Analysing biological colour patterns from digital images: An introduction to the current toolbox.

Understanding the numerous roles that colouration serves in the natural world has remained a central focus in many evolutionary and ecological studies. However, to accurately characterise and then compare colours or patterns among individuals or species has been historically challenging. In recent years, there have been a myriad of new resources developed that allow researchers to characterise biological colours and patterns, specifically from digital imagery. However, each resource has its own strengths and weaknesses, answers a specific question and requires a detailed understanding of how it functions to be used properly. These nuances can make navigating this emerging field rather difficult. Herein, we evaluate several new techniques for analysing biological colouration, with a specific focus on digital images. First, we introduce fundamental background knowledge about light and perception to be considered when designing and implementing a study of colouration. We then show how numerous modifications can be made to images to ensure consistent formatting prior to analysis. After, we describe many of the new image analysis approaches and their respective functions, highlighting the type of research questions that they can address. We demonstrate how these various techniques can be brought together to examine novel research questions and test specific hypotheses. Finally, we outline potential future directions in colour pattern studies. Our goal is to provide a starting point and pathway for researchers wanting to study biological colour patterns from digital imagery.

Quantum of fear: Herbivore grazing rates not affected by reef shark presence.

Grazing by nominally herbivorous fishes is widely recognised as a critical ecosystem function on coral reefs. However, several studies have suggested that herbivory is reduced in the presence of predators, especially sharks. Nevertheless, the effects of shark presence on grazing, under natural settings, remains poorly resolved. Using ∼200 h of video footage, we quantify the extent of direct disturbance by reef sharks on grazing fishes. Contrary to expectations, grazing rate was not significantly suppressed due to sharks, with fishes resuming feeding in as little as 4 s after sharks passed. Based on our observations, we estimate that an average m2 area of reef at our study locations would be subjected to ∼5 s of acute shark disturbance during daylight hours. It appears the short-term impact of reef shark presence has a negligible effect on herbivore grazing rates, with the variable nature of grazing under natural conditions overwhelming any fear effects.

Meta-analysis reveals weak associations between reef fishes and corals.

Habitat associations underpin species ecologies in high-diversity systems. Within tropical, shallow water coral reefs, the relationship between fishes and corals is arguably the most iconic and highly scrutinized. A strong relationship between fishes and reef-building hard corals is often assumed, a belief supported by studies that document the decline of reef fishes following coral loss. However, the extent of this relationship is often unclear, as evidenced by conflicting reports. Here we assess the strength of this ecological association by relying on literature that has surveyed both fishes and corals synchronously. We quantitatively synthesize 723 bivariate correlation coefficients (from 66 papers), published over 38 years, that relate fish metrics (abundance, biomass and species richness) with the percentage of hard coral cover. Remarkably, despite extensive variation, the pattern of association on a global scale reveals a predominantly positive, albeit weak (|r| < 0.4), correlation. Even for commonly hypothesized drivers of fish-coral associations, fish family and trophic group, associations were consistently weak. These findings question our assumptions regarding the strength and ubiquity of fish-coral associations, and caution against assuming a direct and omnipresent relationship between these two iconic animal groups.

Natural recovery of corals after severe disturbance.

Ecosystem recovery from human-induced disturbances, whether through natural processes or restoration, is occurring worldwide. Yet, recovery dynamics, and their implications for broader ecosystem management, remain unclear. We explored recovery dynamics using coral reefs as a case study. We tracked the fate of 809 individual coral recruits that settled after a severe bleaching event at Lizard Island, Great Barrier Reef. Recruited Acropora corals, first detected in 2020, grew to coral cover levels that were equivalent to global average coral cover within just 2 years. Furthermore, we found that just 11.5 Acropora recruits per square meter were sufficient to reach this cover within 2 years. However, wave exposure, growth form and colony density had a marked effect on recovery rates. Our results underscore the importance of considering natural recovery in management and restoration and highlight how lessons learnt from reef recovery can inform our understanding of recovery dynamics in high-diversity climate-disturbed ecosystems.

Hot spots of bleaching in massive Porites coral colonies.

Coral bleaching events have become more frequent and severe due to ocean warming. While the large-scale impacts of bleaching events are well-known, there is growing recognition of the importance of small-scale spatial variation in bleaching and survival probability of individual coral colonies. By quantifying bleaching in 108 massive Porites colonies spread across Lizard Island, Great Barrier Reef, during the 2016 bleaching event, we investigated how hydrodynamic exposure levels and colony size contribute to local variability in bleaching prevalence and extent. Our results revealed that exposed locations were the least impacted by bleaching, while lagoonal areas exhibited the highest prevalence of bleaching and colony-level bleaching extents. Such patterns of bleaching could be due to prolonged exposure to warm water in the lagoon. These findings highlight the importance of considering location-specific factors when assessing coral health and emphasize the vulnerability of corals in lagoonal habitats to rapid and/or prolonged elevated temperatures.

Spatial and temporal variability in tropical off-reef zooplankton across broad spatial and temporal scales.

Productivity of oligotrophic coral reefs is largely dependent on the constant influx of zooplankton. However, our understanding of how zooplankton communities in tropical reef-associated regions vary over large spatial and temporal scales is limited. Using the Australian continuous plankton recorder dataset, we explored if, and to what extent, the off-reef zooplankton community along the Queensland shelf (including most of the Great Barrier Reef lagoon) varied with latitude, month, and diel time. The zooplankton community was consistently dominated by copepods (∼60%) which, with appendicularians, chaetognaths, non-copepod crustaceans, and thaliaceans, comprised ∼98% of the zooplankton. However, the abundance of these taxonomic groups did not vary predictably across latitude, month, or diel time, with these gradients only explaining 5% of community variation. At the scales sampled herein the composition of zooplankton was highly predictable in terms of broad taxonomic groups but variation in the relative abundance of these groups was not predictable.

Bio-physical determinants of sediment accumulation on an offshore coral reef: A snapshot study.

Sediments are found on all coral reefs around the globe. However, the amount of sediment in different reservoirs, and the rates at which sediments move between reservoirs, can shape the biological functioning of coral reefs. Unfortunately, relatively few studies have examined reef sediment dynamics, and associated bio-physical drivers, simultaneously over matching spatial and temporal scales. This has led to a partial understanding of how sediments and living reef systems are connected, especially on clear-water offshore reefs. To address this problem, four sediment reservoirs/sedimentary processes and three bio-physical drivers were quantified across seven different reef habitats/depths at Lizard Island, an exposed mid-shelf reef on the Great Barrier Reef. Even in this clear-water reef location a substantial load of suspended sediment passed over the reef; a load theoretically capable of replacing the entire standing stock of on-reef turf sediments in just 8 h. However, quantification of actual sediment deposition suggested that just 2 % of this passing sediment settled on the reef. The data also revealed marked spatial incongruence in sediment deposition (sediment trap data) and accumulation (TurfPod data) across the reef profile, with the flat and back reef emerging as key areas of both deposition and accumulation. By contrast, the shallow windward reef crest was an area of deposition but had a limited capacity for sediment accumulation. These cross-reef patterns related to wave energy and reef geomorphology, with low sediment accumulation on the ecologically important reef crest aligning with substantial wave energy. These findings reveal a disconnect between patterns of sediment deposition and accumulation on the benthos, with the 'post-settlement' fate of sediments dependent on local hydrodynamic conditions. From an ecological perspective, the data suggests key contextual constraints (wave energy and reef geomorphology) may predispose some reefs or reef areas to high-load turf sediment regimes.

Research priorities for the sustainability of coral-rich western Pacific seascapes.

Nearly a billion people depend on tropical seascapes. The need to ensure sustainable use of these vital areas is recognised, as one of 17 policy commitments made by world leaders, in Sustainable Development Goal (SDG) 14 ('Life below Water') of the United Nations. SDG 14 seeks to secure marine sustainability by 2030. In a time of increasing social-ecological unpredictability and risk, scientists and policymakers working towards SDG 14 in the Asia-Pacific region need to know: (1) How are seascapes changing? (2) What can global society do about these changes? and (3) How can science and society together achieve sustainable seascape futures? Through a horizon scan, we identified nine emerging research priorities that clarify potential research contributions to marine sustainability in locations with high coral reef abundance. They include research on seascape geological and biological evolution and adaptation; elucidating drivers and mechanisms of change; understanding how seascape functions and services are produced, and how people depend on them; costs, benefits, and trade-offs to people in changing seascapes; improving seascape technologies and practices; learning to govern and manage seascapes for all; sustainable use, justice, and human well-being; bridging communities and epistemologies for innovative, equitable, and scale-crossing solutions; and informing resilient seascape futures through modelling and synthesis. Researchers can contribute to the sustainability of tropical seascapes by co-developing transdisciplinary understandings of people and ecosystems, emphasising the importance of equity and justice, and improving knowledge of key cross-scale and cross-level processes, feedbacks, and thresholds.

On the evolution of fish-coral interactions.

The biodiversity of tropical reefs is typified by the interaction between fishes and corals. Despite the importance of this ecological association, coevolutionary patterns between these two animal groups have yet to be critically evaluated. After compiling a large dataset on the prevalence of fish-coral interactions, we found that only a minority of fish species associate strongly with live corals (~5%). Furthermore, we reveal an evolutionary decoupling between fish and coral lineage trajectories. While fish lineages expanded in the Miocene, the bulk of coral diversification occurred in the Pliocene/Pleistocene. Most importantly, we found that coral association did not drive major differences in fish diversification. These results suggest that the Miocene fish diversification is more likely related to the development of novel, wave-resistant reef structures and their associated ecological opportunities. Macroevolutionary patterns in reef fishes are thus more strongly correlated with the expansion of reefs than with the corals themselves.

The evolution of fast-growing coral reef fishes.

Individual growth is a fundamental life history trait1-4, yet its macroevolutionary trajectories have rarely been investigated for entire animal assemblages. Here we analyse the evolution of growth in a highly diverse vertebrate assemblage-coral reef fishes. We combine state-of-the-art extreme gradient boosted regression trees with phylogenetic comparative methods to detect the timing, number, location and magnitude of shifts in the adaptive regime of somatic growth. We also explored the evolution of the allometric relationship between body size and growth. Our results show that the evolution of fast growth trajectories in reef fishes has been considerably more common than the evolution of slow growth trajectories. Many reef fish lineages shifted towards faster growth and smaller body size evolutionary optima in the Eocene (56-33.9 million years ago), pointing to a major expansion of life history strategies in this Epoch. Of all lineages examined, the small-bodied, high-turnover cryptobenthic fishes shifted most towards extremely high growth optima, even after accounting for body size allometry. These results suggest that the high global temperatures of the Eocene5 and subsequent habitat reconfigurations6 might have been critical for the rise and retention of the highly productive, high-turnover fish faunas that characterize modern coral reef ecosystems.

Limited cross-species virus transmission in a spatially restricted coral reef fish community.

The Great Barrier Reef (GBR)-the largest coral reef ecosystem in the world-supports over 1,200 fish species with some of the highest population densities and diversities observed in vertebrates, offering a high potential for virus transmission among species. As such, the GBR represents an exceptional natural ecosystem to determine the impact of host community diversity on virus evolution and emergence. In recent decades, the GBR has also experienced significant threats of extinction, making it one of the most vulnerable ecosystems on the planet. Despite the global importance of the GBR, our understanding of virus diversity and connectivity in tropical reef fishes remains poor. Here, we employed metatranscriptomic sequencing to reveal the viromes of sixty-one reef fish species. This identified transcripts representing 132 putative viral sequences, 38 of which exhibited strong phylogenetic relationships with known vertebrate-associated viral genera, including a novel Santee-Cooper ranavirus (Iridoviridae). We found little evidence for virus transmission between fish species living within a very restricted geographical space-a 100-m2 coral reef ecosystem-suggesting that there might be important host barriers to successful cross-species transmission despite regular exposure. We also identified differences in virome composition among reef fish families, such that cryptobenthic reef fishes-characterized by small body sizes and short life spans-exhibited greater virome richness compared to large reef fishes. This study suggests that there are important barriers to cross-species virus transmission and that successful emergence in a reef fish community likely requires active host adaptation, even among closely related host species.

Broad-scale analysis of fish community data suggests critical need to support regional connectivity of coral reefs.

Connectivity is vital for the biodiversity and functioning of marine ecosystems. It is known to be important for coral reefs, but the scales at which connectivity effects matter-and, correspondingly, the scales at which management responses are needed-are poorly understood in marine systems. We used 23 years of fish monitoring data collected from ~50 different coral reefs by the Australian Institute of Marine Science, together with a range of geographic data layers (including the Allen Coral Atlas) and additional network analysis, to explore the balance of local and regional influence on fish communities. Variance partitioning indicated that 42% of the variance in fish community composition could be explained by regional effects or their interaction with coarse-grained local influences (habitat). The variance explained by regional influences was divided evenly between measures that capture location on environmental gradients (e.g., proximity to coastal shelf, latitude) and cross-scale centrality measures of reef location within a broader reef network. A total of 11% of variance could be directly or indirectly attributed to management. Our results provide clear evidence that management and restoration of reefs across the globe must consider both local and regional influences on reef-associated organisms and highlight the potential benefits of improving connectivity in human-dominated coastal seascapes.

On the Challenges of Identifying Benthic Dominance on Anthropocene Coral Reefs.

The concept of dominance is frequently used to describe changes in rapidly reconfiguring ecosystems, but the definition of dominance can vary widely among studies. Using coral reefs as a model, we use extensive benthic composition data to explore how variability in applying dominance concepts can shape perceptions. We reveal that coral dominance is sensitive to the exclusion of key algal groups and the categorization of other benthic groups, with ramifications for detecting an ecosystem phase shift. For example, ignoring algal turf inflates the dominance of hard and soft corals in the benthic habitats underpinning reef ecosystems. We need a consensus on how dominance concepts are applied so that we can build a more comprehensive understanding of ecosystem shifts across a broad range of aquatic and terrestrial settings. For reefs, we highlight the benefits of comprehensive and inclusive surveys for evaluating and managing the altered ecosystem states that are emerging in the Anthropocene.

Benthic composition changes on coral reefs at global scales.

Globally, ecosystems are being reconfigured by a range of intensifying human-induced stressors. Coral reefs are at the forefront of this environmental transformation, and if we are to secure their key ecosystem functions and services, it is important to understand the likely configuration of future reefs. However, the composition and trajectory of global coral reef benthic communities is currently unclear. Here our global dataset of 24,468 observations spanning 22 years (1997-2018) revealed that particularly marked declines in coral cover occurred in the Western Atlantic and Central Pacific. The data also suggest that high macroalgal cover, widely regarded as the major degraded state on coral reefs, is a phenomenon largely restricted to the Western Atlantic. At a global scale, the raw data suggest decreased average (± standard error of the mean) hard coral cover from 36 ± 1.4% to 19 ± 0.4% (during a period delineated by the first global coral bleaching event (1998) until the end of the most recent event (2017)) was largely associated with increased low-lying algal cover such as algal turfs and crustose coralline algae. Enhanced understanding of reef change, typified by decreased hard coral cover and increased cover of low-lying algal communities, will be key to managing Anthropocene coral reefs.

To harness traits for ecology, let's abandon 'functionality'.

Traits are measurable features of organisms. Functional traits aspire to more. They quantify an organism's ecology and, ultimately, predict ecosystem functions based on local communities. Such predictions are useful, but only if 'functional' really means 'ecologically relevant'. Unfortunately, many 'functional' traits seem to be characterized primarily by availability and implied importance - not by their ecological information content. Better traits are needed, but a prevailing trend is to 'functionalize' existing traits. The key may be to invert the process, that is, to identify functions of interest first and then identify traits as quantifiable proxies. We propose two distinct, yet complementary, perspectives on traits and provide a 'taxonomy of traits', a conceptual compass to navigate the diverse applications of traits in ecology.

Small predators dominate fish predation in coral reef communities.

Ecosystem processes are challenging to quantify at a community level, particularly within complex ecosystems (e.g., rainforests, coral reefs). Predation is one of the most important types of species interactions, determining several ecosystem processes. However, while it is widely recognised, it is rarely quantified, especially in aquatic systems. To address these issues, we model predation on fish by fish, in a hyperdiverse coral reef community. We show that body sizes previously examined in fish-fish predation studies (based on a metanalysis), only represent about 5% of likely predation events. The average fish predator on coral reefs is just 3.65 cm; the average fish prey just 1.5 cm. These results call for a shift in the way we view fish predation and its ability to shape the species or functional composition of coral reef fish communities. Considered from a functional group approach, we found general agreement in the distribution of simulated and observed predation events, among both predator and prey functional groups. Predation on coral reefs is a process driven by small fish, most of which are neither seen nor quantified.

A snapshot of sediment dynamics on an inshore coral reef.

Sediments are ubiquitous on coral reefs. However, studies of reef sediments have largely focused on isolated reservoirs, or processes, and rarely consider hydrodynamic drivers. We therefore provide a quantitative snapshot of sediment dynamics on a coral reef. Across a depth profile, we simultaneously examined: suspended sediments, sediment deposition and accumulation, and hydrodynamic and biological movement processes. We reveal the marked potential for the water column to deliver sediments. Currents carried 12.6 t of sediment over the 2,314 m2 study area in 6 days. Sediment traps suggested that a surprisingly high percentage of this sediment was potentially deposited (5.2%). Furthermore, wave-driven resuspension and reworking by parrotfishes separated a highly dynamic sediment regime on the shallow reef flat (3 m), from a more stagnant reef slope (4.5 m-12 m). This study provides a comprehensive model of how hydrodynamic forces and on-reef processes may shape sediment dynamics on a coral reef.

The role of nocturnal fishes on coral reefs: A quantitative functional evaluation.

The ecological functions of nocturnal coral reef fishes are poorly known. Yet, nocturnal resources for coral reef consumers are theoretically as abundant and productive, if not more so, than their diurnal counterparts. In this study, we quantify and contrast the energetic dynamics of nocturnal and diurnal fishes in a model coral reef ecosystem, evaluating whether they attain similar levels of biomass production. We integrated a detailed dataset of coral reef fish counts, comprising diurnal and nocturnal species, in sites sheltered and exposed to wave action. We combined somatic growth and mortality models to estimate rates of consumer biomass production, a key ecosystem function. We found that diurnal fish assemblages have a higher biomass than nocturnal fishes: 104% more in sheltered sites and 271% more in exposed sites. Differences in productivity were even more pronounced, with diurnal fishes contributing 163% more productivity in sheltered locations, and 558% more in exposed locations. Apogonidae dominated biomass production within the nocturnal fish assemblage, comprising 54% of total nocturnal fish productivity, which is proportionally more than any diurnal fish family. The substantially lower contributions of nocturnal fishes to biomass and biomass production likely indicate constraints on resource accessibility. Taxa that overcome these constraints may thrive, as evidenced by apogonids. This study highlights the importance of nocturnal fishes in underpinning the flow of energy and nutrients from nocturnal resources to reef communities; a process driven mainly by small, cryptic fishes.

Occurrence and accumulation of heavy metals in algal turf particulates and sediments on coral reefs.

Algal turfs form a critical interface on coral reefs that interacts with several key ecosystem processes. While we know these turfs have a remarkable propensity to accumulate sediments, which can have a range of ecosystem impacts, their role as sinks for heavy metals remains largely unexamined. Here we quantified the concentration of 15 metals in algal turf sediments from Lizard Island and Orpheus Island on the Great Barrier Reef, and specifically explored how the loads of arsenic, cobalt, iron and lead were related to turf length. Metal composition differed markedly between the two islands, with the composition at Orpheus Island suggesting closer links to terrestrial sediment sources. Furthermore, metal loads increased significantly with turf length, suggesting that longer turfs can accumulate these pollutants on reefs. Given that algal turfs are a crucial component of herbivorous/detritivorous trophic pathways, this could represent a key juncture at which these metals enter food chains.