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Biodiversity hotspots are increasingly recognized as areas of high taxonomic and functional diversity. These hotspots are dynamic and shift geographically over time in response to environmental change. To identify drivers of the origin, evolution, and persistence of diversity hotspots,we investigated the diversity patterns of reef-building corals (Scleractinia) in the Central Indo-Pacific, a marine biodiversity hotspot for the last 25 Myr. We used the scleractinian fossil record (based on literature and a newly acquired fossil collection) to examine the taxonomic and functional diversity of corals from the Eocene to Pliocene. Our data identify potential drivers of coral diversity through time (and space) in the Central Indo-Pacific by constraining the timing of taxonomic turnover events and correlating them with known environmental changes. Increases in taxonomic diversity, high origination rates, and changes in abundance of functional character states indicate that the origin of the Central Indo-Pacific hotspot occurred during the Oligocene, initially through proliferation of pre-existing taxa and then by origination of new taxa. In contrast to taxonomic diversity, overall functional diversity of Central Indo-Pacific reef-building corals remained constant from the Eocene to the Pliocene. Our results identify global sea level as a main driver of diversity increase that, together with local tectonics, regulates availability of suitable habitats. Moreover, marine biodiversity hotspots develop from both the accumulation of taxa from older biodiversity hotspots and origination of new taxa. Our study demonstrates the utility of a combined literature-based and new collection approach for gaining new insights into the timing, drivers, and development of tropical biodiversity hotspots.
One of the longest-lived, noncolonial animals on the planet today is a bivalve that attains life spans in excess of 500 years and lives in a cold, seasonally food-limited setting. Separating the influence of temperature and food availability on life span in modern settings is difficult, as these two conditions covary. The life spans of fossil animals can provide insights into the role of environment in the evolution of extreme longevity that are not available from studies of modern taxa. We examine bivalves from the unique, nonanalogue, warm and high-latitude setting of Seymour Island, Antarctica, during the greenhouse intervals of the Late Cretaceous and Paleogene. Despite significant sampling limitations, we find that all 11 species examined are both slow growing and long-lived, especially when compared with modern bivalves living in similar temperature settings. While cool temperatures have long been thought to be a key factor in promoting longevity, our findings suggest an important role for caloric restriction brought about by the low and seasonal light regime of the high latitudes. Our life-history data, spanning three different families, emphasize that longevity is in part governed by environmental rather than solely phylogenetic or ecologic factors. Such findings have implications for both modern and ancient latitudinal diversity gradients, as a common correlate of slow growth and long life is delayed reproduction, which limits the potential for evolutionary change. While life spans of modern bivalves are well studied, data on life spans of fossil bivalves are sparse and largely anecdotal. Life histories of organisms from deep time can not only elucidate the controls on life span but also add a new dimension to our understanding of macroevolutionary patterns.
Recent studies of eurypterid paleoecology suggest that formation of eurypterid Lagerstätten in the mid-Paleozoic of Laurentia was controlled by the presence of an ecological—taphonomic window that recurred predictably in nearshore, marginal environments during transgressions. We tested this hypothesis by performing a high-resolution taxonomic, environmental, and stratigraphic survey and quantitative analysis of all Silurian—Lower Devonian eurypterid-bearing intervals in the Appalachian basin, the most prolific region for eurypterid remains in the world. Canonical correspondence analysis of sedimentological and faunal associations revealed a strong lithologic gradient between groupings of eurypterid genera and associated taxa across the basin, and a significant association of eurypterids with microbialites (thrombolites, stromatolites) and evaporitic structures. Field observations confirmed that, stratigraphically, eurypterids in the basin frequently occur above the microbialite structures and beneath evaporites and other indicators of increased salinity or subaerial exposure. Following interpretation of these features within a sequence stratigraphic framework, we present a preservational model in which (1) eurypterids inhabited nearshore settings following freshening conditions concomitant with minor transgressions, (2) their remains were subsequently buried by storms or microbialite sediment baffling, and (3) subsequent long-term preservation of tissues was facilitated by regression and cyclical shallowing-up successions that promoted hypersalinity and anoxia. In the central and southern region of the basin, where microbial structures and evidence for hypersalinity are less common, a similar pattern of cyclical shallowing-upward deposition within eurypterid-bearing units holds. Thus, eurypterid preservation appears to reflect a combination of ecological preferences and abiotic conditions that promoted inhabitation and eventual preservation within the same setting. This study provides the first quantitative support for a sea level—based control on preservation of eurypterids and adds to the growing body of evidence that suggests that analysis of exceptional preservation in the fossil record benefits from interpretation within a sequence stratigraphic framework.
Sexual dimorphism is thought to have evolved via selection on both sexes. Ostracodes display sexual shape dimorphism in adult valves; however, no previous studies have addressed temporal changes on evolutionary timescales or examined the relationships between sexual shape dimorphism and selection pressure and between sexual shape dimorphism and juvenile shape. Temporal changes in sexually dimorphic traits result from responses of these traits to selection pressure. Using the Gaussian mixture model for the height/length ratio, a valve-shape parameter, we identified sexual differences in the valve shape of Krithe dolichodeira s.l. from deep-sea sediments of the Paleocene (62.6–57.6 Ma) and estimated the proportion of females in the fossil populations at 11 time intervals. Because the proportion of females in a population is altered by the mortality rate of adult males, it is reflective of selection pressure on males. We attempted to correlate the height/length ratios between the sexes with the proportion of females, taking into consideration that the valve shape was not linked with the selection pressure on males. In time-series data of the height/ length ratio, both sexes indicate no significant changes on evolutionary timescales, even though the sex ratio of the population changed from female skewed to male skewed during the late Paleocene. The sexual shape dimorphism was not driven by sexual selection. The static allometry between the height/length ratio and length indicates that the sexual shape dimorphismdid not function for sexual display. The absence of change over time in the female allometric slope suggests that the evolution of valve shape was constrained by stasis.
A submarine cave is a unique environment that is dark, food limited, semi-isolated from the outside, and sheltered from wave action. However, our knowledge of the long-term change in submarine-cave ecosystems remains limited. We document here the community-scale responses toward long-term change in a submarine cave, Daidokutsu in Okinawa in southern Japan. Using both metazoans (ostracods and bivalves) and protozoans (larger benthic foraminiferans) in two sediment cores obtained from the cave, we reconstruct the faunal and diversity changes of the past 7 Kyr. All taxonomic groups showed long-term, gradual linear change of faunal composition from predominantly open-water taxa to predominantly cave taxa, and ostracods showed short-term variability of species diversity. The long-term faunal trend probably reflects gradual isolation of the cave ecosystem due to coral reef development (i.e., development of the cave ceiling) during periods of the Holocene transgression and subsequent sea-level highstand. The short-term diversity changes show substantial similarity to centennial- to millennial-scale Holocene Asian monsoon variability. Ostracod species diversity peaks tend to correspond with periods of strong East Asian winter monsoons. The results indicate that limestone submarine-cave ecosystems, an important cryptic habitat, developed gradually during the Holocene and may be sensitive to rapid climate changes.
Sauropodomorpha included the largest known terrestrial vertebrates and was the first dinosaur clade to achieve a global distribution. This success is associated with their early adoption of herbivory, and sauropod gigantism has been hypothesized to be a specialization for bulk feeding and obligate high-fiber herbivory. Here, we apply a combination of biomechanical character analysis and comparative phylogenetic methods with the aim of quantifying the evolutionary mechanics of the sauropodomorph feeding apparatus. We test for the role of convergence to common feeding function and divergence toward functional optima across sauropodomorph evolution, quantify the rate of evolution for functional characters, and test for coincident evolutionary rate shifts in craniodental functional characters and body mass. Results identify a functional shift toward increased cranial robustness, increased bite force, and the onset of static occlusion at the base of the Sauropoda, consistent with a shift toward bulk feeding. Trends toward similarity in functional characters are observed in Diplodocoidea and Titanosauriformes. However, diplodocids and titanosaurs retain significant craniodental functional differences, and evidence for convergent adoption of a common “adaptive zone” between them is weak. Modeling of craniodental character and body-mass evolution demonstrates that these functional shifts were not correlated with evolutionary rate shifts. Instead, a significant correlation between body mass and characters related to bite force and cranial robustness suggests a correlated-progression evolutionary mode, with positive-feedback loops between body mass and dietary specializations fueling sauropod gigantism.
Numerical and taxonomic resolution of compositional data sets affects investigators' abilities to detect and measure relationships between communities and environmental factors. We test whether varying numerical (untransformed, square-root- and fourth-root-transformed relative abundance and presence—absence data) and taxonomic (species, genera, families) resolutions reveals different insights into early to middle Miocene molluscan communities along bathymetric and salinity gradients. The marine subtidal has a more even species-abundance distribution, a higher number of rare species, and higher species:family and species:genus ratios than the three habitats—marine and estuarine intertidal, estuarine subtidal—with higher fluctuations in salinity and other physical parameters. Taxonomic aggregation and numerical transformation of data result in very different ordinations, although all habitats differ significantly from one another at all taxonomic and numerical levels. Rank correlations between species-level and higher-taxon, among-sample dissimilarities are very high for proportional abundance and decrease strongly with increasing numerical transformation, most notably in the two intertidal habitats. The proportion of variation explained by depth is highest for family-level data, decreases gradually with numerical transformation, and is higher in marine than in estuarine habitats. The proportion of variation explained by salinity is highest for species-level data, increases gradually with numerical transformation, and is higher in subtidal than in intertidal habitats. Therefore, there is no single best numerical and taxonomic resolution for the discrimination of communities along environmental gradients: the “best” resolution depends on the environmental factor considered and the nature of community response to it. Different numerical and taxonomic transformations capture unique aspects of metacommunity assembly along environmental gradients that are not detectable at a single level of resolution. We suggest that simultaneous analyses of community gradients at multiple taxonomic and numerical resolutions provide novel insights into processes responsible for spatial and temporal community stability.
Studying the deep-time origins of macroecological phenomena can help us to understand their long-term drivers. Given the considerable spatiotemporal bias of the terrestrial fossil record, it behooves us to understand how much biological information is lost. The aim of this study is to establish whether latitudinal diversity gradients are detectable in a biased terrestrial fossil record. I develop a simulated fossilization approach, weighting the probability of terrestrial mammal species appearing in the fossil record based on body size and geographic-range size; larger species with larger range sizes are more likely to enter the fossil record. I create simulated fossil localities from the modern North American mammal record. I vary the percentage of species successfully fossilized and estimate the magnitude of the latitudinal diversity gradient (slope of the richness gradient and degree of species turnover). I find that estimates of the latitudinal diversity gradient are sensitive to the loss of species with small body size and geographic-range sizes. In some cases, simulated fossil-record bias completely obliterates evidence of declining richness with latitude, a phenomenon that is not ameliorated by the application of nonparametric richness estimation. However, if the rate of preservation is medium (50% of species) to high (75% of species), the magnitude of the latitudinal diversity gradient can be reliably estimated. Similarly, changes in the diversity gradient estimates are largely explained by differences in the diversity—climate relationship among iterations, suggesting that these relationships may be measurable in the fossil record.
The demonstration of sexual dimorphism in the fossil record can provide vital information about the role that sexual selection has played in the evolution of life. However, statistically robust inferences of sexual dimorphism in fossil organisms are exceedingly difficult to establish, owing to issues of sample size, experimental control, and methodology. This is particularly so in the case of dinosaurs, for which sexual dimorphism has been posited in many species, yet quantifiable data are often lacking. This study presents the first statistical investigation of sexual dimorphism across Dinosauria. It revisits prior analyses that purport to find quantitative evidence for sexual dimorphism in nine dinosaur species. After the available morphological data were subjected to a suite of statistical tests (normality and unimodality tests and mixture modeling), no evidence for sexual dimorphism was found in any of the examined taxa, contrary to conventional wisdom. This is not to say that dinosaurs were not sexually dimorphic (phylogenetic inference suggests they may well have been), only that the available evidence precludes its detection. A priori knowledge of the sexes would greatly facilitate the assessment of sexual dimorphism in the fossil record, and it is suggested that unambiguous indicators of sex (e.g., presence of eggs, embryos, medullary bone) be used to this end.
Most geometric morphometric studies are underpinned by sets of photographs of specimens. The camera lens distorts the images it takes, and the extent of the distortion will depend on factors such as the make and model of the lens and camera and user-controlled variation such as the zoom of the lens. Any study that uses populations of geometric data digitized from photographs will have shape variation introduced into the data set simply by the photographic process. We illustrate the nature and magnitude of this error using a 30-specimen data set of Recent New Zealand Mactridae (Mollusca: Bivalvia), using only a single camera and camera lens with four different photographic setups. We then illustrate the use of retrodeformation in Adobe Photoshop and test the magnitude of the variation in the data set using multivariate Procrustes analysis of variance. The effect of photographic method on the variance in the data set is significant, systematic, and predictable and, if not accounted for, could lead to misleading results, suggest clustering of specimens in ordinations that has no biological basis, or induce artificial oversplitting of taxa. Recommendations to minimize and quantify distortion include: (1) that studies avoid mixing data sets from different cameras, lenses, or photographic setups; (2) that studies avoid placing specimens or scale bars near the edges of the photographs; (3) that the same camera settings are maintained (as much as practical) for every image in a data set; (4) that care is taken when using full-frame cameras; and (5) that a reference grid is used to correct for or quantify distortion.