Aaron O'Dea

Hopping Hotspots: Global Shifts in Marine Biodiversity

Hotspots of high species diversity are a prominent feature of modern global biodiversity patterns. Fossil and molecular evidence is starting to reveal the history of these hotspots. There have been at least three marine biodiversity hotspots during the past 50 million years. They have moved across almost half the globe, with their timing and locations coinciding with major tectonic events. The birth and death of successive hotspots highlights the link between environmental change and biodiversity patterns. The antiquity of the taxa in the modern Indo-Australian Archipelago hotspot emphasizes the role of pre-Pleistocene events in shaping modern diversity patterns.

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.

Environmental change preceded Caribbean extinction by 2 million years

Paleontologists typically treat major episodes of extinction as single and distinct events in which a major environmental perturbation results in a synchronous evolutionary response. Alternatively, the causes of biotic change may be multifaceted and extinction may lag behind the changes ultimately responsible because of nonlinear ecological dynamics. We examined these alternatives for the major episode of Caribbean extinction 2 million years ago (Ma). Isolation of the Caribbean from the Eastern Pacific by uplift of the Panamanian Isthmus was associated with synchronous changes in Caribbean near shore environments and community composition between 4.25 and 3.45 Ma. Seasonal fluctuations in Caribbean seawater temperature decreased 3-fold, carbonate deposition increased, and there was a striking, albeit patchy, shift in dominance of benthic ecosystems from heterotrophic mollusks to mixotrophic reef corals and calcareous algae. All of these changes correspond well with a simple model of decreased upwelling and collapse in planktonic productivity associated with the final stages of the closure of the isthmian barrier. However, extinction rates of mollusks and corals did not increase until 3–2 Ma and sharply peaked between 2 and 1 Ma, even though extinction overwhelmingly affected taxa commonly associated with high productivity. This time lag suggests that something other than environmental change per se was involved in extinction that does not occur as a single event. Understanding cause and effect will require more taxonomically refined analysis of the changing abundance and distribution patterns of different ecological guilds in the 2 million years leading up to the relatively sudden peak in extinction.

Historical biogeography of the Isthmus of Panama

About 3 million years ago (Ma), the Isthmus of Panama joined the Americas, forming a land bridge over which inhabitants of each America invaded the other—the Great American Biotic Interchange. These invasions transformed land ecosystems in South and Middle America. Humans invading from Asia over 12000 years ago killed most mammals over 44 kg, again transforming tropical American ecosystems. As a sea barrier, the isthmus induced divergent environmental change off its two coasts—creating contrasting ecosystems through differential extinction and diversification.

Intracolony variation in zooid size in cheilostome bryozoans as a new technique for investigating palaeoseasonality

Variation in zooid size within colonies of fossil cheilostome bryozoans is presented as a potential source of information on palaeoseasonality. We base our approach on the inverse relationship between temperature and zooid size in bryozoans, and analyse the mean intracolonial coeYcient of variation (CV ) in zooid length, zooid width and zooid area ( length◊width) in a number of Recent bryozoan species collected from many seasonally diVerent environments. A highly significant, positive correlation was obtained between the mean annual range of temperature (MART ) experienced by the colonies and the mean intracolonial CVs in zooid lengths (R2=74.7%), zooid widths (R2=58.9%) and zooid areas (R2=80.0%). An algebraic equation derived from regression analysis of mean intracolonial CV of zooid area and MART is proposed as a new method of investigating the MART of ancient seas by assessing variation in zooid area within fossil cheilostome colonies. This technique is then applied to bryozoan colonies from two Neogene shallow-water deposits in Western Europe. Results from the Coralline Crag in southeastern England reveal a moderate level of seasonality, in keeping with previous estimates of seasonality for British seas during the Pliocene. Results from the middle Miocene ‘faluns’ in north-west France suggest less seasonal variation in temperature than occurs in comparable seas today. We conclude that the technique represents a useful new approach that oVers some benefits over other techniques of assessing seasonality in marine palaeoenvironments.

Paleontological baselines for evaluating extinction risk in the modern oceans

Recognizing the threat of additive risk Humans are accelerating the extinction rates of species in both terrestrial and marine environments. However, species extinctions have occurred across time for a variety of other reasons. Finnegan et al. looked at the extinction rates across marine genera (groups of species) over the past 23 million years to determine intrinsic extinction rates and what traits or regions correspond to the highest rates. Combining patterns of intrinsic extinction with regions of high anthropogenic threat revealed taxa and areas, particularly in the tropics, where the risk of extinction will be especially high. Science, this issue p. 567 Fossils reveal patterns of extinction in marine species, past and present. Marine taxa are threatened by anthropogenic impacts, but knowledge of their extinction vulnerabilities is limited. The fossil record provides rich information on past extinctions that can help predict biotic responses. We show that over 23 million years, taxonomic membership and geographic range size consistently explain a large proportion of extinction risk variation in six major taxonomic groups. We assess intrinsic risk—extinction risk predicted by paleontologically calibrated models—for modern genera in these groups. Mapping the geographic distribution of these genera identifies coastal biogeographic provinces where fauna with high intrinsic risk are strongly affected by human activity or climate change. Such regions are disproportionately in the tropics, raising the possibility that these ecosystems may be particularly vulnerable to future extinctions. Intrinsic risk provides a prehuman baseline for considering current threats to marine biodiversity.

Environmental change drove macroevolution in cupuladriid bryozoans

Most macroevolutionary events are correlated with changes in the environment, but more rigorous evidence of cause and effect has been elusive. We compiled a 10 Myr record of origination and extinction, changes in mode of reproduction, morphologies and abundances of cupuladriid bryozoan species, spanning the time when primary productivity collapsed in the southwestern Caribbean as the Isthmus of Panama closed. The dominant mode of reproduction shifted dramatically from clonal to aclonal, due in part to a pulse of origination followed by extinction that was strongly selective in favour of aclonal species. Modern-day studies predict reduced clonality in increasingly oligotrophic conditions, thereby providing a mechanistic explanation supporting the hypothesis that the collapse in primary productivity was the cause of turnover. However, whereas originations were synchronous with changing environments, extinctions lagged 1–2 Myr. Extinct species failed to become more robust and reduce their rate of cloning when the new environmental conditions arose, and subsequently saw progressive reductions in abundance towards their delayed demise. Environmental change is therefore established as the root cause of macroevolutionary turnover despite the lag between origination and extinction.

Life history and environmental inference through retrospective morphometric analysis of bryozoans : a preliminary study

A preliminary comparative analysis of colony growth and zooid size in the perennial bryozoan Flustra foliacea (Bryozoa: Cheilostomatida) reveals reduced colony growth in the Bay of Fundy relative to growth in the Menai Straits and the Skagerrak, while seasonal fluctuations in zooid size are in synchrony with temperature regimes. Such retrospective morphometric analyses may allow inferences of primary productivity and thermal regimes and provide insights into the life histories of both Recent and fossil bryozoans.

Changes in bivalve functional and assemblage ecology in response to environmental change in the Caribbean Neogene

Abstract We documented changes in the relative abundance of bivalve genera and functional groups in the southwest Caribbean over the past 11 Myr to determine their response to oceanographic changes associated with the closure of the Central American Seaway ca. 3.5 Ma. Quantitative bulk samples from 29 localities yielded 106,000 specimens in 145 genera. All genera were assigned to functional groups based on diet, relationship to the substrate, and mobility. Ordinations of assemblages based on quantitative data for functional groups demonstrated strong shifts in community structure, with a stark contrast between assemblages older than 5 Ma and those younger than 3.5 Ma. These changes are primarily due to an increase in the abundance of attached epifaunal bivalves (e.g., Chama, Arcopsis, and Barbatia) and a decrease in infaunal bivalves (e.g., Varicorbula and Caryocorbula). Taxa associated with seagrasses, including deposit-feeding and chemosymbiotic bivalves (e.g., Lucina), also increased in relative abundance compared to suspension feeders. The composition of bivalve assemblages is correlated with the carbonate content of sediments and the percentage of skeletal biomass that is coral. Our results strongly support the hypothesis that increases in the extent of coral reefs and Thalassia communities were important drivers of biologic turnover in Neogene Caribbean benthic communities.

Evidence of size-selective evolution in the fighting conch from prehistoric subsistence harvesting.

Intensive size-selective harvesting can drive evolution of sexual maturity at smaller body size. Conversely, prehistoric, low-intensity subsistence harvesting is not considered an effective agent of size-selective evolution. Uniting archaeological, palaeontological and contemporary material, we show that size at sexual maturity in the edible conch Strombus pugilis declined significantly from pre-human (approx. 7 ka) to prehistoric times (approx. 1 ka) and again to the present day. Size at maturity also fell from early- to late-prehistoric periods, synchronous with an increase in harvesting intensity as other resources became depleted. A consequence of declining size at maturity is that early prehistoric harvesters would have received two-thirds more meat per conch than contemporary harvesters. After exploring the potential effects of selection biases, demographic shifts, environmental change and habitat alteration, these observations collectively implicate prehistoric subsistence harvesting as an agent of size-selective evolution with long-term detrimental consequences. We observe that contemporary populations that are protected from harvesting are slightly larger at maturity, suggesting that halting or even reversing thousands of years of size-selective evolution may be possible.