Seth Finnegan

Climate change and the past, present, and future of biotic interactions.

Biotic interactions drive key ecological and evolutionary processes and mediate ecosystem responses to climate change. The direction, frequency, and intensity of biotic interactions can in turn be altered by climate change. Understanding the complex interplay between climate and biotic interactions is thus essential for fully anticipating how ecosystems will respond to the fast rates of current warming, which are unprecedented since the end of the last glacial period. We highlight episodes of climate change that have disrupted ecosystems and trophic interactions over time scales ranging from years to millennia by changing species’ relative abundances and geographic ranges, causing extinctions, and creating transient and novel communities dominated by generalist species and interactions. These patterns emerge repeatedly across disparate temporal and spatial scales, suggesting the possibility of similar underlying processes. Based on these findings, we identify knowledge gaps and fruitful areas for research that will further our understanding of the effects of climate change on ecosystems.

The Magnitude and Duration of Late Ordovician–Early Silurian Glaciation

Carbonate isotopes reveal a link between past ocean temperatures and mass extinction. Understanding ancient climate changes is hampered by the inability to disentangle trends in ocean temperature from trends in continental ice volume. We used carbonate “clumped” isotope paleothermometry to constrain ocean temperatures, and thereby estimate ice volumes, through the Late Ordovician–Early Silurian glaciation. We find tropical ocean temperatures of 32° to 37°C except for short-lived cooling by ~5°C during the final Ordovician stage. Evidence for ice sheets spans much of the study interval, but the cooling pulse coincided with a glacial maximum during which ice volumes likely equaled or exceeded those of the last (Pleistocene) glacial maximum. This cooling also coincided with a large perturbation of the carbon cycle and the Late Ordovician mass extinction.

The effect of geographic range on extinction risk during background and mass extinction.

Wide geographic range is generally thought to buffer taxa against extinction, but the strength of this effect has not been investigated for the great majority of the fossil record. Although the majority of genus extinctions have occurred between major mass extinctions, little is known about extinction selectivity regimes during these “background” intervals. Consequently, the question of whether selectivity regimes differ between background and mass extinctions is largely unresolved. Using logistic regression, we evaluated the selectivity of genus survivorship with respect to geographic range by using a global database of fossil benthic marine invertebrates spanning the Cambrian through the Neogene periods, an interval of ≈500 My. Our results show that wide geographic range has been significantly and positively associated with survivorship for the great majority of Phanerozoic time. Moreover, the significant association between geographic range and survivorship remains after controlling for differences in species richness and abundance among genera. However, mass extinctions and several second-order extinction events exhibit less geographic range selectivity than predicted by range alone. Widespread environmental disturbance can explain the reduced association between geographic range and extinction risk by simultaneously affecting genera with similar ecological and physiological characteristics on global scales. Although factors other than geographic range have certainly affected extinction risk during many intervals, geographic range is likely the most consistently significant predictor of extinction risk in the marine fossil record.

Two-phase increase in the maximum size of life over 3.5 billion years reflects biological innovation and environmental opportunity

The maximum size of organisms has increased enormously since the initial appearance of life >3.5 billion years ago (Gya), but the pattern and timing of this size increase is poorly known. Consequently, controls underlying the size spectrum of the global biota have been difficult to evaluate. Our period-level compilation of the largest known fossil organisms demonstrates that maximum size increased by 16 orders of magnitude since life first appeared in the fossil record. The great majority of the increase is accounted for by 2 discrete steps of approximately equal magnitude: the first in the middle of the Paleoproterozoic Era (≈1.9 Gya) and the second during the late Neoproterozoic and early Paleozoic eras (0.6–0.45 Gya). Each size step required a major innovation in organismal complexity—first the eukaryotic cell and later eukaryotic multicellularity. These size steps coincide with, or slightly postdate, increases in the concentration of atmospheric oxygen, suggesting latent evolutionary potential was realized soon after environmental limitations were removed.

Formation of the Isthmus of Panama

Independent evidence from rocks, fossils, and genes converge on a cohesive narrative of isthmus formation in the Pliocene. The formation of the Isthmus of Panama stands as one of the greatest natural events of the Cenozoic, driving profound biotic transformations on land and in the oceans. Some recent studies suggest that the Isthmus formed many millions of years earlier than the widely recognized age of approximately 3 million years ago (Ma), a result that if true would revolutionize our understanding of environmental, ecological, and evolutionary change across the Americas. To bring clarity to the question of when the Isthmus of Panama formed, we provide an exhaustive review and reanalysis of geological, paleontological, and molecular records. These independent lines of evidence converge upon a cohesive narrative of gradually emerging land and constricting seaways, with formation of the Isthmus of Panama sensu stricto around 2.8 Ma. The evidence used to support an older isthmus is inconclusive, and we caution against the uncritical acceptance of an isthmus before the Pliocene.

Extinctions in ancient and modern seas

In the coming century, life in the ocean will be confronted with a suite of environmental conditions that have no analog in human history. Thus, there is an urgent need to determine which marine species will adapt and which will go extinct. Here, we review the growing literature on marine extinctions and extinction risk in the fossil, historical, and modern records to compare the patterns, drivers, and biological correlates of marine extinctions at different times in the past. Characterized by markedly different environmental states, some past periods share common features with predicted future scenarios. We highlight how the different records can be integrated to better understand and predict the impact of current and projected future environmental changes on extinction risk in the ocean.

CARBONATES IN SKELETON-POOR SEAS: NEW INSIGHTS FROM CAMBRIAN AND ORDOVICIAN STRATA OF LAURENTIA

Abstract Calcareous skeletons evolved as part of the greater Ediacaran–Cambrian diversification of marine animals. Skeletons did not become permanent, globally important sources of carbonate sediment, however, until the Ordovician radiation. Representative carbonate facies in a Series 3 (510–501 Ma) Cambrian to Tremadocian succession from western Newfoundland, Canada, and Ordovician successions from the Ibex area, Utah, USA, show that, on average, Cambrian and Tremadocian carbonates contain much less skeletal material than do post-Tremadocian sediments. Petrographic point counts of skeletal abundance within facies and proportional facies abundance in measured sections suggest that later Cambrian successions contain on average <5% skeletal material by volume, whereas the skeletal content of post-Tremadocian Ordovician sections is closer to ∼15%. A compilation of carbonate stratigraphic sections from across Laurentia confirms that post-Tremadocian increase in skeletal content is a general pattern and not unique to the two basins studied. The long interval (∼40 myr) between the initial Cambrian appearance of carbonate skeletons and the subsequent Ordovician diversification of heavily skeletonized organisms provides an important perspective on the Ordovician radiation. Geochemical data increasingly support the hypothesis that later Cambrian oceans were warm and, in subsurface water masses, commonly dysoxic to anoxic. We suggest that surface waters in such oceans would have been characterized by relatively low saturation states for calcite and aragonite. Mid-Ordovician cooling would have raised oxygen concentrations in subsurface water masses, establishing more highly oversaturated surface waters. If correct, these links could provide a proximal trigger for the renewed radiation of heavily skeletonized invertebrates and algae.

Climate change and the selective signature of the Late Ordovician mass extinction

Selectivity patterns provide insights into the causes of ancient extinction events. The Late Ordovician mass extinction was related to Gondwanan glaciation; however, it is still unclear whether elevated extinction rates were attributable to record failure, habitat loss, or climatic cooling. We examined Middle Ordovician-Early Silurian North American fossil occurrences within a spatiotemporally explicit stratigraphic framework that allowed us to quantify rock record effects on a per-taxon basis and assay the interplay of macrostratigraphic and macroecological variables in determining extinction risk. Genera that had large proportions of their observed geographic ranges affected by stratigraphic truncation or environmental shifts at the end of the Katian stage were particularly hard hit. The duration of the subsequent sampling gaps had little effect on extinction risk, suggesting that this extinction pulse cannot be entirely attributed to rock record failure; rather, it was caused, in part, by habitat loss. Extinction risk at this time was also strongly influenced by the maximum paleolatitude at which a genus had previously been sampled, a macroecological trait linked to thermal tolerance. A model trained on the relationship between 16 explanatory variables and extinction patterns during the early Katian interval substantially underestimates the extinction of exclusively tropical taxa during the late Katian interval. These results indicate that glacioeustatic sea-level fall and tropical ocean cooling played important roles in the first pulse of the Late Ordovician mass extinction in Laurentia.

The Ordovician Radiation: A Follow-up to the Cambrian Explosion?'

Abstract There was a major diversification known as the Ordovician Radiation, in the period immediately following the Cambrian. This event is unique in taxonomic, ecologic and biogeographic aspects. While all of the phyla but one were established during the Cambrian explosion, taxonomic increases during the Ordovician were manifest at lower taxonomic levels although ordinal level diversity doubled. Marine family diversity tripled and within clade diversity increases occurred at the genus and species levels. The Ordovician radiation established the Paleozoic Evolutionary Fauna; those taxa which dominated the marine realm for the next 250 million years. Community structure dramatically increased in complexity. New communities were established and there were fundamental shifts in dominance and abundance. Over the past ten years, there has been an effort to examine this radiation at different scales. In comparison with the Cambrian explosion which appears to be more globally mediated, local and regional studies of Ordovician faunas reveal sharp transitions with timing and magnitudes that vary geographically. These transitions suggest a more episodic and complex history than that revealed through synoptic global studies alone. Despite its apparent uniqueness, we cannot exclude the possibility that the Ordovician radiation was an extension of Cambrian diversity dynamics. That is, the Ordovician radiation may have been an event independent of the Cambrian radiation and thus requiring a different set of explanations, or it may have been the inevitable follow-up to the Cambrian radiation. Future studies should focus on resolving this issue.

Terminal Ordovician carbon isotope stratigraphy and glacioeustatic sea-level change across Anticosti Island (Québec, Canada)

Globally documented carbon isotope excursions provide time-varying signals that can be used for high-resolution stratigraphic correlation. We report detailed inorganic and organic carbon isotope curves from carbonate rocks of the Ellis Bay and Becscie Formations spanning the Ordovician-Silurian boundary on Anticosti Island, Quebec, Canada. Strata of the Anticosti Basin record the development of a storm-dominated tropical carbonate ramp. These strata host the well-known Hirnantian positive carbon isotope excursion, which attains maximum values of ~4.5‰ in carbonate carbon of the Laframboise Member or the Fox Point Member of the Becscie Formation. The excursion also occurs in organic carbon, and δ^(13)C_carb and δ^(13)C_org values covary such that no reproducible Δ^(13)C (= δ^(13)C_carb – δ^(13)C_org) excursion is observed. The most complete stratigraphic section, at Laframboise Point in the west, shows the characteristic shape of the Hirnantian Stage excursion at the global stratotype section and point (GSSP) for the Hirnantian Stage in China and the Silurian System in Scotland. We therefore suggest that the entire Hirnantian Stage on Anticosti Island is confined to the Laframboise and lower Fox Point Members. By documenting discontinuities in the architecture of the carbon isotope curve at multiple stratigraphic sections spanning a proximal to distal transect across the sedimentary basin, we are able to reconstruct glacioeustatic sea-level fluctuations corresponding to maximum glacial conditions associated with the end-Ordovician ice age. The combined litho- and chemostratigraphic approach provides evidence for the diachroneity of the oncolite bed and Becscie limestones; the former transgresses from west to east, and the latter progrades from east to west. The sea-level curve consistent with our sequence-stratigraphic model indicates that glacioeustatic sea-level changes and the positive carbon isotope excursion were not perfectly coupled. Although the start of the isotope excursion and the initial sea-level drawdown were coincident, the peak of the isotope excursion did not occur until after sea level had begun to rise. Carbon isotope values did not return to baseline until well after the Anticosti ramp was reflooded. The sea-level–δ13Ccarb relationship proposed here is consistent with the “weathering” hypothesis for the origin of the Hirnantian δ^(13)C_carb excursion.