Reef Of Moorea Research Articles

Non-random orientation of Pocillopora colonies on forereefs of Moorea, French Polynesia

Morphological variation in scleractinian corals has been variously ascribed to genetic differentiation and phenotypic plasticity, with a likely influence of both factors. Experimental approaches have dominated studies of phenotypic plasticity, for example with asymmetrical colony formation resulting from exposure to experimentally imposed unidirectional water flow. However, it has been difficult to disentangle the effects of competition for space between sessile organisms from the influence of physical forcing on coral morphologies in situ. To this end, we investigated morphological variation of coral colonies of the genus Pocillopora on reefs in Moorea, French Polynesia, where recovery (by 2018) after near complete loss of coral cover on the forereef in 2010 was driven predominantly by recruitment and growth of Pocillopora. We employed a large-area imaging approach at 8 study sites in the forereef of Moorea, constructing digital models of 100 m2 benthic plots to describe the orientation of ~400 individual colonies of Pocillopora. Results demonstrate that Pocillopora colonies on the forereef of Moorea exhibit non-random orientations, with a systematic orientation of the major axis that lies approximately perpendicular to the 20-30 m isobath.

Structural complexity shapes the behavior and abundance of a common herbivorous fish, increasing herbivory on a turf-dominated, fringing reef

Coral-dominated reefs typically have abundant refuges and limited resources for herbivores; conversely, algal-dominated reefs have abundant resources but often have limited refuges. This difference results in a trade-off between risk and reward for herbivorous fishes. However, disturbances that kill coral but retain topographic complexity can decouple this tradeoff, challenging this paradigm. Many fringing reefs in Moorea, French Polynesia are algal-dominated with heterogeneous structural complexity likely driven by disturbances associated with crown-of-thorns predation and typhoons, which occurred between 2006 and 2010. Short turfs are a good recruitment surface for coral larvae, and thus has a greater potential to recover to coral dominance than long turfs. To explore how variation in complexity impacts the behavior of herbivorous fishes, we compared abundance, residence time, and time budgets of Acanthurus nigrofuscus, a common herbivore species, in paired high versus low complexity areas. We chose areas in close proximity to maximize the likelihood fish could choose between complexity types. We also measured turf height in high versus low complexity plots. Fish were ~3× more abundant, spent roughly double the time in, and budgeted more than 4× as much time to hiding in high complexity reef areas. Although, on average, individual herbivores spent the same proportion of time foraging in both areas, their greater abundance and longer residence time suggests their local herbivory rates could be 7.5 times higher in high complexity areas. We also found turfs in high complexity plots were roughly 75% the height of turfs in low complexity plots. This provides some evidence for stronger overall herbivory pressure in high complexity areas, albeit differences in absolute turf height were modest (~0.5 mm). Combined, these results suggest reefs that retain structural complexity following disturbance may be more likely to recover due to the higher localized herbivory rates resulting from uncoupling of the tradeoff between structural complexity and resource limitation.

Selective consumption of macroalgal species by herbivorous fishes suggests reduced functional complementarity on a fringing reef in Moorea, French Polynesia

Worldwide, many coral reefs are at risk of shifting to degraded algal-dominated states, due to compromised ecological conditions. Functional diversity of herbivorous fishes maintains coral reef health and promotes reef resilience to disturbances. Given previous evidence, it appears the functional roles of herbivorous fishes differ across geographical locations, indicating a need for further assessment of macroalgal consumption by herbivorous fishes. We assessed functional diversity by examining foraging behavior of herbivorous fish species on macroalgae on a fringing reef in Moorea, French Polynesia. We video-recorded choice experiments containing seven common macroalgae and used Strauss' linear resource selection index to determine macroalgal selectivities. We used cluster analysis to identify any distinct groups within herbivorous fish species, given the macroalgal species they targeted, and fitted generalized linear mixed-effects models to identify factors that best predicted the number of bites taken on macroalgae. Seven species from 3 fish families/tribes took a total of 956 bites. Fish species differed in their selectivity with some species (Naso lituratus, N. unicornis, Calatomus carolinus) strongly preferring one or two macroalgal species, while other fish species (Acanthurus nigrofuscus, Ctenochaetus striatus, Chlorurus sordidus, Balistapus undulatus) were less selective. This resulted in fish species forming two clusters. Only 3 of 7 macroalgae were preferred by any fish species, with two fish species both preferring the same two macroalgae. The limited differences in fish species' preferences for different macroalgae suggests limited functional complementarity. Two models (macroalgal species identity+fish functional group, macroalgal species identity+fish species) best predicted the number of bites taken on macroalgae compared to models incorporating only a single explanatory factor or fish family. In the context of this Moorean fringing reef, there is greater functional redundancy than complementarity of herbivorous fishes consuming macroalgae, and the fishes grouped together according to their relative selectivity. We observed fish species that are not classified as browsers consuming macroalgae, suggesting diets of herbivorous fishes may be broader than previously thought. Finally, we observed macroalgal selectivities and consumption that differed from previous studies for the same fish species. Our results contribute to the understanding of functional diversity of herbivorous fishes across coral reefs, and also highlight the need for additional research to further elucidate the role of context and functional diversity of herbivorous fishes consuming macroalgae on coral reefs.

Herbivory as a limiting factor for seagrass proximity to fringing reefs in Moorea, French Polynesia

Abstract Seagrasses are important foundational species that frequently display distinct depth distributions, although the drivers of these patterns can be spatially and temporally variable. While the pantropical seagrass Halophila decipiens is known from waters as shallow as 1 m deep, in Moorea, French Polynesia we only found it > 6.4 m deep. To explore factors affecting depth distribution, we transplanted H. decipiens into 3 habitats: the existing seagrass bed (control), just outside the seagrass bed, and shallower habitat adjacent to a fringing coral reef. Results showed that growth was not significantly different between the control and just outside of the seagrass bed; however, number of shoots and rhizome length were significantly reduced adjacent to the reef. Transplanting seagrass into the shallow reef site with and without herbivore exclusion cages showed that H. decipiens grew in herbivore exclusion treatments, but lost both shoots and rhizomes in the control. These results indicate that H. decipiens can grow in shallow habitats adjacent to reefs on Moorea, but that herbivory pressure, presumably from the reef, limits its depth distribution.

Epifaunal invertebrate assemblages associated with branching Pocilloporids in Moorea, French Polynesia.

Reef-building corals can harbour high abundances of diverse invertebrate epifauna. Coral characteristics and environmental conditions are important drivers of community structure of coral-associated invertebrates; however, our current understanding of drivers of epifaunal distributions is still unclear. This study tests the relative importance of the physical environment (current flow speed) and host quality (e.g., colony height, surface area, distance between branches, penetration depth among branches, and background partial mortality) in structuring epifaunal communities living within branching Pocillopora colonies on a back reef in Moorea, French Polynesia. A total of 470 individuals belonging to four phyla, 16 families and 39 genera were extracted from 36 Pocillopora spp. colonies. Decapods were the most abundant epifaunal organisms (accounting for 84% of individuals) found living in Pocillopora spp. While coral host characteristics and flow regime are very important, these parameters were not correlated with epifaunal assemblages at the time of the study. Epifaunal assemblages associated with Pocillopora spp. were consistent and minimally affected by differences in host characteristics and flow regime. The consistency in abundance and taxon richness among colonies (regardless of habitat characteristics) highlighted the importance of total habitat availability. With escalating effects of climate change and other localized disturbances, it is critical to preserve branching corals to support epifaunal communities.

Complex and interactive effects of ocean acidification and temperature on epilithic and endolithic coral-reef turf algal assemblages

Turf algal assemblages are ubiquitous primary producers on coral reefs, but little is known about the response of this diverse group to ocean acidification (OA) across different temperatures. We tested the hypothesis that CO2 influences the functional response of epilithic and endolithic turf assemblages to increasing temperature. Replicate carbonate plugs covered by turf were collected from the reef and exposed to ambient and high pCO2 (1000 µatm) conditions for 3 weeks. Each pCO2 treatment was replicated across six temperatures (24.0–31.5 °C) that spanned the full seasonal temperature range on a fringing reef in Moorea, French Polynesia, and included one warming treatment (3 °C above daily average temperatures). Temperature and CO2 enrichment had complex, and sometimes interactive, effects on turf metabolism and growth. Photosynthetic and respiration rates were enhanced by increasing temperature, with an interactive effect of CO2 enrichment. Photosynthetic rates were amplified by high CO2 in the warmest temperatures, while the increase in respiration rates with temperature were enhanced under ambient CO2. Epilithic turf growth rates were not affected by temperature, but increased in response to CO2 enrichment. We found that CO2 and temperature interactively affected the endolithic assemblage, with the highest growth rates under CO2 enrichment, but only at the warmest temperatures. These results demonstrate how OA may influence algal physiology and growth across a range of ecologically relevant temperatures, and indicate that the effects of CO2 enrichment on coral-reef turf assemblages can be temperature dependent. The complex effects of CO2 enrichment and temperature across a suite of algal responses illustrates the importance of incorporating multiple stressors into global change experiments.

The potential for self-seeding by the coral Pocillopora spp. in Moorea, French Polynesia.

Coral reefs in Moorea, French Polynesia, suffered catastrophic coral mortality through predation by Acanthaster planci from 2006 to 2010, and Cyclone Oli in 2010, yet by 2015 some coral populations were approaching pre-disturbance sizes. Using long-term study plots, we quantified population dynamics of spawning Pocillopora spp. along the north shore of Moorea between 2010 and 2014, and considered evidence that population recovery could be supported by self-seeding. Results scaled up from study plots and settlement tiles suggest that the number of Pocillopora spp. colonies on the outer reef increased 1,890-fold between 2010 and 2014/2015, and in the back reef, 8-fold between 2010 and 2014/2015. Assuming that spawning Pocillopora spp. in Moorea release similar numbers of eggs as con-generics in Hawaii, and fertilization success is similar to other spawning corals, the capacity of Pocillopora spp. to produce larvae was estimated. These estimates suggest that Pocillopora spp. in Moorea produced a large excess of larvae in 2010 and 2014 relative to the number required to produce the recruits found in the back reef and outer reef in 2010 and 2014, even assuming that ∼99.9% of the larvae do not recruit in Moorea. Less than a third of the recruits in one year would have to survive to produce the juvenile Pocillopora spp. found in the back and outer reefs in 2010 and 2014/2015. Our first order approximations reveal the potential for Pocillopora spp. on the north shore of Moorea to produce enough larvae to support local recruitment and population recovery following a catastrophic disturbance.

Coral Reef Resilience, Tipping Points and the Strength of Herbivory.

Coral reefs increasingly are undergoing transitions from coral to macroalgal dominance. Although the functional roles of reef herbivores in controlling algae are becoming better understood, identifying possible tipping points in the herbivory-macroalgae relationships has remained a challenge. Assessment of where any coral reef ecosystem lies in relation to the coral-to-macroalgae tipping point is fundamental to understanding resilience properties, forecasting state shifts, and developing effective management practices. We conducted a multi-year field experiment in Moorea, French Polynesia to estimate these properties. While we found a sharp herbivory threshold where macroalgae escape control, ambient levels of herbivory by reef fishes were well above that needed to prevent proliferation of macroalgae. These findings are consistent with previously observed high resilience of the fore reef in Moorea. Our approach can identify vulnerable coral reef systems in urgent need of management action to both forestall shifts to macroalgae and preserve properties essential for resilience.

Spatial patterns of self-recruitment of a coral reef fish in relation to island-scale retention mechanisms.

Oceanographic features influence the transport and delivery of marine larvae, and physical retention mechanisms, such as eddies, can enhance self-recruitment (i.e. the return of larvae to their natal population). Knowledge of exact locations of hatching (origin) and settlement (arrival) of larvae of reef animals provides a means to compare observed patterns of self-recruitment 'connectivity' with those expected from water circulation patterns. Using parentage inference based on multiple sampling years in Moorea, French Polynesia, we describe spatial and temporal variation in self-recruitment of the anemonefish Amphiprion chrysopterus, evaluate the consistency of net dispersal distances of self-recruits against the null expectation of passive particle dispersal and test the hypothesis that larvae originating in certain reef habitats (lagoons and passes) would be retained and thus more likely to self-recruit than those originating on the outer (fore) reef. Estimates of known self-recruitment were consistent across the sampling years (~25-27% of sampled recruits). For most (88%) of these self-recruits, the net distance between hatching and settlement locations was within the maximum dispersal distance expected for a neutrally buoyant passive particle based on the longest duration of the larval dispersive phase and the average direction and speed of current flow around Moorea. Furthermore, a parent of a given body size on the outer (fore) reef of Moorea was less likely to produce self-recruits than those in passes. Our findings show that even a simple dispersal model based on net average flow and direction of alongshore currents can provide insight into landscape-scale retention patterns of reef fishes.

Size matters: experimental partitioning of the strength of fish herbivory on a fringing coral reef in Moorea, French Polynesia

AbstractWhile the importance of top‐down control by grazers in maintaining tropical reef community structure is well known, the effect of ‘fishing down the food web’, which simultaneously changes the abundance and size of herbivorous fishes, has received less attention. As many fishing practices target the biggest fish and regulations often set minimum size limits, understanding size‐dependency of this controlling force is important. We evaluated the hypothesis that reduction in the abundance and size of fish, regardless of species identity, reduces the role of herbivory in controlling algal abundance on reefs and assessed variation in efficacy of this top‐down control on different types of common macroalgae. We quantified herbivory rates within cages of differing opening sizes on assemblages of four common algal species (Padina boryana, Dictyota bartayresiana, Halimeda opuntia and Galaxaura fasciculata) on two fringing reefs in Moorea, French Polynesia. Small acanthurids (<15 cm length) were the dominant herbivorous fish while other herbivorous fish were rare. For the two most palatable algae, the majority of herbivory occurred in open plots, with herbivory reduced by >50% in the largest opening cages (6 × 6 cm) where the maximum size fish entering was 12 cm in length (mean = 7.6, SE = 0.4). Fish entering medium (4.5 × 4.5 cm openings, maximum fish length = 8 cm, mean = 6.3, SE = 0.4) and small (3 × 3 cm openings, no fish observed entering) cages had herbivory rates approximately equal to the control treatment (1 × 1 cm openings). Consumption varied among algal species, with minimal consumption of physically and chemically defended algae and no pattern across treatments. Our results demonstrate a need for management plans to not only maintain the overall abundance of herbivorous fish but to protect the largest sizes for effective top‐down control of algal communities.

Complex environmental forcing across the biogeographical range of coral populations.

Although there is a substantial body of work on how temperature shapes coastal marine ecosystems, the spatiotemporal variability of seawater pH and corresponding in situ biological responses remain largely unknown across biogeographic ranges of tropical coral species. Environmental variability is important to characterize because it can amplify or dampen the biological consequences of global change, depending on the functional relationship between mean temperature or pH and organismal traits. Here, we characterize the spatiotemporal variability of pH, temperature, and salinity at fringing reefs in Moorea, French Polynesia and Nanwan Bay, Taiwan using advanced time series analysis, including wavelet analysis, and infer their potential impact on the persistence and stability of coral populations. Our results demonstrate that both the mean and variance of pH and temperature differed significantly between sites in Moorea and Taiwan. Seawater temperature at the Moorea site passed the local bleaching threshold several times within the ~45 day deployment while aragonite saturation state at the Taiwan site was often below commonly observed levels for coral reefs. Our results showcase how a better understanding of the differences in environmental conditions between sites can (1) provide an important frame of reference for designing laboratory experiments to study the effects of environmental variability, (2) identify the proximity of current environmental conditions to predicted biological thresholds for the coral reef, and (3) help predict when the temporal variability and mean of environmental conditions will interact synergistically or antagonistically to alter the abundance and stability of marine populations experiencing climate change.

Location, location, location: small shifts in collection site result in large intraspecific differences in macroalgal palatability

The role of herbivorous fishes in coral reef resilience has increased interest in the process of herbivory and has focused attention on herbivore feeding behavior, making it important to evaluate experimental methods used to assess herbivore decisions. We tested whether small-scale differences in collection site play a role in within-species palatability of macroalgae. Baseline grazing assays using algae collected on a fringing reef in Moorea, French Polynesia, revealed that herbivore preferences among three common species ranked Padina boryana > Sargassum mangarevense ≫ Amansia rhodantha. Comparing grazing preferences between individual thalli of the same species collected <15 m apart revealed that consumption of intertidal S. mangarevense was nearly six times greater than for conspecifics collected from the adjacent subtidal reef flat. The same trend occurred for P. boryana but was not significant. This demonstrated that algal palatability can vary on a very small spatial scale, presenting a potential trap for the unwary when setting up experiments; we encourage researchers to consider this potential complication in experimental studies of herbivory.

Responses of the metabolism of the larvae of Pocillopora damicornis to ocean acidification and warming.

Ocean acidification and warming are expected to threaten the persistence of tropical coral reef ecosystems. As coral reefs face multiple stressors, the distribution and abundance of corals will depend on the successful dispersal and settlement of coral larvae under changing environmental conditions. To explore this scenario, we used metabolic rate, at holobiont and molecular levels, as an index for assessing the physiological plasticity of Pocillopora damicornis larvae from this site to conditions of ocean acidity and warming. Larvae were incubated for 6 hours in seawater containing combinations of CO2 concentration (450 and 950 µatm) and temperature (28 and 30°C). Rates of larval oxygen consumption were higher at elevated temperatures. In contrast, high CO2 levels elicited depressed metabolic rates, especially for larvae released later in the spawning period. Rates of citrate synthase, a rate-limiting enzyme in aerobic metabolism, suggested a biochemical limit for increasing oxidative capacity in coral larvae in a warming, acidifying ocean. Biological responses were also compared between larvae released from adult colonies on the same day (cohorts). The metabolic physiology of Pocillopora damicornis larvae varied significantly by day of release. Additionally, we used environmental data collected on a reef in Moorea, French Polynesia to provide information about what adult corals and larvae may currently experience in the field. An autonomous pH sensor provided a continuous time series of pH on the natal fringing reef. In February/March, 2011, pH values averaged 8.075±0.023. Our results suggest that without adaptation or acclimatization, only a portion of naïve Pocillopora damicornis larvae may have suitable metabolic phenotypes for maintaining function and fitness in an end-of-the century ocean.

Diel pCO2 oscillations modulate the response of the coral Acropora hyacinthus to ocean acidification

To investigate the effect of diel variation in pCO2 on coral calcification, branches of Acropora hyacinthus were collected in 2 habitats (upstream and downstream in a unidirectional flow) in a shallow back reef in Moorea, French Polynesia, where different diel amplitudes of pCO2 oscillation were expected. Corals were maintained for 6 wk under different pCO2 regimes (con- stant versus oscillatory), each delivered in 3 configurations: constant conditions of 400 µatm, 700 µatm, and 1000 µatm pCO2, or oscillatory conditions varying daily from 280 to 550 µatm, 550 to 1000 µatm, or 400 to 2000 µatm, with minima and maxima during the day and night, respec- tively. Calcification rates in all treatments tended to increase over time, and the interaction between Time and pCO2 regime (i.e. constant versus oscillating) was significant (or close to signif- icant) for upstream corals due to higher calcification in oscillatory pCO2. A significant pCO2 regime effect was detected in the highest pCO2 for downstream corals, with higher calcification in the 400 to 2000 µatm oscillatory pCO2 treatment compared to the 1000 µatm constant pCO2 treat- ment. After 6 wk, calcification of A. hyacinthus was affected significantly by habitat, the pCO2 level, and the pCO2 regime. Calcification generally was reduced by high pCO2 and was ≥21% greater in 400 to 2000 µatm oscillatory pCO2 versus 1000 µatm constant pCO2 treatment. Increased calcification in the 400 to 2000 µatm oscillatory pCO2 treatment suggests that natural diel oscillations in pCO2 could play a role by reducing the locally negative effects of rising pCO2 associated with ocean acidification on coral calcification.

Distributions of coral reef macroalgae in a back reef habitat in Moorea, French Polynesia

On tropical reefs where macroalgae are subjected to continuous herbivore pressure, spatial refuges typically are identified as large-scale, landscape interfaces that limit foraging behavior. However, algal distributions and community assemblages may also rely on the availability of smaller scale spatial refuges within the reef. The results of this study demonstrate that the patterns of macroalgal distribution across the back reef of Moorea, French Polynesia, are maintained by herbivores interacting with the small-scale structural complexities of the coral reef landscape. Although the majority of space available for colonization is composed of exposed surfaces, macroalgae rarely are found in the open. Instead, macroalgal occurrence is highest in the protected narrow crevices and hole microhabitats provided by massive Porites spp. coral heads. These distributions are determined initially by post-settlement mortality of young algal recruits in exposed habitats. Rates of consumption for two of the most common macroalgal species found in refuges across the back reef, Halimeda minima and Amansia rhodantha, indicate that algal recruits in exposed habitats are limited by herbivory. While algal abundance and community structure are highly dependent upon herbivore grazing, the availability of small-scale spatial refuges ultimately shapes the distinct community patterns and distributional boundaries of coral reef macroalgae in the back reefs of Moorea.

Turbinaria ornata as an herbivory refuge for associate algae

Habitat associations are an integral part of coral reef community structure. Commonly, one organism lives in such close association within or near another that a spatial refuge occurs, whereby one of the organisms provides protection to the other. This is often the result of defenses of the host deterring an associate organism’s consumers. In Moorea, French Polynesia, the range and abundance of the brown macroalga, Turbinaria ornata, have increased drastically since 1980 such that dense aggregations of this macroalga are a dominant component of the backreef habitat. Turbinaria ornata is both mechanically and chemically defended from herbivores. Other species of macroalgae grow within aggregations of Turbinaria and may benefit from these defenses. This study investigates whether aggregations of Turbinaria create a refuge from herbivory for associate macroalgae. When Turbinaria aggregations were removed experimentally, there was a significant increase in the number of associate algal species. Moreover, an herbivory assay using the palatable local alga Acanthophora spicifera identified herbivory as the mechanism for lower diversity on bommies lacking Turbinaria aggregations. The local increase in algal richness due to the refuge from herbivory afforded by Turbinaria may be an important contribution to macroalgal and community dynamics on reefs in Moorea, French Polynesia.

Near-surface enrichment of zooplankton over a shallow back reef: implications for coral reef food webs

Zooplankton were 3–8 times more abundant during the day near the surface than elsewhere in the water column over a 1–2.4 m deep back reef in Moorea, French Polynesia. Zooplankton were also significantly more abundant near the surface at night although gradients were most pronounced under moonlight. Zooplankton in a unidirectional current became concentrated near the surface within 2 m of departing a well-mixed trough immediately behind the reef crest, indicating that upward swimming behavior, rather than near-bottom depletion by reef planktivores, was the proximal cause of these gradients. Zooplankton were highly enriched near the surface before and after a full lunar eclipse but distributed evenly throughout the water column during the eclipse itself supporting light as a proximal cue for the upward swimming behavior of many taxa. This is the first investigation of the vertical distribution of zooplankton over a shallow back reef typical of island barrier reef systems common around the world. Previous studies on deeper fringing reefs found zooplankton depletion near the bottom but no enrichment aloft. In Moorea, where seawater is continuously recirculated out the lagoon and back across the reef crest onto the back reef, selection for upward swimming behavior may be especially strong, because the surface serves both as a refuge from predation and an optimum location for retention within the reef system. Planktivorous fish and corals that can forage or grow even marginally higher in the water column might have a substantial competitive advantage over those nearer the bottom on shallow reefs. Zooplankton abundance varied more over a few tens of centimeters vertical distance than it did between seasons or even between day and night indicating that great care must be taken to accurately assess the availability of zooplankton as food on shallow reefs.

Effects of disturbance on coral communities: bleaching in Moorea, French Polynesia

This study examines patterns of susceptibility and short-term recovery of corals from bleaching. A mass coral bleaching event began in March, 1991 on reefs in Moorea, French Polynesia and affected corals on the shallow barrier reef and to >20 m depth on the outer forereef slope. There were significant differences in the effect of the bleaching among common coral genera, with Acropora, Montastrea, Montipora, and Pocillopora more affected than Porites, Pavona, leptastrea or Millepora. Individual colonies of the common species of Acropora and Pocillopora were marked and their fate assessed on a subsequent survey in August, 1991 to determine rates of recovery and mortality. Ninety-six percent of Acropora spp. showed some degree of bleaching compared to 76% of Pocillopora spp. From March to August mortality of bleached colonies of Pocillopora was 17%, 38% recovered completely, and many suffered some partial mortality of the tissue. In contrast, 63% of the Acropora spp. died, and about 10% recovered completely. Generally, those colonies with less than 50% of the colony area affected by the bleaching recovered at a higher rate than did those with more severe bleaching. Changes in community composition four months after the event began included a significant decrease only in crustose algae and an increase in cover of filamentous algae, much of which occupied plate-like and branching corals that had died in the bleaching event. Total coral cover and cover of susceptible coral genera had declined, but not significantly, after the event.

Darwin's Early Notes on Coral Reef Formation

Darwin's letters and some rough notes found in his field notebooks of 1835 confirm the statement in his Autobiography that he had formulated his theory of coral reef formation before the Beagle left South America and before he had seen a coral reef. His geological observations having convinced him of the elevation of the South American continent, Darwin predicted that evidence of a compensatory gradual subsidence of the Pacific Ocean floor would be found in the existence of shallow-water coral genera in the Pacific reef formations. The first draft of the theory was written on board the Beagle shortly after seeing the reefs of Moorea in November 1835. After visiting the Cocos (Keeling) Islands he wrote a summary of his view in a letter of April 1838, in which he expressed his conviction that he had found an explanation which would "put some of the facts in a more simple and connected point of view, than that in which they have hitherto been considered".