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Documenting sexual dimorphism for structures that exhibit indeterminate growth can be more difficult than for structures exhibiting determinate growth. Most proboscidean tusks are ever-growing structures that change size and shape throughout life. Sexual dimorphism is pronounced in tusks of mature individuals, but the external form of tusks offers no clear evidence of maturation, and it is difficult to distinguish a young male's tusk from that of an older female. Thus, with previous approaches, knowledge of age was often required to assess sex from tusk measurements. This study examines sexual dimorphism of American mastodon (Mammut americanum) tusks through principal components analysis to determine which aspects of tusk form contribute most strongly to the variance among measurements and to explore the relationship between tusk form and individual age and sex. Twenty-one mastodon tusks from the Great Lakes region were evaluated in two analyses, the first focusing on geometrically distinct aspects of tusk form and the second adding measurements that reflect ontogenetic changes in a single aspect of morphology (circumference). Both analyses separated mastodons by sex (PC-I) and sorted them by age (PC-II). The distribution of tusks on the PC-II versus PC-I plane provides better discrimination of sex than univariate or bivariate methods because tusks of similar size and opposite sex appear near opposite ends of an age spectrum. The second analysis enhances sorting by age, thereby clarifying assessment of sex. This work contributes to studies of mastodon paleobiology by presenting a reliable method for assessing the sex of an individual from tusk measurements without requiring independent knowledge of age.
The photosynthetic gas exchange capacities of early angiosperms remain enigmatic. Nevertheless, many hypotheses about the causes of early angiosperm success and how angiosperms influenced Mesozoic ecosystem function hinge on understanding the maximum capacity for early angiosperm metabolism. We applied structure-functional analyses of leaf veins and stomatal pore geometry to determine the hydraulic and diffusive gas exchange capacities of Early Cretaceous fossil leaves. All of the late Aptian–early Albian angiosperms measured possessed low vein density and low maximal stomatal pore area, indicating low leaf gas exchange capacities in comparison to modern ecologically dominant angiosperms. Gas exchange capacities for Early Cretaceous angiosperms were equivalent or lower than ferns and gymnosperms. Fossil leaf taxa from Aptian to Paleocene sediments previously identified as putative stem-lineages to Austrobaileyales and Chloranthales had the same gas exchange capacities and possibly leaf water relations of their living relatives. Our results provide fossil evidence for the hypothesis that high leaf gas exchange capacity is a derived feature of later angiosperm evolution. In addition, the leaf gas exchange functions of austrobaileyoid and chloranthoid fossils support the hypothesis that comparative research on the biology of living basal angiosperm lineages reveals genuine signals of Early Cretaceous angiosperm ecophysiology.
Studies of crinoid morphology have been pivotal in understanding the constraints on the range of morphology within a clade as well as the patterns of disparity throughout the Phanerozoic. Newly discovered and described faunas and recent study of early Paleozoic crinoid diversity provide an ideal opportunity to reanalyze Ordovician through Early Silurian crinoid disparity with more complete taxonomic coverage and finer stratigraphic resolution. Using the coarse stratigraphic binning of Foote (1999), the updated morphologic data set has a similar disparity pattern to those previously reported for the early Paleozoic. However, with the more resolved stratigraphic binning used by Peters and Ausich (2008), a significant difference exists between the original and current data sets. Both data sets have a pronounced disparity high during the late Middle Ordovician. However, the updated disparity curve has a much higher initial disparity during the Early Ordovician and a pronounced rise in disparity during the Silurian recovery. Examination of differential sampling, proportions of the crinoid orders through time, and methods of coding characters indicate these factors have little effect on the pattern of crinoid disparity. The Silurian morphospace expansion occurs primarily within disparids and coincides with the origination of the myelodactylids. These findings corroborate the rapid expansion of morphospace during the Ordovician. However, crinoid disparity did not remain static and, although less frequent than during the initial radiation, new body plans evolved following the Ordovician Extinction (e.g., the myelodactylids). These results are consistent with the hypothesis of ecology constraining the limits on morphologic disparity at the class level.
We have devised a simple model for assessing the role of development in shaping the evolution of morphological disparity. Disparity of a clade at any given time is expressed in terms of the developmental dynamics that lead to the variety of adult morphotypes observed. We use assumed phenotypic manifestations of developmental processes, as they could be detected from allometric characterizations, to distinguish a few, nonexclusive types of evolutionary changes in ontogeny. On the basis of this formalization, we describe the diversification of hypothetical clades, using the standard curve of adult morphological disparity, the curve of juvenile disparity, and the curve of allometric disparity, the latter quantifying the diversification of clades in allometric space. Contrasts of these curves reflect the underlying developmental scheme that drives temporal changes in disparity. We then vary the parameters of the model to assess the expected signature of each metric under specific conditions: changes in the relative frequencies of the types of evolutionary developmental changes, changes in the transition magnitude attached to each of them, and effects of temporal variation in average adult size on disparity curves and patterns of morphospace occupation. Results emphasize the potential contribution of these proxies for developmental dynamics—juvenile morphological disparity, allometric disparity, and average adult size—in enriching the interpretation of standard disparity curves and the description of clade histories, with possible process-oriented inferences.
The modern structure of marine benthic ecosystems was largely established during the Jurassic and Early Cretaceous (200-100 Ma), a transition that has been termed the Mesozoic Marine Revolution (MMR). Although it has been suggested that the MMR marks an increase in the average energy consumption of marine animal ecosystems, this hypothesis has not been evaluated quantitatively. In this study, we integrate body size and abundance data from the fossil record with physiological data from living representatives to estimate mean per capita metabolic rates of tropical to subtropical assemblages of shallow-marine gastropods—a major component of marine ecosystems throughout the Meso-Cenozoic—both before and after the MMR. We find that mean per capita metabolic rate rose by ∼150% between the Late Triassic and Late Cretaceous and remained relatively stable thereafter. The most important factor governing the increase in metabolic rate was an increase in mean body size. In principle, this size increase could result from secular changes in sampling and taphonomic biases, but these biases are suggested to yield decreases rather than increases in mean size. Considering that post-MMR gastropod diversity is dominated by predators, the net primary production required to supply the energetic needs of the average individual increased by substantially more than 150%. These data support the hypothesis that benthic energy budgets increased during the MMR, possibly in response to rising primary productivity.
Niche conservatism is increasingly recognized in diverse modern ecological settings, and it influences many aspects of modern ecosystems, including speciation mechanisms, community structure, and response to climate change. Here, we investigate the stability of niches with benthic marine invertebrates along a Late Ordovician onshore-offshore gradient on the Cincinnati Arch in the eastern United States. Using a Gaussian niche model characterized by peak abundance, preferred environment, and environmental tolerance, with these parameters estimated through weighted averaging and logistic regression, we find evidence of strong niche conservatism in peak abundance and preferred environment, particularly for abundant taxa. This conservatism is maintained in successive depositional sequences and through the nearly 9–10 Myr study interval. Environmental tolerance shows no evidence of conservatism, although numerical simulations suggest that the error rates in estimates of this parameter are so high that they could overwhelm evidence of conservatism. These numerical simulations also indicate that both weighted averaging and logistic regression produce useful estimates of peak abundance and preferred environment, with slightly better results for weighted averaging. This evidence for niche conservatism suggests that long-term shifts of higher taxa of marine invertebrates into deeper water are primarily the result of differential rates of origination and extinction. These results also add to the evidence of long periods of relatively stable ecosystems despite regional environmental perturbations, and they constrain the causes of peaked patterns in occupancy.
Colony-wide feeding currents are a common feature of many bryozoan colonies. These feeding currents are centered on excurrent macular chimneys that expel previously filtered water away from the colony surface. In some bryozoans these macular chimneys consist of a branching channel network that converges at a point in the center of the chimney. The bifurcating channels of the maculae are analogous to a stream channel network in a closed basin with centripetal drainage. The classical methods of stream channel network analysis from geomorphology are here used to quantitatively analyze the number and length of macular channels in bryozoans. This approach is applied to a giant branch of the trepostome bryozoan Tabulipora from the Early Permian Kim Fjelde Formation in North Greenland. Its large size allowed 18 serial tangential peels to be made through the 8-mm-thick exozone. The peels intersected two stellate maculae as defined by contiguous exilapores. The lengths of 1460 channels radiating from the maculae were measured and their Horton-Strahler stream order and Shreve magnitude scored.
We hypothesize that if fossil bryozoan maculae function as excurrent water chimneys, then they should conform to Horton's laws of stream networks and behave like closed basins with centripetal drainage. Results indicate that the stellate maculae in this bryozoan behaved liked stream channel networks exhibiting landscape maturation and stream capture. They conformed to the Law of Stream Number. They have a Bifurcation Ratio that falls within the range of natural stream channel networks. They showed a pattern opposite that expected by the Law of Stream Lengths in response to behavior characteristic of a centripetal drainage pattern in a closed basin. Thus, the stellate maculae in this bryozoan probably functioned as excurrent water chimneys with the radiating channels serving to efficiently collect the previously filtered water, conducting it to the central chimney for expulsion away from the colony surface.
Phanerozoic trends in shell and life habit traits linked to postmortem durability were evaluated for the most common fossil brachiopod, gastropod, and bivalve genera in order to test for changes in taphonomic bias. Using the Paleobiology Database, we tabulated occurrence frequencies of genera for 48 intervals of ∼11 Myr duration. The most frequently occurring genera, cumulatively representing 40% of occurrences in each time bin, were scored for intrinsic durability on the basis of shell size, reinforcement (ribs, folds, and spines), life habit, and mineralogy.
Shell durability is positively correlated with the number of genera in a time bin, but durability traits exhibit different temporal patterns across higher taxa, with notable offsets in the timing of changes in these traits. We find no evidence for temporal decreases in durability that would indicate taphonomic bias at the Phanerozoic scale among commonly occurring genera. Also, all three groups show a remarkable stability in mean shell size through the Phanerozoic, an unlikely pattern if strong size-filtering taphonomic megabiases were affecting the fossil record of shelly faunas. Moreover, small shell sizes are attained in the early Paleozoic in brachiopods and in the latest Paleozoic in gastropods but are steady in bivalves; unreinforced shells are common to all groups across the entire Phanerozoic; organophosphatic and aragonitic shells dominate only the oldest and youngest time bins; and microstructures having high organic content are most common in the oldest time bins.
In most cases, the timing of changes in durability-related traits is inconsistent with a late Mesozoic Marine Revolution. The post-Paleozoic increase in mean gastropod reinforcement occurs in the early Triassic, suggesting either an earlier appearance and expansion of durophagous predators or other drivers. Increases in shell durability hypothesized to be the result of increased predation in the late Mesozoic are not evident in the common genera examined here. Infaunal life habit does increase in the late Mesozoic, but it does not become more common than levels already attained during the Paleozoic, and only among bivalves does the elevated late Mesozoic level persist through the Holocene.
These temporal patterns suggest control on the occurrence of durability-related traits by individual evolutionary histories rather than taphonomic megabiases. Our findings do not mean taphonomic biases are absent from the fossil record, but rather that their effects apparently have had little net effect on the relative occurrence of shell traits generally thought to confer higher preservation potential over long time scales.
Quantifying the effects of taphonomic processes on species abundances in time-averaged death assemblages (DAs) is pivotal for paleoecological inference. However, fidelity estimates based on conventional “live-dead” comparisons are fundamentally ambiguous: (1) data on living assemblages (LAs) are based on a very short period of sampling and thus do not account for biological variability in the LA, (2) LAs are sampled at the same time as the DA and thus do not necessarily reflect past LAs that contributed to the DA, (3) compositions of LAs and DAs can be autocorrelated owing to shared cohorts, and (4) fidelity estimates are cross-scale estimates because DAs are time-averaged and LAs are not. Some portion of raw (total) live-dead (LD) variation in species composition thus arises from incomplete sampling of LAs and from biological temporal variation among LAs (together = premortem component of LD variation), as contrasted with new variation created by interspecific variation in population turnover and preservation rates and by the time-averaging of skeletal input (together = postmortem component of LD variation). To tackle these problems, we introduce a modified test for homogeneity of multivariate dispersions (HMD) in order to (1) account for temporal autocorrelation in composition between LAs and DAs and (2) decompose total LD compositional variation into premortem and postmortem components, and we use simulations to evaluate the contribution of within-habitat time-averaging on the postmortem component. Applying this approach to 31 marine molluscan data sets, each consisting of spatial replicates of LAs and DAs in a single habitat, we find that total LD variation is driven largely by variation among LAs. However, genuinely postmortem processes have significant effects on composition in 25–65% of data sets (depending on the metric) when the effects of temporal autocorrelation are taken into account using HMD. Had we ignored the effects of autocorrelation, the effects of postmortem processes would have been negligible, inflating the similarity between LAs and DAs. Simulations show that within-habitat time-averaging does not increase total LD variation to a large degree—it increases total LD variation mainly via increasing species richness, and decreases total LD variation by reducing dispersion among DAs. The postmortem component of LD variation thus arises from differential turnover and preservation and multi-habitat time-averaging. Moreover, postmortem processes have less effect on the compositions of DAs in habitats characterized by high variability among LAs than they have on DAs in temporally stable habitats, a previously unrecognized first-order factor in estimating postmortem sources of compositional variation in DAs.