removed did new Paleogene nannoplankton assemblages become abundant globally. Our results indicate that competition from regionally incumbent survivors was as an important control on the K/Pg recovery of nannoplankton.
Drillholes made by naticid and muricid gastropods are frequently used in evolutionary and ecological studies because they provide direct, preservable evidence of predation. The muricid Ecphora is common in many Neogene Atlantic Coastal Plain assemblages in the United States, but is frequently ignored in studies of naticid predation. We used a combination of Pliocene fossil, modern beach, and experimentally derived samples to evaluate the hypothesis that Ecphora was an important source of drillholes in infaunal bivalve prey shared with naticids. We focused on the large, thick-shelled venerid, Mercenaria, which is commonly drilled by naticids today. Laboratory experiments, modern beach samples, and the published literature confirm that naticids preferentially drill near the umbo (significant clumping of holes), show a significant correlation between prey size and predator size (estimated by outer borehole diameter), and prefer Mercenaria <50 mm antero-posterior width when other prey are present. Fossil samples containing Ecphora (with or without other large muricids) show no drillhole site stereotypy (no significant clumping, greater variability in placement), no significant predator: prey size correlation, drilled prey shells larger than the largest modern naticids could produce in an experimental setting, and drillholes larger in diameter than those estimated for the largest Pliocene naticids, thus supporting our hypothesis. Substantial overlap in the placement of holes drilled by naticids and muricids, however, made identifying predators from drillhole position problematic. The lack of overlapping ranges of prey shell thickness between fossil and other samples precluded the use of drill- hole morphology to establish predator identity (e.g., ratio of inner borehole diameter to outer borehole diameter, drillhole angle). Whereas the difficulty in determining predator identity from drillholes limits the types of analyses that can be reliably performed in mixed-predator assemblages, recognizing Ecphora as a prominent drilling predator creates the opportunity to investigate previously unrecognized questions.
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