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Biologically mediated sediment fluxes on coral reefs: sediment removal and off-reef transportation by the surgeonfish 'Ctenochaetus striatus'

2010, Goatley, Christopher, Bellwood, David R

Off-reef sediment transport by the surgeonfish 'Ctenochaetus striatus' (Acanthuridae) was quantified on the reef crest at Lizard Island, Great Barrier Reef. Three independent methods were implemented to estimate sediment ingestion rates. These considered (1) the bite rate and bite volume, (2) the defecation rate and faecal pellet size, and (3) the average gut contents and throughput rate. The 3 methods provided a broad range of estimates of sediment ingestion from 8.8 ± 2.4, to 66.1 ± 14.4 g fish-1 d-1 (mean ± SE). Nevertheless, these estimates were comparable to rates of sediment ingestion by parrotfishes (Labridae), the other major sediment-moving group on reefs. Overall, 36.5% of all sediment ingested was transported from the upper reef crest into deeper water, equating to a removal rate of 28.6 ± 6.2 kg 100 m-2 yr-1 at the study site. By brushing the reef, 'C. striatus' reduces the sediment loading in the epilithic algal matrix (EAM) while causing little damage to the algal turf. Reducing sediments in EAMs provides favourable settlement surfaces for benthic organisms and increases the palatability of the EAM to herbivorous reef fishes, thus supporting reef resilience. The ecological importance of 'C. striatus', which is abundant on reefs throughout the IndoPacific, appears to have been underestimated, particularly when considering reef sediment dynamics.

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The Rise of Jaw Protrusion in Spiny-Rayed Fishes Closes the Gap on Elusive Prey

2015, Bellwood, David R, Goatley, Christopher, Bellwood, Orpha, Delbarre, Daniel J, Friedman, Matt

Jaw protrusion is one of the most important innovations in vertebrate feeding over the last 400 million years [1, 2]. Protrusion enables a fish to rapidly decrease the distance between itself and its prey [2, 3]. We assessed the evolution and functional implications of jaw protrusion in teleost fish assemblages from shallow coastal seas since the Cretaceous. By examining extant teleost fishes, we identified a robust morphological predictor of jaw protrusion that enabled us to predict the extent of jaw protrusion in fossil fishes. Our analyses revealed increases in both average and maximum jaw protrusion over the last 100 million years, with a progressive increase in the potential impact of fish predation on elusive prey. Over this period, the increase in jaw protrusion was initially driven by a taxonomic restructuring of fish assemblages, with an increase in the proportion of spiny-rayed fishes (Acanthomorpha), followed by an increase in the extent of protrusion within this clade. By increasing the ability of fishes to catch elusive prey [2, 4], jaw protrusion is likely to have fundamentally changed the nature of predator-prey interactions and may have contributed to the success of the spiny-rayed fishes, the dominant fish clade in modern oceans [5].

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Clarifying functional roles: algal removal by the surgeonfishes Ctenochaetus striatus and Acanthurus nigrofuscus

2017, Tebbett, Sterling B, Goatley, Christopher, Bellwood, David R

The lined bristletooth, Ctenochaetus striatus, and the brown surgeonfish, Acanthurus nigrofuscus, are among the most abundant surgeonfishes on Indo-Pacific coral reefs. Yet, the functional role of these species has been the focus of an ongoing debate lasting at least six decades. Specifically, to what extent are C. striatus herbivorous like the visually similar A. nigrofuscus? To address this question, we used natural feeding surfaces, covered with late successional stage reef-grown algal turfs, to examine turf algal removal by the two species. Surfaces exposed to C. striatus in laboratory experiments exhibited no significant reductions in turf length or area covered by turfing algae. In marked contrast, A. nigrofuscus reduced turf length by 51% and area covered by turfing algae by 15% in 1 h. The gut contents of specimens from the reef revealed that A. nigrofuscus predominantly ingests algae (the dominant item in 79.6–94.7% of gut content quadrats), while C. striatus ingests detritus and sediments (dominant in 99.6–100% of quadrats). Therefore, C. striatus ingests detritus and sediment, leaving mature algal turfs relatively intact, while A. nigrofuscus directly removes and ingests turf algae. The function of C. striatus differs from cropping herbivorous surgeonfishes such as A. nigrofuscus. On coral reefs, C. striatus brush detrital aggregates from algal turfs, removing microorganisms, organic detritus and inorganic sediment. Confusion over the functional role of C. striatus may stem from an inability to fit it into a single functional category.

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Morphological structure in a reef fish assemblage

2009, Goatley, Christopher, Bellwood, D R

Two key morphological traits, horizontal gape and eye diameter, were measured in a large representative group of coral reef fishes. These morphological traits were used concurrently to assess their utility in exploring abilities of coral reef fishes at an assemblage level. A total of 1,218 specimens from 181 species found on the Great Barrier Reef were examined. Cryptobenthic fishes were included to provide a broader representation of reef fish groups. In the analyses, a clear morphological distinction was found between nocturnal and diurnal fishes. Nocturnal fishes had larger relative horizontal gapes and relative eye diameters by factors of 1.6 and 1.5, respectively. A bivariate plot separated into quadrants was used to assess the implications of morphological variation. The morphological measures reflected distinct ecological traits in each quadrant. Whilst nocturnal fishes had large relative gapes and eye diameters, diurnal predators and detritivores had the same wide gapes, but small relative eye diameters. Highly selective, visual feeders such as the Chaetodontidae and Pseudochromidae had large eyes and small gapes, whilst non-selective feeders with low visual dependence such as the grazing herbivores (Acanthuridae, Siganidae, etc.) had both small eye diameters and gape sizes. The analysis proved to be robust enough to apply to a wide assemblage, but with enough subtlety to distinguish morphological differences within individual families. The methods used in this study may have broad applications to other fish assemblages, both fossil and extant.

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The role of fishes as food: A functional perspective on predator-prey interactions

2021-05, Mihalitsis, Michalis, Hemingson, Christopher R, Goatley, Christopher H R, Bellwood, David R

1. Every animal dies. In nature, mortality usually occurs due to predation by other animals. One of the fundamental consequences of mortality is the transfer of energy and nutrients from one organism (prey) to another (predator). On coral reefs, these key interactions and processes, that are essential for ecosystem functioning, are primarily mediated by fishes; up to 53% of fishes on coral reefs can be regarded as piscivorous.

2. To date, piscivory on coral reefs has been primarily studied with regard to the species piscivores feed on, and how piscivores control populations. Consequently, understanding prey selectivity by piscivorous fishes has been a major goal. However, prey functional traits may also be important in understanding these ‘energy transactions’, especially in complex ecosystems such as coral reefs.

3. Our goal, therefore, was to quantify-at a community level-functional traits of prey that have been shown to influence predator-prey interactions. We found that, on average, deep‐bodied, social fishes occupy higher positions in the water column, whereas solitary species are usually elongate and more closely associated with the benthos. On closer examination, we found that solitary species have a size‐dependent relationship, with substratum associations shifting to water column associations, at approximately 50 mm body length.

4. Our results reveal three distinct prey functional groups: cryptobenthic substratum dwellers, solitary epibenthics and social fishes. These groups display significant differences in their morphologies and behaviours. Furthermore, based on a meta‐analysis of published mortality rates of small‐bodied (<100 mm TL) reef fishes, we show that the three groups display different mortality rates, possibly due to differential exposure to, and potential to be captured by, different predator types.

5. Although fishes are widely available on coral reefs, they may not be equally available as prey to all piscivore types. Prey are not simply victims; they are capable of influencing potential predation through functional traits.

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Low-quality sediments deter grazing by the parrotfish 'Scarus rivulatus' on inner-shelf reefs

2016, Gordon, Sophie E, Goatley, Christopher, Bellwood, David R

Elevated sediment loads in the epilithic algal matrix (EAM) deter grazing by herbivorous fishes and may compromise their critical roles on coral reefs. However, the properties of sediments that drive herbivore deterrence are unknown. Binary choice trials in aquaria were used to examine the effects of three sediment attributes-sediment source, grain size and organic load-on grazing by the abundant inner-shelf parrotfish, Scarus rivulatus. Fish were presented with a choice between EAM-covered rocks treated with (a) terrigenous or reefal sediments, (b) fine or coarse sediments or (c) sediments with high or low organic loads. Scarus rivulatus did not show a preference for sediments from different sources (terrigenous vs. reefal); however, a clear preference was evident for fine-grained sediments over coarse (109 % more bites) and sediments with high organic loads over low (147 % more bites). The avoidance of coarse sediments is likely to be a key factor driving the inhibition of grazing on mid-shelf reefs, which are dominated by coarse sediments. In contrast, on innershelf reefs, grazing by parrotfishes may be deterred primarily by high sediment loads, which reduce the proportional organic content in EAM sediments. Our study highlights the potential impact of sediments on critical ecological processes and the threats posed by changing sediment loads on inner-shelf reefs.

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Evolution of long-toothed fishes and the changing nature of fish-benthos interactions on coral reefs

2014, Bellwood, David R, Hoey, Andrew S, Bellwood, Orpha, Goatley, Christopher

Interactions between fishes and the benthos have shaped the development of marine ecosystems since at least the early Mesozoic. Here, using the morphology of fish teeth as an indicator of feeding abilities, we quantify changes over the last 240 million years of reef fish evolution. Fossil and extant coral reef fish assemblages reveal exceptional stasis in tooth design over time, with one notable exception, a distinct long-toothed form. Arising only in the last 40 million years, these long-toothed fishes have bypassed the invertebrate link in the food chain, feeding directly on benthic particulate material. With the appearance of elongated teeth, these specialized detritivores have moved from eating invertebrates to eating the food of invertebrates. Over evolutionary time, fishes have slid back down the food chain.

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Fishes on coral reefs: changing roles over the past 240 million years

2010, Goatley, Christopher, Bellwood, David R, Bellwood, Orpha

Key morphological traits reveal changes in functional morphospace occupation of reef fish assemblages over time. We used measurements of key functional attributes (i.e., lower jaw length and orbit diameter) of 208 fossil fish species from five geological periods to create bivariate plots of functional morphological traits through time. These plots were used to examine possible function and ecological characteristics of fossil reef fish assemblages throughout the Mesozoic and Cenozoic. A previously unknown trend of increasing orbit diameter over time became apparent. The Teleostei are the principal drivers of this change. The Eocene appears to mark a dramatic increase in two previously rare feeding modes in fishes: nocturnal feeding and high-precision benthic feeding. Interestingly, members of the Pycnodontiformes had relatively large eyes since the Triassic and appear to be the ecological precursors of their later teleost counterparts and may have been among the earliest nocturnal feeding fishes. Our results highlight potential changes in the roles of fishes on coral reefs through time.

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Reef-scale partitioning of cryptobenthic fish assemblages across the Great Barrier Reef, Australia

2016, Goatley, Christopher, Gonzalez-Cabello, Alonso, Bellwood, David R

Onshore to offshore gradients in marine assemblages have been well documented on coral reefs, with most studies showing a distinct separation between onshore and offshore locations. Here we use enclosed anaesthetic sampling of small, cryptobenthic reef fishes to assess changes in assemblage composition across the Great Barrier Reef continental shelf. The cryptobenthic fishes exhibited fine-scale partitioning across the shelf. Three dominant species of goby accounted for over 55% of all fishes collected, with 1 species characterising each of the 3 key shelf positions: inner-, mid- and outer-shelf. Multivariate analyses of assemblage composition revealed further separation of reefs within the inner- and mid-shelf positions, highlighting the exceptional sensitivity of cryptobenthic reef fish assemblages to shelf position, with a progressive separation of individual reef assemblages with distance from the shore. These among-reef patterns contrast markedly with other reef fish taxa which invariably show 2 broad assemblages across the continental shelf (inner- vs. a composite mid- and outer-shelf community). As a result of this exceptional sensitivity to environmental conditions, cryptobenthic reef fish communities may represent good subjects for high-resolution monitoring of disturbances on coral reefs.

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

2018-01-31, Tebbett, Sterling B, Goatley, Christopher H R, Huertas, Víctor, Mihalitsis, Michalis, Bellwood, David R

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.