Sexual reproduction by scleractinian reef corals is important for maintaining coral populations and evolutionary processes. The ongoing global renaissance in coral reproduction research is providing a wealth of new information on this topic, and has almost doubled the global database on coral reproductive patterns during the past two decades. Information on sexual reproduction is now available for 444 scleractinian species, and confirms that hermaphroditic broadcast spawning is the dominant pattern among coral species studied to date. Relatively few hermaphroditic or gonochoric brooding species have been recorded. Multispecific coral spawning has been recorded on many reefs, but the degree of reproductive synchrony varies greatly within and among species at different geographic locations.

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Sexual Reproduction of Scleractinian Corals

With only about one-sixth of the original coral cover left, most Caribbean coral reefs may disappear in the next 20 years, primarily due to the loss of grazers in the region, according to this latest report, Status and Trends of Caribbean Coral Reefs: 1970-2012. This is the most detailed and comprehensive study of its kind published to date – the result of the work of 90 experts over the course of three years. It contains the analysis of more than 35,000 surveys conducted at 90 Caribbean locations since 1970, including studies of corals, seaweeds, grazing sea urchins and fish.

Status and trends of Caribbean coral reefs : 1970-2012

Global-scale deteriorations in coral reef health have caused major shifts in species composition. One projected consequence is a lowering of reef carbonate production rates, potentially impairing reef growth, compromising ecosystem functionality and ultimately leading to net reef erosion. Here, using measures of gross and net carbonate production and erosion from 19 Caribbean reefs, we show that contemporary carbonate production rates are now substantially below historical (mid- to late-Holocene) values. On average, current production rates are reduced by at least 50%, and 37% of surveyed sites were net erosional. Calculated accretion rates (mm year � 1 ) for shallow fore-reef habitats are also close to an order of magnitude lower than Holocene averages. A live coral cover threshold of B10% appears critical to maintaining positive production states. Below this ecological threshold carbonate budgets typically become net negative and threaten reef accretion. Collectively, these data suggest that recent ecological declines are now suppressing Caribbean reef growth potential.

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Caribbean-wide decline in carbonate production threatens coral reef growth

Trait-based approaches advance ecological and evolutionary research because traits provide a strong link to an organism’s function and fitness. Trait-based research might lead to a deeper understanding of the functions of, and services provided by, ecosystems, thereby improving management, which is vital in the current era of rapid environmental change. Coral reef scientists have long collected trait data for corals; however, these are difficult to access and often under-utilized in addressing large-scale questions. We present the Coral Trait Database initiative that aims to bring together physiological, morphological, ecological, phylogenetic and biogeographic trait information into a single repository. The database houses species- and individual-level data from published field and experimental studies alongside contextual data that provide important framing for analyses. In this data descriptor, we release data for 56 traits for 1547 species, and present a collaborative platform on which other trait data are being actively federated. Our overall goal is for the Coral Trait Database to become an open-source, community-led data clearinghouse that accelerates coral reef research.

The Coral Trait Database, a curated database of trait information for coral species from the global oceans

Dispersal influences ecological dynamics, evolution, biogeography, and biodiversity conservation, but models of larval dispersal in marine organisms make simplifying assumptions that are likely to approximate poorly the temporal dynamics of larval survival and capacity for settlement. In particular, larval mortality rates are typically assumed to be constant throughout larval life; and all larvae are frequently assumed to acquire and lose competence at the same time. To improve upon these assumptions, we here develop simple models of dispersal potential that incorporate rates of mortality, and acquisition and loss of settlement competence. We fit these models to empirical competence and survival data for five scleractinian coral species, to test the models' ability to characterize empirical survival and competence patterns, and to estimate the dispersal potential implied by those patterns. The models fit the data well, incorporating qualitative features of competence and survival that traditional approaches to modeling dispersal do not, with important implications for dispersal potential. Most notably, there was high within-cohort variation in the duration of the competent period in all species, and this variation increases both self-recruitment and long-distance dispersal compared with models assuming a fixed competent period. These findings help to explain the seeming paradox of high genetic population structure, coupled with large geographic range size, observed in many coral species. More broadly, our approach offers a way to parsimoniously account for variation in competence dynamics in dispersal models, a phenomenon that our results suggest has important effects on patterns of connectivity in marine metapopulations.

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Estimating dispersal potential for marine larvae: dynamic models applied to scleractinian corals.

Due to the importance of preserving the genetic integrity of populations, strategies to restore damaged coral reefs should attempt to retain the allelic diversity of the disturbed population; however, genetic diversity estimates are not available for most coral populations. To provide a generalized estimate of genetic diversity (in terms of allelic richness) of scleractinian coral populations, the literature was surveyed for studies describing the genetic structure of coral populations using microsatellites. The mean number of alleles per locus across 72 surveyed scleractinian coral populations was 8.27 (±0.75 SE). In addition, population genetic datasets from four species (Acropora palmata, Montastraea cavernosa, Montastraea faveolata and Pocillopora damicornis) were analyzed to assess the minimum number of donor colonies required to retain specific proportions of the genetic diversity of the population. Rarefaction analysis of the population genetic datasets indicated that using 10 donor colonies randomly sampled from the original population would retain >50% of the allelic diversity, while 35 colonies would retain >90% of the original diversity. In general, scleractinian coral populations are genetically diverse and restoration methods utilizing few clonal genotypes to re-populate a reef will diminish the genetic integrity of the population. Coral restoration strategies using 10–35 randomly selected local donor colonies will retain at least 50–90% of the genetic diversity of the original population.

Restoration of coral populations in light of genetic diversity estimates

The rapid decline of Acropora cervicornis and Acropora palmata has often been linked with coral reef deterioration in the Caribbean; yet, it remains controversial whether these species are currently recovering or still declining. In this study, the status of ten populations of A. palmata in Los Roques National Park (LRNP), Venezuela is presented. Six of these populations showed signs of recovery. Ten 80 m2 belt-transects were surveyed at each of the ten reef sites. Within belt-transects, each colony was measured (maximum diameter and height) and its status (healthy, diseased or injured) was recorded. Populations in recovery were defined by a dominance of small to medium-sized colonies in densities >1 colony per 10 m2, together with 75% undamaged colonies, a low prevalence of diseases (<10%), and a low density of predators (0.25 snails per colony). Based on allozyme analysis of seven polymorphic loci in four populations (N = 30), a moderate to high-genetic connectivity among these populations (F
 ST = 0.048) was found with a predominance of sexual over asexual reproduction (N* : N = 1; N
 go : N = 0.93–1). Both ecological and molecular data support a good prognosis for the recovery of this species in Los Roques.

Ecological and genetic data indicate recovery of the endangered coral Acropora palmata in Los Roques, Southern Caribbean

This study aimed to evaluate potential differences in coral spawning behavior between a fringing coastal reef and an offshore reef in the southern Caribbean. For this, scleractinian and gorgonian colonies (N = 324) of 21 species were mapped along eight transects, each 10-m long, at two study sites located in Morrocoy and Los Roques National Parks, Venezuela. Observations were made between 19:30 and 23:00 from August 23 to 30 and from September 26 to 30, 2002. Ninety one colonies belonging to six hard coral and seven octocoral species spawned or planulated during this period. We were unable to observe any signs of reproductive activity in 95 colonies of nine species different from those that reproduced. Despite the differences in environmental conditions between the two sites, we observed high synchrony in the spawning behavior of seven coral species common to both reefs. The most striking result was the ability of colonies of Montastraea faveolata and Eusmilia fastigiata to split spawn up to three times, either in consecutive nights or in different months.

Coral mass- and split-spawning at a coastal and an offshore Venezuelan reefs, southern Caribbean

Studies of genetic diversity and population genetic structure in marine organisms are relevant to understanding populations’ variability, and therefore their ability to withstand environmental perturbations, their potential for resistance to local extinction and their natural rate of recovery. Population structure and genetic diversity were assessed at a regional spatial scale (i.e., Mesoamerican Barrier Reef System, MBRS) in two major reef building coral species Orbicella (formerly Montastraea) faveolata and Acropora palmata, and at a larger spatial scale (i.e., Caribbean-wide; MBRS, Panama, Venezuela and Puerto Rico) for A. palmata only. The most significant findings were as follows: (1) high genetic diversity and low clonality were found for both species, which is expected for O. faveolata but not for A. palmata, (2) both species showed low-to-moderate, yet significant population structure among populations along the MBRS; in particular, O. faveolata and A. palmata from Ambergris (Belize) and O. faveolata from Calabash (Belize) and A. palmata from Puerto Morelos (Mexico) showed some genetic differentiation from the rest of the MBRS populations, and (3) A. palmata from MBRS, Panama, Puerto Rico and Venezuela were grouped into four subregions that could be considered as management units. A more spatially detailed sampling program and the inclusion of recruits will be necessary to get a comprehensive understanding of coral population structure and current gene flow patterns in these two species.

Population structure of the corals Orbicella faveolata and Acropora palmata in the Mesoamerican Barrier Reef System with comparisons over Caribbean basin-wide spatial scale

Eight years ago (2007), the distribution and status of Acropora palmata was quantified throughout Los Roques archipelago in Venezuela. The aim was to produce a baseline study for this species which combined population genetics with demographic data. The results highlighted that A. palmata had the potential to recover in at least 6 out of 10 sites surveyed. Recovery potential was assumed to be high at sites with a relatively high abundance of the coral, low disease prevalence, high genetic diversity, and high rates of sexual reproduction. However, as noted, Zubillaga et al. (2008) realized recovery was still strongly dependent on local and regional stressors. In 2014 (this study), the status of A. palmata was re-evaluated at Los Roques. We increased the number of sites from 10 in the original baseline study to 106. This allowed us to assess the population status throughout the entirety of the MPA. Furthermore, we also identified local threats that may have hindered population recovery. Here, we show that A. palmata now has a relatively restricted distribution throughout the park, only occurring in 15% of the sites surveyed. Large stands of old dead colonies were common throughout the archipelago; a result which demonstrates that this species has lost almost 50% of its original distribution over the past decades. The majority of corals recorded were large adults (∼2 m height), suggesting that these older colonies might be less susceptible or more resilient to local and global threats. However, 45% of these surviving colonies showed evidence of partial mortality and degradation of living tissues. Interestingly, the greatest increase in partial mortality occurred at sites with the lowest levels of protection ([Formula: see text]; df = 4, p < 0.05). This may suggest there is a positive role of small scale marine management in assisting reef recovery. We also recorded a significant reduction ([Formula: see text]; df = 8; p < 0.05) in the density of A. palmata in sites that had previously been categorized as having a high potential for recovery. One explanation for this continued decline may be due to the fact that over the past 10 years, two massive bleaching events have occurred throughout the Caribbean with records showing that Los Roques has experienced unprecedented declines in overall coral cover. We therefore conclude that although local protection could promote recovery, the impacts from global threats such as ocean warming may hamper the recovery of this threatened species.

Ainhoa L. Zubillaga

Papers

Restoration of coral populations in light of genetic diversity estimates

Ecological and genetic data indicate recovery of the endangered coral Acropora palmata in Los Roques, Southern Caribbean

Coral mass- and split-spawning at a coastal and an offshore Venezuelan reefs, southern Caribbean

Population structure of the corals Orbicella faveolata and Acropora palmata in the Mesoamerican Barrier Reef System with comparisons over Caribbean basin-wide spatial scale

Is Acropora palmata recovering? A case study in Los Roques National Park, Venezuela