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As the climate changes and ecosystems shift toward novel combinations of species, the methods and metrics of conservation science are becoming less species-centric. To meet this growing need, marine conservation paleobiologists stand to benefit from the addition of new, taxon-free benthic indices to the live–dead analysis tool kit. These indices, which were developed to provide actionable, policy-specific data, can be applied to the readily preservable component of benthic communities (e.g., mollusks) to assess the ecological quality status of the entire community. Because these indices are taxon-free, they remain applicable even as the climate changes and novel communities develop—making them a potentially valuable complement to traditionally applied approaches for live–dead analysis, which tend to focus on maintaining specific combinations of species under relatively stable environmental conditions. Integrating geohistorical data with these established indices has potential to increase the salience of the live–dead approach in the eyes of resource managers and other stakeholders.
Noel A. Heim, Saket H. Bakshi, Loc Buu, Stephanie Chen, Shannon Heh, Ashli Jain, Christopher Noll, Ameya Patkar, Noah Rizk, Sriram Sundararajan, Isabella Villante, Matthew L. Knope, Jonathan L. Payne
The typical marine animal has increased in biovolume by more than two orders of magnitude since the beginning of the Cambrian, but the causes of this trend remain unknown. We test the hypothesis that the efficiency of intra-organism oxygen delivery is a major constraint on body-size evolution in marine animals. To test this hypothesis, we compiled a dataset comprising 13,723 marine animal genera spanning the Phanerozoic. We coded each genus according to its respiratory medium, circulatory anatomy, and feeding mode. In extant genera, we find that respiratory medium and circulatory anatomy explain more of the difference in size than feeding modes. Likewise, we find that most of the Phanerozoic increase in mean biovolume is accounted for by size increase in taxa that accomplish oxygen delivery through closed circulatory systems. During the Cambrian, water-breathing animals with closed circulatory systems were smaller, on average, than contemporaries with open circulatory systems. However, genera with closed circulatory systems superseded in size genera with open circulatory systems by the Middle Ordovician, as part of their Phanerozoic-long trend of increasing size. In a regression analysis, respiratory and circulatory anatomy explain far more size variation in the living fauna than do feeding modes, even after accounting for taxonomic affinity at the class level. These findings suggest that ecological and environmental drivers of the Phanerozoic increase in the mean size of marine animals operated within strong, anatomically determined constraints.
Principal coordinates analysis (PCoA) is a statistical ordination technique commonly applied to morphology-based cladistic matrices to study macroevolutionary patterns, morphospace occupation, and disparity. However, PCoA-based morphospaces are dissociated from the original data; therefore, whether such morphospaces accurately reflect body-plan disparity or extrinsic factors, such as body size, remains uncertain. We collated nine character–taxon matrices of dinosaurs together with body-mass estimates for all taxa and tested for relationships between body size and both the principal axis of variation (i.e., PCo1) and the entire set of PCo scores. The possible effects of body size on macroevolutionary hypotheses derived from ordinated matrices were tested by reevaluating evidence for the accelerated accumulation of avian-type traits indicated by a strong directional shift in PCo1 scores in hypothetical ancestors of modern birds. Body mass significantly accounted for, on average, approximately 50% and 16% of the phylogenetically corrected variance in PCo1 and all PCo scores, respectively. Along the avian stem lineage, approximately 30% of the morphological variation is attributed to the reconstructed body masses of each ancestor. When the effects of body size are adjusted, the period of accelerated trait accumulation is replaced by a more gradual, additive process. Our results indicate that even at low proportions of variance, body size can noticeably affect macroevolutionary hypotheses generated from ordinated morphospaces. Future studies should thoroughly explore the nature of their character data in association with PCoA-based morphospaces and use a residual/covariate approach to account for potential correlations with body size.
A thorough understanding of how communities respond to extreme changes, such as biotic invasions, is essential to manage ecosystems today. Here we constructed fossil food webs to identify changes in Late Ordovician (Katian) shallow-marine paleocommunity structure and functioning before and after the Richmondian invasion, a well-documented ancient invasion. Food webs were compared using descriptive metrics and cascading extinction on graphs models. Richness at intermediate trophic levels was underrepresented when using only data from the Paleobiology Database relative to museum collections, resulting in a spurious decrease in modeled paleocommunity stability. Therefore, museum collections and field sampling may provide more reliable sources of data for the reconstruction of trophic organization in comparison to online data repositories. The invasion resulted in several changes in ecosystem dynamics. Despite topological similarities between pre- and postinvasion food webs, species loss occurred corresponding to a minor decrease in functional groups. Invaders occupied all of the preinvasion functional guilds, with the exception of four incumbent guilds that were lost and one new guild, corroborating the notion that invaders replace incumbents and fill preexisting niche space. Overall, models exhibited strong resistance to secondary extinction, although the postinvasion community had a lower threshold of collapse and more variable response to perturbation. We interpret these changes in dynamics as a decrease in stability, despite similarities in overall structure. Changes in food web structure and functioning resulting from the invasion suggest that conservation efforts may need to focus on preserving functional diversity if more diverse ecosystems are not inherently more stable.
The classical taxonomy of fossil invertebrates is based on subjective judgments of morphology, which can cause confusion, because there are no codified standards for the classification of genera. Here, we explore the validity of the genus taxonomy of 75 species and morphospecies of the Follicucullidae, a late Paleozoic family of radiolarians, using a new method, Hayashi's quantification theory II (HQT-II), a general multivariate statistical method for categorical datasets relevant to discriminant analysis. We identify a scheme of 10 genera rather than the currently accepted 3 genera (Follicucullus, Ishigaconus, and Parafollicucullus). As HQT-II cannot incorporate stratigraphic data, a phylogenetic tree of Follicucullidae was reconstructed for 38 species using maximum parsimony. Six lineages emerged, roughly in concordance with the results of HQT-II. Combined with parsimony ancestral state reconstruction, the ancestral group of this family is Haplodiacanthus. Five other groups were discriminated, the Parafollicucullus, Curvalbaillella, Pseudoalbaillella, Longtanella, and Follicucullus–Cariver lineages. The morphological evolution of these lineages comprises a minimum essential list of eight states of four traits. HQT-II is a novel discriminant analytical multivariate method that may be of value in other taxonomic problems of paleobiology.
Taphonomic processes are informative about the magnitude and timing of paleoecological changes but remain poorly understood with respect to freshwater invertebrates in spring-fed rivers and streams. We compared taphonomic alteration among freshwater gastropods in live, dead (surficial shell accumulations), and fossil (late Pleistocene–early Holocene in situ sediments) assemblages from two Florida spring-fed systems, the Wakulla and Silver/Ocklawaha Rivers. We assessed taphonomy of two gastropod species: the native Elimia floridensis (n = 2504) and introduced Melanoides tuberculata (n = 168). We quantified seven taphonomic attributes (aperture condition, color, fragmentation, abrasion, juvenile spire condition, dissolution, and exterior luster) and combined those attributes into a total taphonomic score (TT). Fossil E. floridensis specimens exhibited the greatest degradation (highest TT scores), whereas live specimens of both species were least degraded. Specimens of E. floridensis from death assemblages were less altered than fossil specimens of the same species. Within death assemblages, specimens of M. tuberculata were significantly less altered than specimens of E. floridensis, but highly degraded specimens dominated in both species. Radiocarbon dates on fossils clustered between 9792 and 7087 cal BP, whereas death assemblage ages ranged from 10,692 to 1173 cal BP. Possible explanations for the observed taphonomic patterns include: (1) rapid taphonomic shell alteration, (2) prolonged near-surface exposure to moderate alteration rates, and/or (3) introduction of reworked fossil shells into surficial assemblages. Combined radiocarbon dates and taphonomic analyses suggest that all these processes may have played a role in death assemblage formation. In these fluvial settings, shell accumulations develop as a complex mixture of specimens derived from multiple sources and characterized by multimillennial time-averaging. These findings suggest that, when available, fossil assemblages may be more appropriate than death assemblages for assessing preindustrial faunal associations and recent anthropogenic changes in freshwater ecosystems.
Hadrosaurid dinosaurs, the dominant large-bodied terrestrial herbivores in most Laurasian Late Cretaceous ecosystems, have an exceptional fossil record consisting of many species known from partial ontogenetic series, making them an ideal clade with which to conduct life-history studies. Previous research considered the Dinosaur Park Formation (DPF) of Alberta as an attritional, or time-averaged, sample and interpreted size–frequency distribution of long bones collected from the DPF with three size classes to suggest that hadrosaurids from the DPF attained near-asymptotic body size in under 3 years. This conflicted with previously published osteohistological estimates of 6+ years for penecontemporaneous hadrosaurids from the Two Medicine Formation (TMF) of Montana, suggesting either extreme variation in hadrosaurid growth rates or that size–frequency distributions and/or osteohistology and growth modeling inaccurately estimate ontogenetic age.
We tested the validity of the previously proposed size–age relationship of hadrosaurids from the DPF by significantly increasing sample size and combining data from size–frequency distributions and osteohistology across multiple long-bone elements. The newly constructed size–frequency distributions typically reveal four relatively distinct size–frequency peaks that, when integrated with the osteohistological data, aligned with growth marks. The yearling size class was heavily underrepresented in the size–frequency distribution. If not due to preservation, this suggests that either juvenile (<2 years of age) hadrosaurids from the DPF had increased survivorship following an initially high nestling mortality rate or that yearlings were segregated from adults. A growth-curve analysis revealed asymptotic body size was attained in approximately 7 years, which is consistent with hadrosaurids from the TMF. The data suggest size–frequency distributions of attritional samples underestimate age and overestimate growth rates, but when paired with osteohistology can provide unique life-history insights.
The Strophomenata, which includes two large orders, the Strophomenida and Productida, is the largest group of Paleozoic brachiopods. Nearly all uncemented strophomenatans possessed an unusual concave brachial valve. Most of these have been considered to have lived epifaunally, but had they rested on the seafloor, not only would they have faced intense predation, but their physical instability would have been fatal. I conclude that nearly all strophomenatans, like similar concavo-convex pectinid bivalves, lived infaunally by ejecting water to create a pit into which they descended, to be protected by sediment covering the concave valve. Strophomenatans have been discovered with this sediment preserved in place. If exhumed and turned upside down, a strophomenatan could have righted itself by squirting water. Many productides had anchoring spines, and some had hinge areas with stabilizing flanges. Small spines on the brachial valves of some productides served to trap disguising sediment. Evolutionary loss of hinge teeth within both the Strophomenida and Productida reduced the friction of valve clapping. Partly because of their slender shape, strophomenides were typically more vulnerable to exhumation than productides. Strophomenides also ejected water less effectively than productides and would have been less adept at righting themselves. The virtual disappearance of the strophomenides during the Devonian can be attributed to their vulnerability to intensifying benthic bulldozing and predation. The success of the productides during the late Paleozoic can be attributed to their relatively deep sequestration in the sediment and ability to right themselves and reburrow effectively when exhumed and overturned.
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