HomeBiologySmall predators dominate fish predation in coral reef communities

Small predators dominate fish predation in coral reef communities


For a lot of animals and vegetation in excessive variety programs, akin to coral reefs, inhabitants dynamics are pushed by youth stage mortality, with recruitment functioning as a inhabitants bottleneck [13]. Certainly, predation-based mortality is extensively thought to be some of the essential species interactions figuring out fish inhabitants constructions on coral reefs [46]. On reefs, many, if not most fishes, are eaten by different fishes (7). Whereas the results of excessive mortality in these ecosystems is effectively documented [8,9], the predators that drive this course of are largely unknown.

In the previous few years, the principle focus of fish predation research on coral reefs has been on trophic cascades [1013], its results on prey abundance [14,15], or oblique “fear-effects” on prey behaviour [1619]. The fish predators investigated in these fields have been primarily sharks or different giant mesopredators. Nevertheless, latest work has highlighted the potential trophic significance of small-bodied coral reef fishes [20,21]. This raises questions over the identification and dimension of fish predators and their relative significance from an ecosystem perform perspective. Principally, huge questions nonetheless stay: Who’re the principle predators of fishes on coral reefs? At which dimension does most predation occur? And, whereas all people in a group undergo the predation gauntlet, how do these interactions scale at a group degree?

Certainly, quantifying fish predation at a group degree poses some challenges. Typically, fish–fish predation has been quantified via intestine content material analyses [7,22,23]. Nevertheless, this evaluation requires a excessive variety of specimens for a small yield of information, as most frequently, piscivore guts are empty [24,25]. Whereas fish–fish predation is extensively acknowledged, it’s logistically difficult to quantify in situ, given the period of those occasions solely lasting few milliseconds [26,27]. Moreover, many predator–prey estimates are based mostly on size–size relationships. Nevertheless, particularly inside coral reef fishes, physique depth (and due to this fact a determinant of gape limitation [28]) is a major axis of variation [29,30]; assuming predator–prey relationships based mostly on size, dangers the lack of vital variation within the knowledge. Lastly, approaches incorporating species abundance alone, might result in the lack of info on the ontogenetic historical past of fishes (for instance, a juvenile predator can also be prey throughout its youth phases), or, to incorrect estimates of interplay charges (i.e., presence doesn’t at all times equate to perform) [31]. Subsequently, for the explanations above, there’s a must quantify predation occasions at a group degree, utilizing a special method.

Given the latest advances in, and promising outcomes from, a practical group method [32] to the investigation of ecosystem processes [3335], we apply this method to coral reef fishes, with a selected give attention to predation on fishes, by fishes. We first surveyed a coral reef fish group and constructed an algorithm to mannequin predator–prey interactions based mostly on the practical constraints imposed by each predators and prey (following [36,37]). Predators have been labeled by their practical group (grabbers versus engulfers) and dimension, whereas prey have been labeled by physique depth (which determines the dimensions of predators in a position to feed on them) and prey practical group (cryptobenthic, epibenthic, social) [36]. This produced a standardised surveyed group of 32,218 fish that have been simulated 1 million occasions to supply 349,000 potential predation occasions (i.e., functionally viable occasions, by eradicating forbidden interactions [38,39]). These outcomes have been then in comparison with the documented consumption of reef fish prey by fish predators, based mostly on a metanalysis of intestine content material knowledge (n = 1,677 predation occasions) throughout Indo-Pacific coral reef ecosystems.


We discovered that fish predation on coral reefs is overwhelmingly dominated by small, diminutive predators. The typical fish predator that feeds on different fish on reefs is simply 3.6 cm, and the common prey simply 1.5 cm. By combining surveys at completely different spatial scales, to generate as full a census as potential, our standardised surveyed reef fish group contained 32,218 fishes from 266 species. Simulating 1 million potential predator–prey interactions (i.e., predation occasions) inside this group, by making use of size-based practical constraints (prey physique depth/predator gape dimension), we obtained 349,081 potential (i.e., functionally possible) predation occasions. On this intensive pool of potential occasions, the median dimension of a predator fish was simply 3.65 cm (95% CI: 2.38 to fifteen) complete size (TL) (imply: 5.6 cm) (Fig 1A). Basically, 95% of potential predation occasions contain predators lower than 15 cm. When simulated predation occasions from our surveyed group have been in comparison with a literature-based dataset of 1,677 noticed predation occasions, by dimension, there was solely 5% overlap (Fig 1B). In essence, the overwhelming majority of research have completely quantified predation by exceptionally giant predators; most predation occasions go unobserved and unrecorded. Our outcomes counsel there’s a want for a shift in the way in which we think about fish predation, and the way trophic interactions form the species, and practical, composition of coral reef fish communities.


Fig 1. Cryptopredators, not “giant” reef fishes, dominate predation occasions on coral reefs.

(A) Neighborhood-level predation of coral reef fishes, alongside a predator dimension gradient, based mostly on simulations from reef surveys. Examples of cryptopredators that seemingly form group composition in coral reef ecosystems, prime to backside: Pseudochromis cyanotaenia, Cypho purpurascens, Plesiops sp. (B) Predation occasions simulated from our surveyed group (identical as (A)) vs. noticed predation occasions (metanalysis of literature). A number of traces in noticed predation occasions replicate attracts from a distribution (see Supplies and strategies). (C) Abundance estimates of cryptopredator species, relative to juveniles of “giant” reef fish predator species. The colored field represents the dimensions vary inside which most predation occasions are estimated. Authentic pictures for clipart in (A) have been initially sourced from C.R. Hemingson, with permission. Authentic photographs for clipart in (B) have been taken by J.E. Randall and sourced from [40]. The info underlying this determine could be present in DOI: 10.5281/zenodo.6772154.


Whereas exponentially declining mortality charges seem like the norm for reef fishes [8,9,41], the predators that drive these curves have remained largely unknown. Our outcomes emphasize the small dimension of those predators. Moreover, based mostly on abundance-based encounter likelihoods, these predators are unlikely to be juveniles of “giant” reef fish predators; the predators driving the method are predominantly cryptopredators (Fig 1C), outlined herein as carnivorous fishes under 10 cm.

The identical patterns apply to prey fish. The estimated median dimension of prey fish was simply 1.5 cm TL (95% CI: 0.8 to three.65) (imply: 1.75 cm) (Fig 2A); 95% of potential predation occasions contain prey sizes lower than 3.65 cm. Functionally possible predation occasions have been simulated based mostly on prey physique depth versus gape dimension relationships. These simulations resulted in 349,081 functionally possible predation occasions. Of those occasions, the prey concerned have been: 90.4% cryptobenthic substratum dwellers (referred to hereafter as cryptobenthic), 8.4% social prey, and 1.2% solitary epibenthic (known as epibenthic) (Fig 2B) (for particulars on practical teams, see S1 Desk). When these predation occasions are in comparison with values of printed reef fish mortality charges, their distribution matches carefully (Fig 2).


Fig 2. Cryptobenthic fishes are the principle prey fishes on coral reefs.

(A) Neighborhood degree predation simulated in our examine alongside a prey dimension gradient (brown), mirrors the exponentially declining line that represents noticed reef fish mortality charges from an impartial metanalysis on reef fish mortality [8]. Examples of major contributors to this density distribution, from prime to backside: Eviota queenslandica, Salarias alboguttatus, Enneapterygius tutuilae. (B) The identical density curve, in 2A, break up based on prey practical teams, specifically: cryptobenthic = crimson, epibenthic = yellow, social = blue. Authentic pictures for clipart in (A) have been initially sourced from C.R. Hemingson, with permission. The info underlying this determine could be present in DOI: 10.5281/zenodo.6772154.


Evaluating size-specific simulated predation to noticed predation occasions at a group degree

The distribution of potential predation occasions amongst completely different dimension lessons of predators in our simulated group, was discovered to carefully replicate the distributions in our metanalysis. This utilized to each predator practical teams (grabbers and engulfers) (Fig 3A and 3B) and all 3 prey practical teams (Fig 3C–3E), aside from small predator physique sizes. In essence, for small predators (i.e., ≤20 to 25 cm), cryptobenthic prey are underrepresented (Fig 3E), whereas epibenthic and social prey are overrepresented (Fig 3C–3E). This can be linked to the practical traits of those prey teams and the predators concerned on this predation.


Fig 3. Purposeful teams of predator–prey fishes reveal patterns of community-level predation.

Simulated vs. noticed relative contributions to the method of piscivory on reefs, based mostly on predator practical teams (A, B) and prey practical teams (C–E). Total, the trajectories of simulated contributions (dashed traces) and noticed contributions (strong traces) have been in settlement for predator and prey practical teams. Disparity was solely discovered between the two estimates for small predator sizes, when outcomes are based mostly on prey practical teams: Social and epibenthic prey have been overrepresented within the food regimen of small predators, whereas cryptobenthic prey have been underrepresented. Stuffed circles point out means, whereas vertical bars point out the vary of values (minimal, most) for a selected dimension bin. Gray zones point out dimension bins the place a distinction between predicted and noticed relative contribution was discovered to be vital. Fish silhouettes redrawn from [36] and [37]. The info underlying this determine could be present in DOI: 10.5281/zenodo.6772154.



Cryptopredators have solely just lately been recognized as vital fish shoppers [21]. Their overwhelming abundance (relative to juveniles of enormous reef fish predators), together with our outcomes of simulated community-level predation, spotlight the potential of those beforehand neglected cryptopredators to be the first contributors to the method of fish predation in coral reef ecosystems. Frequent examples of cryptopredators (cryptobenthic reef fishes sensu Brandl, Goatley [42] that are carnivorous) embody the Pseudochromidae, Plesiopidae, Gobiidae, and Apogonidae. Many of the species inside these households stay beneath 15 cm all through their lives [42].

The fishes consumed by cryptopredators, based mostly on our simulations, are overwhelmingly cryptobenthic (approx. 90% of predation occasions) (Fig 2). Our outcomes mirror earlier empirical research exhibiting a excessive consumption of, and excessive turnover in, cryptobenthic fishes; a “crypto-pump,” fuelling coral reef ecosystems [20]. In essence, our outcomes illuminate the “dark-productivity” (sensu 20) that fuels coral reefs, by figuring out their most probably predators. These fishes maintain a few of the most essential trophic pathways on coral reefs (e.g., the detrital and piscivory-cryptobenthic pump). This will likely add to the number of mechanisms of power recycling, which seem like a necessary attribute of oligotrophic ecosystems with excessive species variety and biomass [43,44]. We present that one of many key pathways that hyperlinks these fishes to the remainder of the food-chain is thru cryptopredators.

Though the underestimation of cryptobenthic prey within the food regimen of predators could also be related to methodological challenges akin to excessive digestion charges and visible identification of prey in guts [20,45], we propose that there may additionally be underlying causes related to the precise options of this prey practical group (cryptobenthic prey). There’s overwhelming proof, that almost all mortality in fishes happens throughout the youth phases at small physique sizes, and that this is because of predation [6,8,46,47]. That is at odds with the life historical past of cryptobenthic fishes. How can cryptobenthic fishes, the shortest residing vertebrates [48,49], with extraordinarily small physique sizes and intensely excessive mortality charges, preserve viable populations? Sustained temporal replica [50], quick development [51], and considerable larvae [20] definitely all assist to facilitate the acute cryptobenthic way of life. Nevertheless, our knowledge strongly counsel that their success may additionally be depending on their potential to scale back relative predation threat under what could be anticipated based mostly on their dimension alone. By lowering predation, they might be higher in a position to unfold predation-based mortality all through their life on the reef, sustaining a better variety of reproducing people throughout this weak interval (S2 Fig). These advantages could also be straight associated to the traits of the cryptobenthic practical prey group such because the behaviour of “sitting” on the benthos [36] and cryptic colouration [52]. The drab colouration and “sitting nonetheless” might certainly be a extremely profitable antipredatory technique.

The outcomes mentioned above, solely grew to become evident when investigating the group from a practical group perspective. Our simulation method permits us to make estimates on the sizes of predators and prey concerned, in addition to the potential relationships between the predator and prey teams. Such outcomes have key implications for our understanding and function of practical teams inside their habitat, one thing which has been proven for different ecosystems as effectively [5355]. Certainly, we present that the practical group method is a strong device in elucidating the complexities of hyperdiverse programs akin to coral reefs [5658]. This practical method might certainly, clarify how cryptobenthic prey are in a position to exist in any case.

You will need to be aware that different elements of coral reef environments (e.g., habitat complexity, publicity) [5961] might form predator–prey interactions for cryptopredators. Particularly, the fractal dimension/scale [62] at which these predation occasions happen are seemingly not the identical as for giant predatory fishes. For giant predatory fishes (e.g., Lutjanidae or Serranidae), the presence of enormous coral constructions are more likely to decide the stability between escape and susceptibility to predation [63]. Nevertheless, for predation by small-bodied and inconspicuous predators (i.e., cryptopredation), it’s seemingly that coral rubble (or small corals), as an alternative, would be the constructions creating the identical benefits or disadvantages, as giant coral constructions for bigger predators. Earlier research have discovered a excessive abundance of cryptobenthic fishes in rubble areas (versus coral-covered areas) [60,61,64], suggesting an intricate matrix happening at a smaller scale, the place habitat might affect predation between cryptopredators and their prey [65]. Our outcomes might assist to place rubble habitats within the context of habitat complexity and the fractal dimension [62] for the fishes residing inside these areas. In essence, there’s a must additional examine the affect of abiotic variables on practical traits referring to cryptopredation.

Total, we present that the overwhelming majority of fish predation occasions on coral reefs is more likely to contain predators under 15 cm. The overwhelming majority of prey in these predation occasions is under 5 cm. “Typical” predators on reefs, akin to jacks, barracudas, and groupers, are usually not those finishing up most predation on reefs. Most fish are eaten by cryptopredators on the reef. We spotlight the overwhelming significance of cryptopredators as drivers of predation at a group degree. Moreover, our knowledge means that, opposite to expectations, a small physique dimension might certainly perform as an antipredatory mechanism whether it is related to “sitting” on the benthos. Our practical teams method revealed that predation occasions are additionally ruled by prey practical traits. Total, predation on coral reefs is a sport of small fishes, and cryptobenthic prey fishes seem like profitable the sport.

Supplies and strategies

Quantifying ecosystem processes at a group degree is a logistically tough and time-consuming course of. Often, species interactions are inferred based mostly on collected empirical knowledge, most frequently recorded as presence/absence or abundance knowledge, or via simulation-based approaches. Right here, we evaluate and distinction these 2.

Quantifying predator abundance and prey availability at a group degree

We first surveyed a coral reef fish group at Lizard Island, a marine reserve with no fishing, positioned on the Nice Barrier Reef, Australia, following [66]. Fish surveys have been performed in all historically recognised reef zones (again, flat, crest, slope). Each underwater visible surveys and enclosed clove oil stations have been used to maximise the proportion of the fish group surveyed. The two approaches have been chosen for various teams of fishes, based mostly on their physique sizes and behaviours [67]. Physique sizes have been estimated by a single diver throughout visible surveys for larger-bodied fishes and have been measured within the laboratory beneath a stereomicroscope for smaller-bodied fishes (see under). Visible surveys have been executed with a diver initially conducting a 50 × 5 m transect tape survey to rely giant (>25 cm TL), water column-positioned or quick swimming fishes more likely to be scared away by the diver. Upon return alongside the tape, the identical diver performed a 30 × 5 m survey focusing on smaller-bodied fishes which are much less cellular. The diver then performed one other 30 × 5 m survey over the identical space to rely small-bodied, non-cryptic fishes often discovered simply above the reef benthos. Lastly, the diver performed a final 30 × 1 m survey to rely cryptic people (e.g., inside or beneath crevices) that will not have been surveyed utilizing conventional visible surveying methods [66]. As well as, to offer extra correct abundance estimates of cryptobenthic reef fishes, a set of 8 enclosed clove oil stations (following [67]) have been deployed in every habitat. A complete of three units of visible surveys and eight clove oil stations have been performed in every reef zone at every of the websites (n = 3 websites). For extra detailed info on sampling strategies, see [66]. Fish surveys have been performed in accordance with the James Prepare dinner College Ethics Committee (A2375) and the Nice Barrier Reef Marine Park Authority (G17/38142.1).

To account for the completely different spatial extents of the completely different surveying strategies, a resampling algorithm was constructed. This allowed scaling the noticed fish abundance to a standardised frequent space amongst surveys. This process generated 1 commonplace, 1,200 m2 group, which could be interpreted as a reef part spanning the completely different reef zones, and with equal space in every of those zones. This “multihabitat” coral reef fish group had 32,235 fish people from 266 species. We then assigned all fishes to their respective prey practical group (based mostly on their practical traits, see S1 Desk) following [36].

Our quantification of potential predation occasions at a group degree, began with a group dataset together with fish species and physique dimension. Based mostly on physique dimension and beforehand printed relationships between physique dimension and practical traits of each predator and prey fishes [36,37], we transformed physique sizes to practical trait values straight associated to predator–prey interactions (i.e., prey physique depth, predator gape dimension). We then performed repeated simulations of potential predation occasions by sampling people (1 predator and 1 prey at a time) inside the group. Solely real looking interactions, based mostly on the practical trait relationships, have been thought-about (e.g., if the prey might match within the predators’ gape), thus taking into consideration the presence of forbidden hyperlinks [38,39]. Following simulations, we in contrast our outcomes of potential predation, to noticed consumption patterns, based mostly on a metanalysis of the intestine contents of coral reef fish predators.

The implementation of a practical group method (based mostly on practical traits) has lowered the preliminary complexity of coral reef piscivores [37,68,69]. Current research have recognized 2 practical teams of predators, grabbers and engulfers, which differ of their morphology, placing, capturing, and prey processing behaviour [37]. Reef fish prey may also be divided into cryptobenthic substratum dwellers (referred to herein as “cryptobenthic”), solitary epibenthic (“epibenthic” herein), and social fishes, which differ in antipredatory morphological and behavioural traits, in addition to in habitat use (e.g., place in water column) [36]. We be aware right here that the time period “cryptobenthic” is barely completely different than that of Brandl, Goatley [42]. For an in depth description of those practical teams, see S1 Desk.

Every particular person fish species was assigned as a predator if that species has been discovered to feed on elusive prey within the literature or different on-line sources (e.g. 40). All fishes have been thought-about as potential prey. Prey physique dimension was then reworked to physique depth based mostly on the practical group of the species following [36].

the place f is 1 of the three practical teams, and i an index denoting a person fish,

af = (−1.58, −0.78, −0.9), bf = (1.04, 0.74, 0.95), for f = (cryptobenthic, epibenthic, social), respectively. L is the full size of a person fish. Predator physique sizes have been then reworked to gape sizes following relationships obtained from coral reef fish specimens [28,70,71], based mostly on the practical group to which the surveyed predator belonged to:

the place f is 1 of the two practical teams, and i an index denoting a person fish,

af = (0.93, 0.04), bf = (0.17,0.17), for f = (engulfers, grabbers), respectively. L is the full size of a person fish.

We then performed a collection of simulations, whereby a person predator from the group (together with its respective practical trait values) was randomly matched in opposition to a person prey fish (together with its respective practical trait values). Every simulation consisted of 10,000 potential piscivory occasions, and the simulation was performed 100 occasions with repetition. We then calculated the connection between predator gape dimension and prey physique depth, for every potential predation occasion (following 28), by dividing prey physique depth to predator gape dimension. We solely stored cases through which the obtained ratios have been inside the vary of 0.14 to 0.7, as this has been discovered to be the relative prey dimension inside which 95% of predation happens [37], and eradicated some other cases (i.e., forbidden hyperlinks [38,39]). Subsequent, we binned these occasions into respective dimension bins of predators (from 5 to 50 cm, at 5 cm intervals), and calculated the relative contribution of every prey practical group to the general prey availability for every predator dimension bin:

the place C is the relative contribution of accessible prey of a selected practical group f at a selected dimension bin b, A is the abundance of people of the precise prey practical group, for the given dimension bin, and the denominator is the summation of the abundances of all n practical teams. These contributions have been then in comparison with the noticed consumption of every prey practical group based mostly on the metanalysis of intestine content material knowledge (see under).

Noticed food regimen of predators (metanalysis)

Food regimen info was collected from printed literature on the intestine contents of piscivorous coral reef teleost fishes within the Indo-Pacific realm (S1 Fig). Data extracted from the literature was: Vary of predator physique sizes sampled from every examine, predator species, prey species, and variety of occurrences that the prey species was present in predator guts. Prey species have been then labeled into practical teams (as above). We eliminated pelagic predators as they seemingly function at a broader spatial scale than extra benthic related predators.

Particular person physique sizes weren’t out there, as solely dimension ranges have been reported within the literature. Subsequently, for each predation occasion recorded from the literature, we drew particular person physique sizes from a uniform distribution delimited by the vary of sizes supplied by the respective examine. This course of was executed for every predation occasion recorded (n = 1,224) and was simulated 1,000 occasions with repetition. In some cases, the precise dimension of the predator was recorded, and was due to this fact used as the one potential physique dimension for the given predation occasions, all through the simulations. These noticed predation occasions have been in comparison with our simulated predation occasions based mostly on the overlap coefficient from the R-package “bayestestR” [72]. These predation occasions have been then assigned into the identical physique dimension bins as those utilized in our group survey (see above). We then summarised the relative contribution of every prey practical group to every predator physique dimension bin (as above).


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