Investigating Biotic Interactions in Deep Time

Recent renewed interest in using fossil data to understand how biotic interactions have shaped the evolution of life is challenging the widely held assumption that long-term climate changes are the primary drivers of biodiversity change. New approaches go beyond traditional richness and co-occurrence studies to explicitly model biotic interactions using data on fossil and modern biodiversity. Important developments in three primary areas of research include analysis of (i) macroevolutionary rates, (ii) the impacts of and recovery from extinction events, and (iii) how humans (Homo sapiens) affected interactions among non-human species. We present multiple lines of evidence for an important and measurable role of biotic interactions in shaping the evolution of communities and lineages on long timescales.Peer reviewe

A framework for evaluating the influence of climate, dispersal limitation, and biotic interactions using fossil pollen associations across the late Quaternary

Environmental conditions, dispersal lags, and interactions among species are major factors structuring communities through time and across space. Ecologists have emphasized the importance of biotic interactions in determining local patterns of species association. In contrast, abiotic limits, dispersal limitation, and historical factors have commonly been invoked to explain community structure patterns at larger spatiotemporal scales, such as the appearance of late Pleistocene no-analog communities or latitudinal gradients of species richness in both modern and fossil assemblages. Quantifying the relative influence of these processes on species co-occurrence patterns is not straightforward. We provide a framework for assessing causes of species associations by combining a null-model analysis of co-occurrence with additional analyses of climatic differences and spatial pattern for pairs of pollen taxa that are significantly associated across geographic space. We tested this framework with data on associations among 106 fossil pollen taxa and paleoclimate simulations from eastern North America across the late Quaternary. The number and proportion of significantly associated taxon pairs increased over time, but only 449 of 56 194 taxon pairs were significantly different from random. Within this significant subset of pollen taxa, biotic interactions were rarely the exclusive cause of associations. Instead, climatic or spatial differences among sites were most frequently associated with significant patterns of taxon association. Most taxon pairs that exhibited co-occurrence patterns indicative of biotic interactions at one time did not exhibit significant associations at other times. Evidence for environmental filtering and dispersal limitation was weakest for aggregated pairs between 16 and 11 kyr BP, suggesting enhanced importance of positive species interactions during this interval. The framework can thus be used to identify species associations that may reflect biotic interactions because these associations are not tied to environmental or spatial differences. Furthermore, temporally repeated analyses of spatial associations can reveal whether such associations persist through time

Spatiotemporal relationships among Late Pennsylvanian plant assemblages: Palynological evidence from the Markley Formation, West Texas, U.S.A.

The Pennsylvanian lowlands of western Pangea are best known for their diverse wetland floras of arborescent and herbaceous ferns, and arborescent horsetails and clubmosses. In apparent juxtaposition, a very different kind of flora, dominated by a xerophilous assemblage of conifers, taeniopterids and peltasperms, is occasionally glimpsed. Once believed to represent upland or extrabasinal floras from well-drained portions of the landscape, these dryland floras more recently have been interpreted as lowland assemblages growing during drier phases of glacial/interglacial cycles. Whether Pennsylvanian dryland and wetland floras were separated spatially or temporally remains an unsettled question, due in large part to taphonomic bias toward preservation of wetland plants. Previous paleobotanical and sedimentological analysis of the Markley Formation of latest Pennsylvanian (Gzhelian) age, from north central Texas, U.S.A, indicates close correlation between lithofacies and distinct dryland and wetland megaflora assemblages. Here we present a detailed analysis one of those localities, a section unusual in containing abundant palynomorphs, from the lower Markley Formation. Paleobotanical, palynological and lithological data from a section thought to represent a single interglacial/glacial phase are integrated and analyzed to create a complex picture of an evolving landscape. Megafloral data from throughout the Markley Formation show that conifer-dominated dryland floras occur exclusively in highly leached kaolinite beds, likely eroded from underlying soils, whereas a mosaic of wetland floras occupy histosols, ultisols, and fluvial overbank deposits. Palynological data largely conform to this pattern but reveal a more complex picture. An assemblage of mixed wetland and dryland palynofloral taxa is interpolated between a dryland assemblage and an overlying histosol containing wetland taxa. In this section, as well as elsewhere in the Markley Formation, kaolinite and overlying organic beds appear to have formed as a single genetic unit, with the kaolinite forming an impermeable aquiclude upon which a poorly drained wetland subsequently formed. Within a single inferred glacial/interglacial cycle, lithological data indicate significant fluctuations in water availability tracked by changes in palynofloral and megafloral taxa. Palynology reveals that elements of the dryland floras appear at low abundance even within wetland deposits. The combined data indicate a complex pattern of succession and suggest a mosaic of dryland and wetland plant communities in the Late Pennsylvanian. Our data alone cannot show whether dryland and wetland assemblages succeed one another temporally, or coexisted on the landscape. However, the combined evidence suggests relatively close spatial proximity within a fragmenting and increasingly arid environment

Oxygen-containing aromatic compounds in a Late Permian sediment

In northern Italy the base of the Tesero Oolite Horizon represents the lithological boundary between the Bellerophon and Werfen Formations and marks one of the most dramatic environmental perturbations in Phanerozoic time. The marl, which directly underlies the Tesero Horizon, contains many indicators of environmental change during the end-Permian mass extinction. At Vigo Meano, near Trento, the marl is relatively thermally immature and contains abundant terrestrially-sourced oxygen containing aromatic compounds such as xanthones, dibenzofurans and dibenzo-p-dioxin as well as abundant alkylated naphthalenes

Data from: When conifers took flight: A biomechanical evaluation of an imperfect evolutionary takeoff

Manifera talaris, a voltzian conifer from the late early to middle Permian (ca. 270 Ma) of Texas, is the earliest known conifer to produce winged seeds indicative of autorotating flight. In contrast to autorotating seeds and fruits of extant plants, the ones of M. talaris are exceptional in that they have variable morphology. They bore two wings that produced a range of wing configurations, from seeds with two equal-sized wings to single-winged specimens, via various stages of underdevelopment of one of the wings. To examine the effects of various seed morphologies on aerodynamics and dispersal potential, we studied the flight performance of paper models of three morphotypes: symmetric double-winged, asymmetric double-winged, and single-winged. Using a high-speed camera we identified the mode of descent (plummeting, gliding, autorotation) and quantified descent speed, autorotation frequency, and other flight characteristics. To validate such modeling as an inferential tool, we compared descent of extant analogues (kauri; Agathis australis) with descent of similarly constructed seed models. All three seed morphotypes exhibited autorotating flight behavior. However, double-winged seeds, especially symmetric ones, failed to initiate slow autorotative descent more frequently than single-winged seeds. Even when autorotating, symmetric double-winged seeds descend faster than asymmetric double-winged ones, and descent is roughly twice as fast compared to single-winged seeds. Moreover, the relative advantage that (effectively) single-winged seeds have in slowing descent during autorotation becomes larger as seed weight increases. Hence, the range in seed wing configurations in M. talaris produced a wide variation in potential dispersal capacity. Overall, our results indicate that the evolutionarily novel autorotating winged seeds must have improved conifer seed dispersal, in a time when animal vectors for dispersion were virtually absent. Because of the range in wing configuration, the early evolution of autorotative flight in conifers was a functionally imperfect one, which provides us insight into the evolutionary developmental biology of autorotative seeds in conifers

Chemical consitution of a Permian-Triassic disaster species

One of the most controversial biological proxies of environmental crisis at the close of the Permian is the organic microfossil Reduviasporonites. The proliferation of this disaster species coincides with the mass extinction and numerous geochemical disturbances. Originally Reduviasporonites was assigned to fungi, opportunistically exploiting dying end-Permian forests, but subsequent geochemical data have been used to suggest an algal origin. We have used high-sensitivity equipment, partly designed to detect interstellar grains in meteorites, to reexamine the geochemical signature of Reduviasporonites. Organic chemistry, carbon and nitrogen isotopes, and carbon/nitrogen ratios are consistent with a fungal origin. The use of this microfossil as a marker of terrestrial ecosystem collapse should not be merely discounted. Unequivocally diagnostic data, however, may have been precluded by post-burial replacement of its organic constituents

Kungurian (Cisuralian) conifers and environmental changes: a negative δ13C shift in the flora of Tregiovo (Northern Italy)

The Le Fraine fossil locality, near the Tregiovo village (Trento Province, northern Italy) yields two of the best documented Kungurian (early Permian) plant fossil assemblages of eastern palaeoequatorial Pangea. Both plant assemblages (Tregiovo A and B) are dominated by walchian and voltzian conifers but differ in the relative abundance of the major plant groups. Analyses of the δ13C of the bulk organic carbon throughout the entire stratigraphic succession of Tregiovo show a negative trend towards the upper part of the succession. Several conifer taxa are recognized and taxon-specific δ13C analyses of the conifer remains in both plant fossil assemblages shows the same trend in all taxa. This correlation between the negative shift of the bulk organic carbon and the taxon-specific δ13C analysis shows that the negative δ13C shift recorded along the stratigraphic section is not the result of change in relative abundance within the plant assemblages but instead has an external cause, likely the δ13C values of atmospheric CO2. These results agree well with a negative shift observed in other Kungurian continental successions (e.g., China, South Africa, Australia), supporting a global perturbation in the carbon cycle. The negative shift in δ13C could very well record the last deglaciation phases of the Late Paleozoic Ice Age and the rise of the atmospheric pCO2