M. Julian Caley’s research while affiliated with James Cook University and other places

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Publications (177)


Assuming stationarity compromises understanding of the dynamics and management of open marine populations
  • Preprint

March 2022

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7 Reads

Carla Chen

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M. Julian Caley

Understanding connectivity of marine species is crucial for their management. This connectivity, however, is difficult to quantify; propagules of marine species are typically small, numerous, and can travel large distances. Consequently, oceanographic models are often used to simulate larval dispersal. To avoid high computational costs, these models typically use, pooled or cross-sectional data and fixed biological parameters. Here we explore how temporal and spatial variation in current velocities, and pelagic larval duration and buoyancy can alter patterns of marine connectivity of Crown-of-Thorns starfish on Australia’s Great Barrier Reef. Our results reveal highly variable reef connectivity in space and time mediated by all three factors. No individual reef acted consistently as a population source or sink. It is, therefore, important to consider many factors concurrently when estimating connectivity for understanding these population dynamics, especially where such estimates are relied on for evidence-based decision making.


Fig. 1. A hypothetical agrivoltaic system in Australia. Solar panels are co-located within croplands and on existing grazing land.
Fig. 2. Conceptual model of components and interactions that influence land condition in agrivoltaic systems. Five major components are summarized through their interactions (represented by a -f and described in the text) by: Biodiversity (wildlife habitat, species diversity, refugia and cover, pollinator and predators, and ecosystem services); Soil and Water (run-off, erosion, soil moisture, soil compaction, dust accumulation, nutrients, fertilizers); Vegetation (ground cover, vegetation complexity, weeds, food production); Maintenance (clearing, infrastructure, mowing, herbicide use, pesticide use, livestock health, cultivation and harvest); Financial returns (carbon credits, electricity production, jobs, site maintenance, pest control, profit through livestock/crop yields, supplemental feeding for livestock).
Designing solar farms for synergistic commercial and conservation outcomes
  • Article
  • Full-text available

November 2021

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3,893 Reads

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39 Citations

Solar Energy

Competition among land uses is making it increasingly difficult to set aside adequate space for wildlife and nature conservation, so it is imperative that opportunities that simultaneously achieve commercial and conservation outcomes be identified and seized. Such opportunities exist in the renewable energy industry. It is widely recognized that renewable energy generation benefits the ecosphere through reduced carbon emissions, but currently, further opportunities for realising direct and indirect conservation benefits through the design of solar farms are less well known. Among other opportunities, solar farm designs that deliver environmental credits through carbon sequestration and biodiversity improvements can deliver higher financial returns. Other opportunities to improve local hydrology, pollination, and pest-control services could be available depending on site-specific characteristics where solar farms are built, and the other land use practices that exist, or are possible, in the immediate vicinity. Here, we explore opportunities among renewable energy generation, agriculture, and conservation, through the co-location and innovative design of PV solar energy farms on grazing and croplands. These forms of land sharing can achieve higher land-equivalent ratios (LERs), a quantitative metric of the reduction in land use. We identify opportunities whereby solar farms can be designed to improve biodiversity, land condition, and conservation outcomes, while maintaining or increasing commercial returns. Much work remains, however, to understand the suite of opportunities available for achieving simultaneously the best commercial and conservation outcomes through solar farm designs in agricultural landscapes.

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Map of study site; (a) regional map showing study region (red square); (b) close‐up of the Oceanic Shoals Australian Marine Park showing location of study sites
Example showing differences in predicted likelihood of occurrence between raised geomorphic features1a). Depth (a); predicted likelihood of occurrence for sponges (b); predicted likelihood of occurrence for hard corals (c); total number of taxa predicted for section of the western Oceanic Shoals AMP. Predictions cover Area 4 from regional overview (Figure 1)
Contribution of each predictor variable to the total deviance explained by boostred regression tree models for each of the four taxa
Transferable, predictive models of benthic communities informs marine spatial planning in a remote and data‐poor region

July 2020

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138 Reads

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8 Citations

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M. Julian Caley

Systematic conservation planning requires spatial information on biodiversity. Such information is often unavailable, forcing spatial planning to rely on assumed relationships between species and environmental features. This problem is particularly acute in large, remote marine protected areas that are proliferating rapidly. Here, we use models to predict whether (a) macrobenthic biodiversity across four taxa (gorgonians, soft corals, hard corals, and sponges) with different life histories are congruent within seascape features through regional space; and (b) models generated in an intensively‐sampled area in one region can predict the occurrence of habitat‐forming macrobenthos in neighboring ones. All four taxa studied showed similar habitat preferences, but high variability in distributions among and within features suggesting factors other than simple geomorphology influence these regional biodiversity patterns. Nonetheless, models derived from one region accurately predicted the presence and absence of the same taxa hundreds of kilometers away. This transferability of models of species occurrences has the potential to deliver improved reserve design in data‐deficient regions.


Latitude and protection affect decadal trends in reef trophic structure over a continental scale

June 2020

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155 Reads

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7 Citations

Abstract The relative roles of top‐down (consumer‐driven) and bottom‐up (resource‐driven) forcing in exploited marine ecosystems have been much debated. Examples from a variety of marine systems of exploitation‐induced, top‐down trophic forcing have led to a general view that human‐induced predator perturbations can disrupt entire marine food webs, yet other studies that have found no such evidence provide a counterpoint. Though evidence continues to emerge, an unresolved debate exists regarding both the relative roles of top‐down versus bottom‐up forcing and the capacity of human exploitation to instigate top‐down, community‐level effects. Using time‐series data for 104 reef communities spanning tropical to temperate Australia from 1992 to 2013, we aimed to quantify relationships among long‐term trophic group population density trends, latitude, and exploitation status over a continental‐scale biogeographic range. Specifically, we amalgamated two long‐term monitoring databases of marine community dynamics to test for significant positive or negative trends in density of each of three key trophic levels (predators, herbivores, and algae) across the entire time series at each of the 104 locations. We found that trophic control tended toward bottom‐up driven in tropical systems and top‐down driven in temperate systems. Further, alternating long‐term population trends across multiple trophic levels (a method of identifying trophic cascades), presumably due to top‐down trophic forcing, occurred in roughly fifteen percent of locations where the prerequisite significant predator trends occurred. Such alternating trophic trends were significantly more likely to occur at locations with increasing predator densities over time. Within these locations, we found a marked latitudinal gradient in the prevalence of long‐term, alternating trophic group trends, from rare in the tropics (


Fig. 1.3. A. Spatial locations of surveyed reefs within region along the Great Barrier Reef. B Cumulative sum of disturbances across all types per sub-region.
1. Values of DIC for the three published versions of the semi-parametric Bayesian hierarchical models
Thresholds of Coral Cover That Support Coral Reef Biodiversity

May 2020

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604 Reads

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1 Citation

Lecture Notes in Mathematics -Springer-verlag-

Global environmental change, such as ocean warming and increased cyclone activity, is driving widespread and rapid declines in the abundance of key ecosystem engineers, reef-building corals, on the Great Barrier Reef. Our ability to understand how coral associated species, such as reef fishes, respond to coral loss can be impeded by uncertainty surrounding natural spatio-temporal variability of coral populations. To address this issue, we developed a semi-parametric hierarchical Bayesian model to estimate long-term trajectories of habitat-forming coral cover as a function of three spatial scales (sub-region, habitat and site) and environmental disturbances. The relationships between coral cover trajectories and fish community structure were examined using posterior predictive distributions of estimated coral cover from the statistical model. In the absence of direct observations of fish community structure, we used the probability of coral cover being above some ecological threshold values as a proxy for potential disruptions of fish community structure. Threshold values were derived from published field studies that estimated changes in the structure of coral-reef fish communities and coral cover after major disturbances. In these studies, fish community structure did not change where post-disturbance coral cover was > 20%. Disruptions in the structure of these communities were observed when coral cover dropped to between 10–20% and declines in fish diversity were typical where coral cover ranged from between 5 and 10%. Based on these thresholds values, posterior probabilities of coral cover being above 20% and between 10 and 20% and between 5 and 10% were calculated across spatial scales on the Great Barrier Reef (GBR) from 1995 to 2011. At the GBR scale, probabilities of coral cover being above these thresholds remained relatively stable through time. Across years, probabilities of coral cover being at least > 20% remained null for the sub-regions of Cairns, Townsville, Whitsundays and Swain but highly variable between reef sites within these sub-regions, with the exception of Townsville. In the Townsville area, probabilities of coral cover being between 10–20% and 5–10% declined from 0.75 to 0 during the study period. This finding highlights potential sub-regional fish community structure disruptions which have not yet been observed at this spatial scale. As frequency and intensity of disturbance events continue to rise, and consequently, as coral cover declines further, the probabilistic Bayesian approach presented in this chapter could be used to help provide early warnings of major ecological shifts at management relevant scales in the absence of direct observations.


Bayesian Learning of Biodiversity Models Using Repeated Observations

May 2020

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33 Reads

Lecture Notes in Mathematics -Springer-verlag-

Predictive biodiversity distribution models (BDM) are useful for understanding the structure and functioning of ecological communities and managing them in the face of anthropogenic disturbances. In cases where their predictive performance is good, such models can help fill knowledge gaps that could only otherwise be addressed using direct observation, an often logistically and financially onerous prospect. The cornerstones of such models are environmental and spatial predictors. Typically, however, these predictors vary on different spatial and temporal scales than the biodiversity they are used to predict and are interpolated over space and time. We explore the consequences of these scale mismatches between predictors and predictions by comparing the results of BDMs built to predict fish species richness on Australia’s Great Barrier Reef. Specifically, we compared a series of annual models with uninformed priors with models built using the same predictors and observations, but which accumulated information through time via the inclusion of informed priors calculated from previous observation years. Advantages of using informed priors in these models included (1) down-weighting the importance of a large disturbance, (2) more certain species richness predictions, (3) more consistent predictions of species richness and (4) increased certainty in parameter coefficients. Despite such advantages, further research will be required to find additional ways to improve model performance.


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Figure 6: Graphical Abstract
Forecasting intensifying disturbance effects on coral reefs

March 2020

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339 Reads

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69 Citations

Global Change Biology

Anticipating future changes of an ecosystem’s dynamics requires knowledge of how its key communities respond to current environmental regimes. The Great Barrier Reef (GBR) is under threat, with rapid changes of its reef‐building hard coral community structure already evident across broad spatial scales. While many of the underlying relationships between hard corals and multiple disturbances have been documented, responses of other benthic communities to disturbances are not well understood. Here, we used statistical modelling to explore the effects of broad‐scale climate‐related disturbances on benthic communities to predict their structure under scenarios of increasing disturbance frequency. We parameterized a multivariate model using the composition of benthic communities estimated by 145,000 observations from the northern GBR between 2012 and 2017. During this time, surveyed reefs were variously impacted by two tropical cyclones and two heat stress events that resulted in extensive hard coral mortality. This unprecedented sequence of disturbances was used to estimate the effects of discrete versus interacting disturbances on the compositional structure of hard corals, soft corals, and algae. Discrete disturbances increased the prevalence of algae relative to hard corals while the interaction between cyclones and heat stress was the main driver of the increase in soft corals relative to algae and hard corals. Predictions from disturbance scenarios included relative increases in algae versus soft corals that varied by the frequency and types of disturbance interactions. However, high un‐certainty of compositional changes in the presence of several disturbances shows that responses of algae and soft corals to the decline in hard corals needs further research. Better understanding the effects of multiple disturbances on benthic communities as a whole is essential for predicting the future status of coral reefs and managing them in the light of new environmental regimes. The approach we develop here opens new opportunities for reaching this goal. Model code available at: https://github.com/JulieVercelloni/multivariate_abundance_model_GBR


of five hierarchies of evidence drawn from the medical, health care and epidemiological literature. Text and enumeration are taken from the original references. Stevens and Milne (1997) is reproduced from Pullin and Knight (2001). With the exception of non-randomized controlled trials, the structure of each of the five hierarchies is remarkably equivalent allowing studies to be allocated to one of five broadly equivalent groups, ranked from highest (top) to weakest (bottom). We have added a color scheme to help emphasize this similarity and also identify one outlier: Stevens and Milne (1997) rank well designed controlled trials without randomization relatively high, whereas Wilson et al. (2015) rank this type of study design as providing a much weaker type of evidence.
Schematic illustration of the evidence hierarchy for randomized controlled trials, non-randomized controlled trials and cross over studies. The die represents randomization at the level of treatment. The strength of evidence is always stronger with treatment randomization (first column on the left) than without it (second column on the left). The colors in the columns to the left reflect the evidence hierarchies presented in Figure 1. The strength of evidence is ranked from highest (1) to lowest (5). The “treatment” in this context could be the designation of a new MPA, a change in zonation status, or the implementation of new education or compliance regime (Table 1). The figure is adapted from Wilson et al. (2015) who adapted Walshe et al. (2013).
Schematic illustration of the evidence hierarchy for observational studies and expert judgment. The die represents selection of a representative sample by, for example, a randomized sampling scheme. The strength of evidence is diminished if the sample is not representative of the target population (second column on the left). The colors in the columns to the left reflect the evidence hierarchies presented in Figure 1. The strength of evidence is ranked from highest (1) to lowest (5). This figure is adapted from Wilson et al. (2015).
Directed acyclic graph highlighting the effects of confounding variables on the observed effect of declaring a MPA. Single-headed arrows represent causal links. Stochastic nodes (variables) are represented by filled circles, deterministic nodes by filled squares. The node FT is the deterministic management action of declaring a no-take MPA. The resulting MPA (node T) is defined by characteristics (such as fishing effort, meta-population effects) that determine the probability of observed outcomes such as the biomass of demersal fish (node Y). A confounding variable is anything that affects both the probability of treatment (the link marked “a”) and the outcomes that the treatment is designed to alter (the link marked “b”). The effect of treatment T on outcome Y could be confounded by U if (for example) the management action is more likely to declare small MPAs, in a region isolated from propagules sources, with relatively poor habitat or in regions that have different levels of localized fishing effort inside versus outside of the proposed MPA boundaries. Monitoring programs that do not control for these variables could over- or under-estimate the effect of management. Non-confounding variables (node G), such as long term climatic shifts, are things that influence the treatment outcomes but do not influence the probability of treatment. They cause conditions inside and outside reserve boundaries to vary independently of the reserve treatment and thereby tend to reduce the power of statistical tests to detect a reserve effect. Measurement error is typically considered to be a non-confounding variable, but it can act as a confounding variable if it interacts with the treatment effect. For example, fish in a no-take zone of an MPA may be less disturbed by SCUBA observers and so more likely to be counted when compared to (spear)fished individuals outside the no-take zone.
of the adequacy of the evidence base, and associated key issues, when considering the types of questions that marine reserve managers might try to answer with data obtained from different levels of the evidence hierarchy. Study design codes are as follows: (1) Expert judgment (EJ), (2) Uncontrolled trial (UT), (3) Uncontrolled time series (UTS), (4) Uncontrolled before after study (UBAS), (5) Uncontrolled interrupted time series (UITS), (6) Controlled trial with non-randomized treatment (CTNRT), (7) Controlled time series with non-randomised treatment (CTNRT), (8) Cross sectional analysis with non-representative sample (CSNRS), (9) Cohort study with non-randomised sample (CNRS), (10) Case control study with non-representative sample (CCSNRS), (11) Cross over study with non-randomised treatment (COSNRT), (12) Controlled before after study with non-randomised treatment (CBASNRT), (13) Controlled interrupted time series with non-randomised treatment (CITSNRT), (14) Cross sectional analysis with representative sample (CSRS), (15) Cohort study with representative sample (CRS), (16) Case control study with representative sample (CCSRS), (17) Cross over study with randomised treatment (COSRT), (18) Randomised controlled trial (RCT), (19) Randomised controlled time series (RCTS), (20) Randomised controlled before-after study (RCBAS), and (21) Randomised controlled interrupted time series (RCITS).
Designing Monitoring Programs for Marine Protected Areas Within an Evidence Based Decision Making Paradigm

November 2019

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327 Reads

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28 Citations

The Evidence Based Decision Making (EBDM) paradigm encourages managers to base their decisions on the strongest available evidence, but it has been criticized for placing too much emphasis on the choice of study design method without considering the types of questions that are being addressed as well as other relevant factors such as how well a study is implemented. Here we review the objectives of Australia’s Marine Park network, and identify the types of questions and data analysis that would address these objectives. Critically, we consider how the design of a monitoring program influences our ability to adequately answer these questions, using the strength of evidence hierarchy from the EBDM paradigm to assess the adequacy of different design strategies and other sources of information. It is important for conservation managers to recognize that the types of questions monitoring programs are able to answer depends on how they are designed and how the collected data are analyzed. The socio-political process that dictates where protected areas are placed typically excludes the strongest types of evidence, Random Controlled Trials (RCTs), for certain questions. Evidence bases that are stronger than ones commonly employed to date, however, could be used to provide a causal inference, including for those questions where RCTs are excluded, but only if appropriate designs such as cohort or case-control studies are used, and supported where relevant by appropriate sample frames. Randomized, spatially balanced sampling, together with careful selection of control sites, and more extensive use of propensity scores and structured elicitation of expert judgment, are also practical ways to improve the evidence base for answering the questions that underlie marine park objectives and motivate long-term monitoring programs.


Monitoring through many eyes: Integrating disparate datasets to improve monitoring of the Great Barrier Reef

November 2019

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359 Reads

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22 Citations

Environmental Modelling & Software

Numerous organisations collect data in the Great Barrier Reef (GBR), but they are rarely analysed together due to different program objectives, methods, and data quality. We developed a weighted spatio-temporal Bayesian model and used it to integrate image-based hard-coral data collected by professional and citizen scientists, who captured and/or classified underwater images. We used the model to predict coral cover across the GBR with estimates of uncertainty; thus filling gaps in space and time where no data exist. Additional data increased the model's predictive ability by 43%, but did not affect model inferences about pressures (e.g. bleaching and cyclone damage). Thus, effective integration of professional and high-volume citizen data could enhance the capacity and cost-efficiency of monitoring programs. This general approach is equally viable for other variables collected in the marine environment or other ecosystems; opening up new opportunities to integrate data and provide pathways for community engagement/stewardship.


Testing biodiversity theory using species richness of reef-building corals across a depth gradient

October 2019

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120 Reads

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17 Citations

Natural environmental gradients encompass systematic variation in abiotic factors that can be exploited to test competing explanations of biodiversity patterns. The species-energy (SE) hypothesis attempts to explain species richness gradients as a function of energy availability. However, limited empirical support for SE is often attributed to idiosyncratic, local-scale processes distorting the underlying SE relationship. Meanwhile, studies are also often confounded by factors such as sampling biases, dispersal boundaries and unclear definitions of energy availability. Here, we used spatially structured observations of 8460 colonies of photo-symbiotic reef-building corals and a null-model to test whether energy can explain observed coral species richness over depth. Species richness was left-skewed, hump-shaped and unrelated to energy availability. While local-scale processes were evident, their influence on species richness was insufficient to reconcile observations with model predictions. Therefore, energy availability, either in isolation or in combination with local deterministic processes, was unable to explain coral species richness across depth. Our results demonstrate that local-scale processes do not necessarily explain deviations in species richness from theoretical models, and that the use of idiosyncratic small-scale factors to explain large-scale ecological patterns requires the utmost caution.


Citations (51)


... The same is true for Environmental Stewardship schemes, and at all buffers sizes there is one scheme type (Entry Level plus Higher Level) significantly smaller around solar farms than around the random points. Landowners inclined to set up land conservation initiatives might favour also incorporating a renewable energy source, such as solar farms, particularly due to the additional grants and subsidies available such as environmental credits for carbon sequestration and biodiversity improvements (Nordberg et al., 2021). However, in the UK, land owners and businesses are being steered financially toward rooftop solar panelling (Rural Payments Agency, 2024). ...

Reference:

The location of solar farms within England's ecological landscape: Implications for biodiversity conservation
Designing solar farms for synergistic commercial and conservation outcomes

Solar Energy

... The seasonal conditions and vast, remote, and inaccessible landscapes of Alaska are significant challenges for bat research. Yet, predictive modeling has proven to be a useful and reliable tool in planning research and conservation efforts in data-poor regions (Santos et al. 2013;Wood et al. 2018;Bridge et al. 2020;Becker et al. 2022). Here, we present a model predicting the spatial distribution of MYLU foraging habitat to a large, understudied region of coastal Alaska. ...

Transferable, predictive models of benthic communities informs marine spatial planning in a remote and data‐poor region

... Generally, as water temperature decreases omnivorous and carnivorous (e.g. planktivore, invertivore and piscivore) foraging strategies appear to become relatively more common (Longo et al., 2019;Madin et al., 2020), whereas a higher diversity and abundance of herbivorous fish species is found in tropical waters (e.g. >15°C annual SST; Floeter et al., 2005;Vejříková et al., 2016). ...

Latitude and protection affect decadal trends in reef trophic structure over a continental scale

... Anthropogenic and natural disturbances are increasingly exerting pressures on coral reef ecosystems worldwide, resulting in widespread coral mortality, impaired growth, and reduced fecundity (Bellwood et al. 2004;Bauman et al. 2013;Vercelloni et al. 2020; Emslie et al. 2024). Under low to moderate levels of disturbance, coral reefs can be expected to recover (Wilkinson et al. 1999; Graham et al. 2015), but on reefs that experience frequent or prolonged disturbances, the cumulative impact of coral loss may diminish recovery capacity and promote changes in community composition (Connell et al. 1997 Mudge et al. 2023). ...

Forecasting intensifying disturbance effects on coral reefs

Global Change Biology

... Unlike in experimental studies, because observational studies such as this one do not experimentally manipulate specific variables and control for others, there is uncertainty over the inferred responses (causal impacts, such as a species-specific catch rate) to a single predictor (Hayes et al., 2019). However, applying quasi-experimental statistical modeling approaches to infer responses to an intervention using standardized fisheries data provide estimates with lower risk of error and bias than observational studies with nominal estimates that do not explicitly account for simultaneous variability in potentially informative predictors of a response (Potts & Rose, 2018;Venables & Dichmont, 2004). ...

Designing Monitoring Programs for Marine Protected Areas Within an Evidence Based Decision Making Paradigm

... VRD employs a weighted likelihood method for combining data from disparate sources in order to obtain combined estimates of coral cover on the GBR. This approach, described by Peterson et al. (2020), involves two steps. In the first step, a set of mechanistic weights is determined for each input source, based on a pre-specified measure of their respective accuracy obtained using a gold standard dataset. ...

Monitoring through many eyes: Integrating disparate datasets to improve monitoring of the Great Barrier Reef

Environmental Modelling & Software

... The full model considered was: depth + depth 2 + location + depth:location + depth 2 :location. A parabolic relationship with depth (i.e., a quadratic term of depth) was included in the models to account for potential hump-shaped relationships between depth on macro-benthic cover and coral richness 110 . The most parsimonious combinations of fixed effects were found using model selection based on AICc (models fitted using maximum likelihood). ...

Testing biodiversity theory using species richness of reef-building corals across a depth gradient

... The morphology and the degree of branching in coral colonies affect not only the light field but also the water flow around the colonies. The flow determines the thickness and shape of boundary layers for momentum transfer (momentum boundary layer, MBL, i.e., water velocity change near the coral surface), mass transfer (diffusive boundary layer, DBL, i.e., concentration change in the diffusion-dominated region adjacent to the coral tissue) (Chan et al., 2016), as well as heat transfer (thermal boundary layer, TBL, referring to temperature changes) (Jimenez et al., 2008;Ong et al., 2019). The coral consumes part of the O 2 produced by the symbionts under light (Al-Horani et al., 2003), while an eventual surplus of oxygen and heat generated in the tissue is transported into the surrounding water column and the coral skeleton. ...

The effect of small-scale morphology on thermal dynamics in coral microenvironments
  • Citing Article
  • October 2019

Journal of Thermal Biology

... Numerous studies have deployed BRUVs within and around MPAs and they have been used to demonstrate changes linked to management measures as well as identifying suitable sites for protection (Letessier et al., 2019). Although historically used for imaging benthic and demersal species, there has been increasing interest in deploying BRUVs in the water column. ...

Correction: Remote reefs and seamounts are the last refuges for marine predators across the Indo-Pacific

... Principal component analysis (PCA) is a widely used tool in ecological applications to interpret high-dimensional data (Huettmann & Diamond, 2001;Janžekovič & Novak, 2012;Jolliffe, 1986;Pearson, 1901;Wu et al., 2019). However, PCA has significant drawbacks in the nonlinear, multivariate setting of the current application. ...

Analysing the dynamics and relative influence of variables affecting ecosystem responses using functional PCA and boosted regression trees: A seagrass case study
  • Citing Article
  • August 2019