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.

Are fish communities on coral reefs becoming less colourful?

An organism's colouration is often linked to the environment in which it lives. The fishes that inhabit coral reefs are extremely diverse in colouration, but the specific environmental factors that support this extreme diversity remain unclear. Interestingly, much of the aesthetic and intrinsic value humans place on coral reefs (a core ecosystem service they provide) is based on this extreme diversity of colours. However, like many processes on coral reefs, the relationship between colouration and the environment is likely to be impacted by global environmental change. Using a novel community-level measure of fish colouration, as perceived by humans, we explore the potential links between fish community colouration and the environment. We then asked if this relationship is impacted by human-induced environmental disturbances, e.g. mass coral bleaching events, using a community-level dataset spanning 27 years on the Great Barrier Reef. We found that the diversity of colours found within a fish community is directly related to the composition of the local environment. Areas with a higher cover of structurally complex corals contained fish species with more diverse and brighter colourations. Most notably, fish community colouration contracted significantly in the years following the 1998 global coral bleaching event. Fishes with colouration directly appealing to human aesthetics are becoming increasingly rare, with the potential for marked declines in the perceived colour of reef fish communities in the near future. Future reefs may not be the colourful ecosystems we recognize today, representing the loss of a culturally significant ecosystem service.

Feeding kinematics of a surgeonfish reveal novel functions and relationships to reef substrata.

Biting to obtain attached benthic prey characterizes a large number of fish species on coral reefs, and is a feeding mode that contributes to important ecosystem functions. We use high-speed video to reveal the mechanisms used by a surgeonfish, Acanthurus leucosternon, to detach algae. After gripping algae in its jaws, the species pulls it by ventrally rotating both the head and the closed jaws, in a novel use of the intra-mandibular joint. These motions remain in the plane of the fish, reducing the use of a lateral head flick to detach the algae. The novel ability to bite and pull algae off the substrate without bending the body laterally minimizes exposure to high water flows, and may be an adaptation to feeding in challenging reef habitats such as the crest and flat. Therefore, our results could potentially represent a key milestone in the evolutionary history of coral reef trophodynamics.

Forensic odontology: Assessing bite wounds to determine the role of teeth in piscivorous fishes.

Teeth facilitate the acquisition and processing of food in most vertebrates. However, relatively little is known about the functions of the diverse tooth morphologies observed in fishes. Piscivorous fishes (fish-eating fish) are crucial in shaping community structure and rely on their oral teeth to capture and/or process prey. However, how teeth are utilized in capturing and/or processing prey remains unclear. Most studies have determined the function of teeth by assessing morphological traits. The behavior during feeding, however, is seldom quantified. Here, we describe the function of teeth within piscivorous fishes by considering how morphological and behavioral traits interact during prey capture and processing. This was achieved through aquarium-based performance experiments, where prey fish were fed to 12 species of piscivorous fishes. Building on techniques in forensic odontology, we incorporate a novel approach to quantify and categorize bite damage on prey fish that were extracted from the piscivore's stomachs immediately after being ingested. We then assess the significance of morphological and behavioral traits in determining the extent and severity of damage inflicted on prey fish. Results show that engulfing piscivores capture their prey whole and head-first. Grabbing piscivores capture prey tail-first using their teeth, process them using multiple headshakes and bites, before spitting them out, and then re-capturing prey head-first for ingestion. Prey from engulfers sustained minimal damage, whereas prey from grabbers sustained significant damage to the epaxial musculature. Within grabbers, headshakes were significantly associated with more severe damage categories. Headshaking behavior damages the locomotive muscles of prey, presumably to prevent escape. Compared to non-pharyngognaths, pharyngognath piscivores inflict significantly greater damage to prey. Overall, when present, oral jaw teeth appear to be crucial for both prey capture and processing (immobilization) in piscivorous fishes.

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.

Functional groups in piscivorous fishes.

Piscivory is a key ecological function in aquatic ecosystems, mediating energy flow within trophic networks. However, our understanding of the nature of piscivory is limited; we currently lack an empirical assessment of the dynamics of prey capture and how this differs between piscivores. We therefore conducted aquarium-based performance experiments, to test the feeding abilities of 19 piscivorous fish species. We quantified their feeding morphology, striking, capturing, and processing behavior. We identify two major functional groups: grabbers and engulfers. Grabbers are characterized by horizontal, long-distance strikes, capturing their prey tailfirst and subsequently processing their prey using their oral jaw teeth. Engulfers strike from short distances, from high angles above or below their prey, engulfing their prey and swallowing their prey whole. Based on a meta-analysis of 2,209 published in situ predator-prey relationships in marine and freshwater aquatic environments, we show resource partitioning between grabbers and engulfers. Our results provide a functional classification for piscivorous fishes delineating patterns, which transcend habitats, that may help explain size structures in fish communities.

Functional implications of dentition-based morphotypes in piscivorous fishes.

Teeth are crucial in elucidating the life history of vertebrates. However, most studies of teeth have focused on mammals. In heterodont mammals, tooth function is based on tooth shape and position along the jaw. However, the vast majority of vertebrates are homodont, and tooth function might not be based on the same principles (in homodonts, tooth shape is broadly similar along the jaw). We provide a quantitative framework and establish dentition-based morphotypes for piscivorous fishes. We then assess how these morphotypes relate to key functional feeding traits. We identified three broad morphotypes: edentulate, villiform and macrodont, with edentulate and villiform species displaying considerable functional overlap; macrodont species are more distinct. When analysing macrodonts exclusively, we found a major axis of variation between 'front-fanged' and 'back-fanged' species. The functional interpretations of this axis suggest that tooth-based functional decoupling could exist, even in homodont vertebrates, where teeth have similar shapes. This diversity is based not only on tooth shape but also solely on the position along the jaw.

A functional evaluation of feeding in the surgeonfish Ctenochaetus striatus: the role of soft tissues.

Ctenochaetus striatus is one of the most abundant surgeonfishes on Indo-Pacific coral reefs, yet the functional role and feeding ecology of this species remain unclear. This species is reported to possess a rigid structure in its palate that is used for scraping, but some authors have reported that this element is comprised of soft tissue. To resolve the nature and role of this structure in the feeding ecology of C. striatus we examined evidence from anatomical observations, scanning electron microscopy, histology, X-ray micro-computed tomography scanning, high-speed video and field observations. We found that C. striatus from the Great Barrier Reef possess a retention plate (RP) on their palates immediately posterior to the premaxillary teeth which is soft, covered in a thin veneer of keratin with a papillate surface. This RP appears to be used during feeding, but does not appear to be responsible for the removal of material, which is achieved primarily by a fast closure of the lower jaw. We infer that the RP acts primarily as a 'dustpan', in a 'dustpan and brush' feeding mechanism, to facilitate the collection of particulate material from algal turfs.

The role of the reef flat in coral reef trophodynamics: Past, present, and future.

The reef flat is one of the largest and most distinctive habitats on coral reefs, yet its role in reef trophodynamics is poorly understood. Evolutionary evidence suggests that reef flat colonization by grazing fishes was a major innovation that permitted the exploitation of new space and trophic resources. However, the reef flat is hydrodynamically challenging, subject to high predation risks and covered with sediments that inhibit feeding by grazers. To explore these opposing influences, we examine the Great Barrier Reef (GBR) as a model system. We focus on grazing herbivores that directly access algal primary productivity in the epilithic algal matrix (EAM). By assessing abundance, biomass, and potential fish productivity, we explore the potential of the reef flat to support key ecosystem processes and its ability to maintain fisheries yields. On the GBR, the reef flat is, by far, the most important habitat for turf-grazing fishes, supporting an estimated 79% of individuals and 58% of the total biomass of grazing surgeonfishes, parrotfishes, and rabbitfishes. Approximately 59% of all (reef-wide) turf algal productivity is removed by reef flat grazers. The flat also supports approximately 75% of all grazer biomass growth. Our results highlight the evolutionary and ecological benefits of occupying shallow-water habitats (permitting a ninefold population increase). The acquisition of key locomotor and feeding traits has enabled fishes to access the trophic benefits of the reef flat, outweighing the costs imposed by water movement, predation, and sediments. Benthic assemblages on reefs in the future may increasingly resemble those seen on reef flats today, with low coral cover, limited topographic complexity, and extensive EAM. Reef flat grazing fishes may therefore play an increasingly important role in key ecosystem processes and in sustaining future fisheries yields.

A morphological and functional basis for maximum prey size in piscivorous fishes.

Fish predation is important in shaping populations and community structure in aquatic systems. These predator-prey interactions can be influenced by environmental, behavioural and morphological factors. Morphological constraints influence the feeding performance of species, and interspecific differences can thus affect patterns of resource use. For piscivorous fishes that swallow prey whole, feeding performance has traditionally been linked to three key morphological constraints: oral gape, pharyngeal gape, and the cleithral gape. However, other constraints may be important. We therefore examine 18 potential morphological constraints related to prey capture and processing, on four predatory species (Cephalopholis urodeta, Paracirrhites forsteri, Pterois volitans, Lates calcarifer). Aquarium-based experiments were then carried out to determine capture and processing behaviour and maximum prey size in two focal species, C. urodeta and P. forsteri. All four species showed a progressive decrease in gape measurements from anterior to posterior with oral gape ≥ buccal ≥ pharyngeal ≥ pectoral girdle ≥ esophagus ≥ stomach. C. urodeta was able to process prey with a maximum depth of 27% of the predators' standard length; for P. forsteri it was 20%. C. urodeta captured prey head-first in 79% of successful strikes. In P. forsteri head-first was 16.6%, mid-body 44.4%, and tail-first 38.8%. Regardless of capture mode, prey were almost always swallowed head first and horizontally in both focal species. Most internal measurements appeared too small for prey to pass through. This may reflect the compressibility of prey, i.e. their ability to be dorsoventrally compressed during swallowing movements. Despite examining all known potential morphological constraints on prey size, horizontal maxillary oral gape in a mechanically stretched position appears to be the main morphological variable that is likely to affect maximum prey size and resource use by these predatory species.