measuring-structure-composition-change-underwater-environments-with

Measuring the Structure, Composition, and Change of Underwater Environments with Large-area Imaging

Digital imaging and processing technology have evolved to facilitate the expansion of large-area imaging surveys, which increase our capacity to study the ...

UC San Diego

The effects of three forms of density-dependent regulation were explored in model coral reef fish populations: top-down (predation), bottom-up (competition for food), and pelagic (non-reef-based mechanisms) control. We describe the demographic responses of both biomass and numbers of adult fish, predicting the mean and the variance of temporal fluctuations resulting from stochastic recruitment of juveniles. We find that top-down control acts by suppressing variability of numbers of fish, which in turn suppresses the variability of biomass. Bottom-up control has no effect on fluctuations of numbers of fish, though it strongly reduces fluctuations of biomass. Because fecundity of fish is directly linked to body mass, the regulation of biomass tightly regulates reproductive output independently of the number of individuals in the population. Finally, populations under pelagic control experience bounded fluctuations of biomass and numbers directly proportional to the bounded fluctuations of recruitment. The demographic signatures predicted from both bottom-up and pelagic control are consistent with current evidence supporting the recruitment limitation hypothesis in reef fish ecology. We propose tests to discriminate the dominant mode of density-dependent regulation using qualitative trends in time series demographic data across environmental clines.

Demographic theory of coral reef fish populations with stochastic recruitment: comparing sources of population regulation.

The community structure of sedentary organisms is largely controlled by the outcome of direct competition for space. Understanding factors defining competitive outcomes among neighbors is thus critical for predicting large-scale changes, such as transitions to alternate states within coral reefs. Using a spatially explicit model, we explored the importance of variation in two spatial properties in benthic dynamics on coral reefs: (1) patterns of herbivory are spatially distinct between fishes and sea urchins and (2) there is wide variation in the areal extent into which different coral species can expand. We reveal that the size-specific, competitive asymmetry of corals versus fleshy algae highlights the significance of spatial patterning of herbivory and of coral growth. Spatial dynamics that alter the demographic importance of coral recruitment and maturation have profound effects on the emergent structure of the reef benthic community. Spatially constrained herbivory (as by sea urchins) is more effective than spatially unconstrained herbivory (as by many fish) at opening space for the time needed for corals to settle and to recruit to the adult population. Further, spatially unconstrained coral growth (as by many branching coral species) reduces the number of recruitment events needed to fill a habitat with coral relative to more spatially constrained growth (as by many massive species). Our model predicts that widespread mortality of branching corals (e.g., Acropora spp) and herbivorous sea urchins (particularly Diadema antillarum) in the Caribbean has greatly reduced the potential for restoration across the region.

Spatial dynamics of benthic competition on coral reefs.

Geographical comparisons suggest that coral reef communities can vary as a function of their environmental context, differing not just in terms of total coral cover but also in terms of relative abundance (or coverage) of coral taxa. While much work has considered how shifts in benthic reef dynamics can shift dominance of stony corals relative to algal and other benthic competitors, the relative performance of coral types under differing patterns of environmental disturbance has received less attention. We construct an empirically-grounded numerical model to simulate coral assemblage dynamics under a spectrum of disturbance regimes, contrasting hydrodynamic disturbances (which cause morphology-specific, whole-colony mortality) with disturbances that cause mortality independently of colony morphology. We demonstrate that the relative representation of morphological types within a coral assemblage shows limited connection to the intensity, and essentially no connection to the frequency, of hydrodynamic disturbances. Morphological types of corals that are more vulnerable to mortality owing to hydrodynamic disturbance tend to grow faster, with rates sufficiently high to recover benthic coverage during inter-disturbance intervals. By contrast, we show that factors causing mortality without linkage to morphology, including those that cause only partial colony loss, more dramatically shift coral assemblage structure, disproportionately favouring fast-growing tabular morphologies. Furthermore, when intensity and likelihood of such disturbances increases, assemblages do not adapt smoothly and instead reveal a heightened level of temporal variance, beyond which reefs demonstrate drastically reduced coral coverage. Our findings highlight that adaptation of coral reef benthic assemblages depends on the nature of disturbances, with hydrodynamic disturbances having little to no effect on the capacity of reef coral communities to resist and recover with sustained coral dominance.

Modelling the linkage between coral assemblage structure and pattern of environmental forcing.

Reef-building coral taxa demonstrate considerable flexibility and diversity in reproduction and growth mechanisms. Corals take advantage of this flexibility to increase or decrease size through clonal expansion and loss of live tissue area (i.e. via reproduction and mortality of constituent polyps). The biological lability of reef-building corals may be expected to map onto varying patterns of demography across environmental contexts which can contribute to geographic variation in population dynamics. Here we explore the patterns of growth of two common coral taxa, corymbose Pocillopora and massive Porites, across seven islands in the central and south Pacific. The islands span a natural gradient of environmental conditions, including a range of pelagic primary production, a metric linked to the relative availability of inorganic nutrients and heterotrophic resources for mixotrophic corals, and sea surface temperature and thermal histories. Over a multi-year sampling interval, most coral colonies experienced positive growth (greater planar area of live tissue in second relative to first time point), though the distributions of growth varied across islands. Island-level median growth did not relate simply to estimated pelagic primary productivity or temperature. However, at locations that experienced an extreme warm-water event during the sampling interval, most Porites colonies experienced net losses of live tissue and nearly all Pocillopora colonies experienced complete mortality. While descriptive statistics of demographics offer valuable insights into trends and variability in colony change through time, simplified models predicting growth patterns based on summarized oceanographic metrics appear inadequate for robust demographic prediction. We propose that the complexity of life history strategies among colonial reef-building corals introduces unique demographic flexibility for colonies to respond to a wide breadth of environmental conditions.

Considering the rates of growth in two taxa of coral across Pacific islands.

Predation is ubiquitous on coral reefs. Among the most charismatic group of reef predators are the top predatory fishes, including sharks and large-bodied bony fishes. Despite the threat presented by top predators, data describing their realized effects on reef community structure and functioning are challenging to produce. Many innovative studies have capitalized on natural experimental conditions to explore predator effects on reefs. Gradients in predator density have been created by spatial patterning of fisheries management. Evidence of prey release has been observed across some reefs, namely that potential prey increase in density when predator density is reduced. While such studies search for evidence of prey release among broad groups or guilds of potential prey, a subset of studies have sought evidence of release at finer population levels. We find that some groups of fishes are particularly vulnerable to the effects of predators and more able to capitalize demographically when predator density is reduced. For example, territorial damselfish appear to realize reliable population expansion with the reduction in predator density, likely because their aggressive, defensive behavior makes them distinctly vulnerable to predation. Relatedly, individual fishes that suffer from debilitating conditions, such as heavy parasite loads, appear to realize relatively stronger levels of prey release with reduced predator density. Studying the effects of predators on coral reefs remains a timely pursuit, and we argue that efforts to focus on the specifics of vulnerability to predation among potential prey and other context-specific dimensions of mortality hold promise to expand our knowledge.

Emerging insights on effects of sharks and other top predators on coral reefs.

For many organisms, early life stages experience significantly higher rates of mortality relative to adults. However, tracking early life stage individuals through time in natural settings is difficult, limiting our understanding of the duration of these 'mortality bottlenecks', and the time required for survivorship to match that of adults. Here, we track a cohort of juvenile corals (1-5 cm maximum diameter) from 12 taxa at a remote atoll in the Central Pacific from 2013 to 2017 and describe patterns of annual survivorship. Of the 537 juveniles initially detected, 219 (41%) were alive 4 years later, 163 (30%) died via complete loss of live tissue from the skeleton, and the remaining 155 (29%) died via dislodgement. The differing mortality patterns suggest that habitat characteristics, as well as species-specific features, may influence early life stage survival. Across most taxa, survival fit a logistic model, reaching > 90% annual survival within 4 years. These data suggest that mortality bottlenecks characteristic of 'recruitment' extend up to 5 years after individuals can be visually detected. Ultimately, replenishment of adult coral populations via sexual reproduction is needed to maintain both coral cover and genetic diversity. This study provides key insights into the dynamics and time scales that characterize these critical early life stages.

Article	Year
Demographic theory of coral reef fish populations with stochastic recruitment: comparing sources of population regulation. The American naturalist\| PubMed ID: 15729643	2005
Spatial dynamics of benthic competition on coral reefs. Oecologia\| PubMed ID: 22009340	2012
Modelling the linkage between coral assemblage structure and pattern of environmental forcing. Royal Society open science\| PubMed ID: 33204448	2020
Considering the rates of growth in two taxa of coral across Pacific islands. Advances in marine biology\| PubMed ID: 33293010	2020
Emerging insights on effects of sharks and other top predators on coral reefs. Emerging topics in life sciences\| PubMed ID: 35258079	2022
Post-settlement demographics of reef building corals suggest prolonged recruitment bottlenecks. Oecologia\| PubMed ID: 35661251	2022

Stuart A. Sandin

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