Defining variation in pre-human ecosystems can guide conservation: An example from a Caribbean coral reef.

Many Caribbean coral reefs are heavily degraded, yet their pre-human, natural states are often assumed or estimated using space-for-time substitution approaches. Here we use an 11-hectare suite of fossilised mid-Holocene (7.2-5.6 ka) fringing reefs in Caribbean Panama to define natural variation in hard coral community structure before human-impact to provide context to the states of the same reefs today. We collected bulk samples from four trenches dug into the mid-Holocene fossil reef and surficial bulk samples from 2-10 m depths on five adjacent modern reefs extending over 5 km. Analysis of the abundances of coral taxa in fossil bulk samples define the Historical Range of Variation (HRV) in community structure of the reefs. When compared to the community structure of adjacent modern reefs, we find that most coral communities today fall outside the HRV, identifying them as novel ecosystems and corroborating the well-documented transition from acroporid-dominated Caribbean reefs to reefs dominated by stress-tolerant taxa (Porites and Agaricia). We find one modern reef, however, whose community composition remains within the HRV showing that it has not transitioned to a novel state. Reef-matrix cores extracted from this reef reveal that the coral community has remained in this state for over 800 years, suggesting long-term stability and resistance to the region-wide shift to novel states. Without these data to provide historical context, this potentially robust and stable reef would be overlooked since it does not fulfil expectations of what a Caribbean coral reef should look like in the absence of humans. This example illustrates how defining past variation using the fossil record can improve our understanding of modern degradation and guide conservation.

Species differences drive nonneutral structure in pleistocene coral communities.

Although ecological assemblages frequently depart from neutral model predictions, these discrepancies have not been unambiguously attributed to neutral theory's core assumption: that community structure is primarily the result of chance variation in birth, death, speciation, and dispersal, rather than the manifestation of demographic differences among species. Using coral communities in Barbados from four time periods during the Pleistocene, we demonstrate that the neutral theory cannot explain coral community similarity distributions, species' regional abundance distributions, or their local occupancy. Furthermore, discrepancies between the neutral theory and the observed communities can be attributed to violation of the core assumption of species equivalence. In particular, species' variable growth rates are driving departures from neutral predictions. Our results reinforce an understanding of reef coral community assembly that invokes trade-offs in species' demographic strategies. The results further suggest that conservation management actions will fail if they are based on the neutral assumption that different coral species are equally able to create live coral cover in the shallow-water reef environment. These findings highlight the importance of developing biodiversity theory that can parsimoniously incorporate species differences in coral reef communities, rather than further elaborating neutral theory.

Evolutionary novelty is concentrated at the edge of coral species distributions.

Conservation priorities are calculated on the basis of species richness, endemism, and threats. However, areas ranked highly for these factors may not represent regions of maximal evolutionary potential. The relationship between geography and evolutionary innovation was analyzed in a dominant complex of Caribbean reef corals, in which morphological and genetic data concur on species differences. Based on geometric morphometrics of Pleistocene corals and genetically characterized modern colonies, we found that morphological disparity varies from the center to the edge of the Caribbean, and we show that lineages are static at well-connected central locations but split or fuse in edge zones where gene flow is limited. Thus, conservation efforts in corals should focus not only on the centers of diversity but also on peripheral areas of species ranges and population connectivity.

Ecological persistence interrupted in Caribbean coral reefs.

The recent mass mortality of Caribbean reef corals dramatically altered reef community structure and begs the question of the past stability and persistence of coral assemblages before human disturbance began. We report within habitat stability in coral community composition in the Pleistocene fossil record of Barbados for at least 95,000 years despite marked variability in global sea level and climate. Results were consistent for surveys of both common and rare taxa. Comparison of Pleistocene and modern community structure shows that Recent human impacts have changed coral community structure in ways not observed in the preceding 220,000 years.