Katie L. Cramer

Anthropogenic mortality on coral reefs in Caribbean Panama predates coral disease and bleaching

Caribbean reef corals have declined precipitously since the 1980s due to regional episodes of bleaching, disease and algal overgrowth, but the extent of earlier degradation due to localised historical disturbances such as land clearing and overfishing remains unresolved. We analysed coral and molluscan fossil assemblages from reefs near Bocas del Toro, Panama to construct a timeline of ecological change from the 19th century-present. We report large changes before 1960 in coastal lagoons coincident with extensive deforestation, and after 1960 on offshore reefs. Striking changes include the demise of previously dominant staghorn coral Acropora cervicornis and oyster Dendrostrea frons that lives attached to gorgonians and staghorn corals. Reductions in bivalve size and simplification of gastropod trophic structure further implicate increasing environmental stress on reefs. Our paleoecological data strongly support the hypothesis, from extensive qualitative data, that Caribbean reef degradation predates coral bleaching and disease outbreaks linked to anthropogenic climate change.

Defining and dividing the greater Caribbean: insights from the biogeography of shorefishes.

The Greater Caribbean biogeographic region is the high-diversity heart of the Tropical West Atlantic, one of four global centers of tropical marine biodiversity. The traditional view of the Greater Caribbean is that it is limited to the Caribbean, West Indies, southwest Gulf of Mexico and tip of Florida, and that, due to its faunal homogeneity, lacks major provincial subdivisions. In this scenario the northern 2/3 of the Gulf of Mexico and southeastern USA represent a separate temperate, “Carolinian” biogeographic region. We completed a comprehensive re-assessment of the biogeography of the Greater Caribbean by comparing the distributions of 1,559 shorefish species within 45 sections of shelf waters of the Greater Caribbean and adjacent areas. This analysis shows that that the Greater Caribbean occupies a much larger area than usually thought, extending south to at least Guyana, and north to encompass the entire Carolinian area. Rather than being homogenous, the Greater Caribbean is divided into three major provinces, each with a distinctive, primarily tropical fauna: (1) a central, tropical province comprising the West Indies, Bermuda and Central America; (2) a southern, upwelling-affected province spanning the entire continental shelf of northern South America; and (iii) a northern, subtropical province that includes all of the Gulf of Mexico, Florida and southeastern USA. This three-province pattern holds for both reef- and soft bottom fishes, indicating a general response by demersal fishes to major variation in provincial shelf environments. Such environmental differences include latitudinal variation in sea temperature, availability of major habitats (coral reefs, soft bottom shorelines, and mangroves), and nutrient additions from upwelling areas and large rivers. The three-province arrangement of the Greater Caribbean broadly resembles and has a similar environmental basis to the provincial arrangement of its sister biogeographic region, the Tropical Eastern Pacific.

Molluscan subfossil assemblages reveal the long-term deterioration of coral reef environments in Caribbean Panama

Caribbean reef corals have declined sharply since the 1980s, but the lack of prior baseline data has hindered identification of drivers of change. To assess anthropogenic change in reef environments over the past century, we tracked the composition of subfossil assemblages of bivalve and gastropod mollusks excavated from pits below lagoonal and offshore reefs in Bocas del Toro, Panama. The higher prevalence of (a) infaunal suspension-feeding bivalves and herbivorous and omnivorous gastropods in lagoons and (b) epifaunal and suspension-feeding bivalves and carnivorous and suspension-feeding gastropods offshore reflected the greater influence of land-based nutrients/sediments within lagoons. Temporal changes indicated deteriorating environmental conditions pre-1960 in lagoons and post-1960 offshore, with offshore communities becoming more similar to lagoonal ones since 1960. Relative abundances of dominant bivalve species tracked those of their coral hosts, revealing broader ecosystem effects of coral community change. The nature and timing of changes implicate land-based runoff in reef deterioration.

Prehistorical and historical declines in Caribbean coral reef accretion rates driven by loss of parrotfish

Caribbean coral reefs have transformed into algal-dominated habitats over recent decades, but the mechanisms of change are unresolved due to a lack of quantitative ecological data before large-scale human impacts. To understand the role of reduced herbivory in recent coral declines, we produce a high-resolution 3,000 year record of reef accretion rate and herbivore (parrotfish and urchin) abundance from the analysis of sediments and fish, coral and urchin subfossils within cores from Caribbean Panama. At each site, declines in accretion rates and parrotfish abundance were initiated in the prehistorical or historical period. Statistical tests of direct cause and effect relationships using convergent cross mapping reveal that accretion rates are driven by parrotfish abundance (but not vice versa) but are not affected by total urchin abundance. These results confirm the critical role of parrotfish in maintaining coral-dominated reef habitat and the urgent need for restoration of parrotfish populations to enable reef persistence.

A 3000 year record of Caribbean reef urchin communities reveals causes and consequences of long‐term decline in Diadema antillarum

Urchins are the last abundant grazers of macroalgae on most Caribbean reefs following the historical overexploitation of herbivorous fishes. The long-spined urchin Diadema antillarum was particularly effective at controlling macroalgae and facilitating coral dominance on Caribbean reefs until its ecological extinction from a catastrophic disease epidemic in the early 1980s. Despite their important role in the structure and functioning of Caribbean reef ecosystems, the natural dynamics of Caribbean reef urchin communities are poorly known due to the paucity of ecological survey data prior to large-scale human disturbances and the Diadema dieoff. To help resolve the baseline abundances and ecological roles of common urchin taxa, we track changes in urchin abundance and composition over the past 3000 yr from analysis of subfossil urchin spines preserved in reef matrix cores collected in Caribbean Panama. Echinometra consistently dominated the subfossil spine assemblage, while Diadema was consistently rare in the subfossil record in this region. Rather than increasing during a period of heightened human exploitation of their fish competitors and predators, Diadema began declining over a millennium ago. Convergent cross mapping (CCM) causality analyses reveal that Diadema abundance is causally related to coral community composition. Diadema is negatively affected by Acropora cervicornis dominance, likely due to the tight association between this coral and the threespot damselfish, an effective Diadema competitor. Conversely, Diadema positively affects the abundance of the coral Madracis mirabilis, possibly via its control of macroalgae. Causal relationships were not detected among abundances of individual urchin taxa, indicating that inter-specific echinoid competition is not a factor limiting Diadema recovery. Our detailed record of prehistorical and historical urchin community dynamics suggests that the failure of Diadema to recover over 30 yr after its mass mortality event may be due in part to the prey release of damselfish following the long-term overfishing of piscivorous fishes.

Methods for isolation and quantification of microfossil fish teeth and elasmobranch dermal denticles (ichthyoliths) from marine sediments

Ichthyoliths—microfossil fish teeth and shark dermal scales (denticles)—are found in nearly all marine sediments. Their small size and relative rarity compared to other microfossil groups means that they have been largely ignored by the paleontology and paleoceanography communities, except as carriers of certain isotopic systems. Yet, when properly concentrated, ichthyoliths are sufficiently abundant to reveal patterns of fish abundance and diversity at unprecedented temporal and spatial resolution, in contrast to the typical millions of years-long gaps in the vertebrate body fossil record. In addition, ichthyoliths are highly resistant to dissolution, making it possible to reconstruct whole fish communities over highly precise and virtually continuous timescales. Here we present methods to isolate and utilize ichthyoliths preserved in the sedimentary record to track fish community structure and ecosystem productivity through geological and historical time periods. These include techniques for isolation and concentration of these microfossils from a wide range of sediments, including deep-sea and coral reef carbonates, clays, shales, and silicate-rich sediments. We also present a novel protocol for ichthyolith staining using Alizarin Red S to easily visualize and distinguish small teeth from debris in the sample. Finally, we discuss several metrics for quantification of ichthyolith community structure and abundance, and their applications to reconstruction of ancient marine food webs and environments. Elizabeth C. Sibert. Society of Fellows, Harvard University, 78 Mount Auburn Street, Cambridge, Massachusetts 02138, USA; Scripps Institution of Oceanography, University of California, San Diego, 9500 Gilman Drive MC 0208, La Jolla, California 92093, USA. esibert@fas.harvard.edu Katie L. Cramer. Scripps Institution of Oceanography, University of California, San Diego, 9500 Gilman Drive MC 0208, La Jolla, California 92093, USA. kcramer@ucsd.edu Philip A. Hastings. Scripps Institution of Oceanography, University of California, San Diego, 9500 Gilman Drive MC 0208, La Jolla, California 92093, USA. phastings@ucsd.edu Richard D. Norris. Scripps Institution of Oceanography, University of California, San Diego, 9500 Gilman Drive MC 0244, La Jolla, California 92093, USA. rnorris@ucsd.edu