The predation gauntlet: early post-settlement mortality in reef fishes

Most marine fishes have pelagic larvae that settle to benthic juvenile/adult habitats. Ecologists have argued that mortality rates are particularly high during the settlement transition, but relevant data have been sparse. Recently, researchers have used several novel techniques to estimate the magnitude of predation mortality during the settlement transition. We used meta-analysis to determine that for 24 taxonomically diverse species in geographically widespread locations, an estimated 55.7% (CI: 43.0–65.5%) of juveniles were consumed within 1–2 days of settlement. Such high mortality highlights this brief period as a key phase in the life history of fishes and supports the view that these communities are strongly influenced by predation. Additionally, we argue that because predators have such strong effects on juvenile survival, the population and community dynamics of reef fishes may be linked to human exploitation of reef predators.     

Aggression among sea urchins on Caribbean coral reefs

The existence and nature of intra- and interspecific aggression were examined for five species of sea urchins inhabiting Caribbean coral reefs. The studies took place in the San Blas Islands of Panama and involved the following species: Echinometra lucunter (Linnaeus), E. viridis Agassiz, Diadema antillarum Phillipi, Lytechinus williamsi Chesher, and Eucidaris tribuloides (Lamarck). An intruder was placed next to an undisturbed resident and the behaviors and responses of both individuals were followed. All pairwise combinations of resident species and intruder species were tested except for combinations involving D. antillarum as intruder.

For E. viridis and E. lucunter, agonistic interactions occurred commonly (46–79% of trials) between conspecifics and congeners. Almost all of the agonistic encounters involved pushing. Additionally, biting occurred in 8–25 % of the trials. Residents were most often the aggressors and usually succeeded in retaining their location. Intruders only succeeded in forcing residents out of their positions if the intruder was equal to or larger in size than the resident. Surveys of an undisturbed population of E. viridis during the daytime indicated that 16% of the individuals were engaged in intraspecific agonistic interactions at any one time.

D. antillarum exhibited biting behavior against both species of Echinometra in 23–24% of the trials. Biting attacks against L. williamsi and E. tribuloides occurred rarely. L. williamsi only once demonstrated pushing behavior and never was observed biting another sea urchin. E. tribuloides occasionally exhibited pushing and biting behaviors, both as residents and as intruders, and was twice observed biting Echinometra.

These studies suggest that two kinds of agonistic interactions may commonly occur among Caribbean reef-dwelling sea urchins: (1) intraspecific and interspecific aggression among Echinometra, and (2) predatory/aggressive attacks against Echinometra by D. antillarum. The former may result in greater dispersion of Echinometra relative to food and shelter resources and control the spatial distribution and concentration of Echinometra grazing pressure within an area. The attacks by D. antillarum may result in the restriction to, or higher densities of, Echinometra in crevices and rugose microhabitats that provide shelter from the larger-bodied D. antillarum.     

Influence of prey body characteristics and performance on predator selection

At the time of settlement to the reef environment, coral reef fishes differ in a number of characteristics that may influence their survival during a predatory encounter. This study investigated the selective nature of predation by both a multi-species predator pool, and a single common predator (Pseudochromis fuscus), on the reef fish, Pomacentrus amboinensis. The study focused on the early post-settlement period of P. amboinensis, when mortality, and hence selection, is known to be highest. Correlations between nine different measures of body condition/performance were examined at the time of settlement, in order to elucidate the relationships between different traits. Single-predator (P. fuscus) choice trials were conducted in 57.4-l aquaria with respect to three different prey characteristics [standard length (SL), body weight and burst swimming speed], whilst multi-species trials were conducted on open patch reefs, manipulating prey body weight only. Relationships between the nine measures of condition/performance were generally poor, with the strongest correlations occurring between the morphological measures and within the performance measures. During aquaria trials, P. fuscus was found to be selective with respect to prey SL only, with larger individuals being selected significantly more often. Multi-species predator communities, however, were selective with respect to prey body weight, with heavier individuals being selected significantly more often than their lighter counterparts. Our results suggest that under controlled conditions, body length may be the most important prey characteristic influencing prey survival during predatory encounters with P. fuscus. In such cases, larger prey size may actually be a distinct disadvantage to survival. However, these relationships appear to be more complex under natural conditions, where the expression of prey characteristics, the selectivity fields of a number of different predators, their relative abundance, and the action of external environmental characteristics, may all influence which individuals survive.     

The influence of habitat characteristics and conspecifics on attraction and survival of coral reef fish juveniles

In marine species with a pelagic larval stage, search behavior and selection of a suitable reef habitat can maximize the settlement success of recently settled juveniles and their subsequent performance (growth and survival of juveniles). Our objective was to test this hypothesis for a single target coral reef fish species (Chromis viridis) at Moorea Island. C. viridis settle on living coral colonies of Porites rus already populated with conspecifics. In the present study (conducted in experimental cages), we found that: 1) mortality rate of recently settled juveniles of C. viridis was lower in the settlement habitat (living coral colonies of P. rus) than in other habitats having physical structure different from those of P. rus colonies; 2) C. viridis juveniles preferentially colonized coral heads of P. rus with conspecifics present rather than uninhabited coral heads and they also preferentially colonized uninhabited coral heads rather than coral heads with heterospecifics; 3) mortality rate of C. viridis juveniles did not vary with the presence or absence of conspecifics or heterospecifics on P. rus colonies. Overall, the study allows us to highlight that site selection by juveniles for habitat containing conspecifics does not benefit their short term mortality rates, suggesting that in the short term at least, site selection has little importance.     

Small-scale recruitment variation in a temperate fish: the roles of macrophytes and food supply

Synopsis The availability of reef-related resources, particularly food and shelter can play a significant role in determining the distribution and abundance of reef fishes. Much of the structure on temperate reefs is provided by macroalgae, and variability in the density of temperate reef fishes at large spatial scales (100's of meters) can often be explained by variation in macroalgal cover or density. In this study I investigated the role of macrophytes and associated food resources on the recruitment of a temperate fish, Tautogolabrus adspersus, at a small spatial scale (0.25 m²). No relationship between the density of new recruits and the percent cover of kelp, foliose or filamentous algae was observed. Multiple regressions revealed that less than 8% of variability in recruitment could be explained by variability in macroalgal cover. However, recruits were found in higher abundance in patches containing many functional forms of seaweeds than in patches dominated by a single form. A wide variety of prey were available for use by cunner recruits; however, crustaceans and mussels were the only common components of their diet, and crustaceans were clearly the most preferred prey. The prey composition in patches where fish were present was compared to randomly selected patches. Significantly greater numbers of isopods, amphipods and newly settled mussels were present in patches where fish were present than in randomly selected patches. The data presented in this study contradict previous work that has shown algal structure to be important in determining patterns of abundance and food supply to be of little significance. A conceptual model is proposed suggesting that settling fish select habitats in a hierarchical manner largely based on their dispersal tendencies. Hierarchical selection of habitats results in different attributes of the habitat being selected during different life-history intervals.     

Kelp gulls Larus dominicanus as predators in kelp Macrocystis pyrifera beds

Summary Minimum daily consumption of bivalves Gaimardia sp by kelp gulls in a Macrocystis kelp bed at Porvenir, Tierra del Fuego, was calculated to be ca. 525 000 individuals, corresponding to an energy removal of ca. 21 kJ·m^-2·d^-1. This is far in excess of energy removal by any other higher-trophic-level predator in southern Chilean Macrocystis beds, but previous work has ignored kelp gulls as a component of the kelp bed community.     

The relative importance of food and shelter for seagrass-associated invertebrates: a latitudinal comparison of habitat choice by isopod grazers

The generality of mechanisms affecting habitat choice and grazing in seagrass meadows was evaluated in a latitudinal comparison of seagrass grazers from the temperate (60°N) Baltic Sea and the subtropical (30°N) Gulf of Mexico. Using similar habitat choice experiment set-ups in Finland and the USA, the role of food type, habitat complexity and predation hazard on habitat choice of the isopods Idotea baltica (Pallas) and Erichsonella attenuata Harger were tested. When shelter was provided by both living and artificial seagrass, epiphytic food resources on artificial vegetation were clearly preferred by both species, although Idotea was attracted to epiphyte-free seagrass when no alternative food was present. When choosing between food and shelter, both species preferred epiphytic food over shelter. However, under predation hazard of fish, Erichsonella clearly switched to the habitat offering shelter, while the presence of a predatory fish produced no preference for shelter by Idotea. Food type may be considered as an universal mechanism that partly determines the presence of grazers in seagrass habitats and is, in the absence of a predator, more important than shelter. Predation risk affected the behaviour of the grazers, but the response varied between species possibly due to varying importance of fish predation in the areas studied. Received: 16 November 1998 / Accepted: 13 February 1999     

Habitat choice in adult longfin damselfish: territory characteristics and relocation times

Habitat selection by coral reef fish during initial settlement has been shown to depend on various biotic and abiotic characteristics. However, relatively little is known of the factors influencing habitat choice by adults during post-settlement processes such as relocation or migration. In this study, we first characterised the habitat of longfin damselfish (Stegastes diencaeus Jordan and Rutter) territories to quantify territory variability. Characteristics such as percentage cover of rock, sand, live coral and distance from sand were highly variable, while territory area, turf and macro algae cover were relatively uniform across territories.We then assessed the importance of specific habitat characteristics by experimentally removing damselfish and measuring recolonisation times in relation to these characteristics. The presence of nest sites markedly increased the speed of territory recolonisation after experimental removals. Other variable territory characteristics such as substrate type, rugosity and the presence of cleaning stations did not affect recolonisation speed. In general, males recolonised territories faster than females, and males were more likely to recolonise territories previously owned by males with an active nest site. Thus, intraspecific competition for high-quality nest sites may generate sex differences in territory relocation and highly stable sex-specific patterns of adult distribution.     

Habitat structure, trophic structure and ecosystem function: interactive effects in a bromeliad–insect community

Although previous studies have shown that ecosystem functions are affected by either trophic structure or habitat structure, there has been little consideration of their combined effects. Such interactions may be particularly important in systems where habitat and trophic structure covary. I use the aquatic insects in bromeliads to examine the combined effects of trophic structure and habitat structure on a key ecosystem function: detrital processing. In Costa Rican bromeliads, trophic structure naturally covaries with both habitat complexity and habitat size, precluding any observational analysis of interactions between factors. I therefore designed mesocosms that allowed each factor to be manipulated separately. Increases in mesocosm complexity reduced predator (damselfly larva) efficiency, resulting in high detritivore abundances, indirectly increasing detrital processing rates. However, increased complexity also directly reduced the per capita foraging efficiency of the detritivores. Over short time periods, these trends effectively cancelled each other out in terms of detrital processing. Over longer time periods, more complex patterns emerged. Increases in mesocosm size also reduced both predator efficiency and detritivore efficiency, leading to no net effect on detrital processing. In many systems, ecosystem functions may be impacted by strong interactions between trophic structure and habitat structure, cautioning against examining either effect in isolation.     

Fish size and habitat depth relationships in headwater streams

Summary Surveys of 262 pools in 3 small streams in eastern Tennessee demonstrated a strong positive relationship between pool depth and the size of the largest fish within a pool (P<0.001). Similarly, the largest colonizers of newly-created deep pools were larger than the colonizers of shallow pools. We explored the role of predation risk in contributing to the bigger fish — deeper habitat pattern, which has been noted by others, by conducting five manipulative field experiments in two streams. Three experiments used stoneroller minnows (Campostoma anomalum); one used creek chubs (Semotilus atromaculatus); and one used striped shiners (Notropis chrysocephalus). The stoneroller experiments showed that survival of fish approximately 100 mm in total length (TL) was much lower in shallow pools (10 cm deep) than in deep (40 cm maximum) pools (19% versus 80% survival over 12 d in one experiment) and added cover markedly increased stoneroller survival in shallow pools (from 49% to 96% in an 11-d experiment). The creek chub experiment showed that, as for stonerollers, pool depth markedly influenced survival: the chubs survived an average of 4.9 d in shallow pools and >10.8 d in deep pools. In the striped shiner experiment in shallow artificial streamside troughs, no individuals 75–100 mm TL survived as long as 13 d, where-as smaller (20–25 mm) fish had 100% survival over 13 d. The results of the experiments show that predation risk from wading/diving animals (e.g., herons and raccoons) is much higher for larger fishes in shallow water than for these fishes in deeper water or for smaller fish in shallow water. We discuss the role of predation risk from two sources (piscivorous fish, which are more effective in deeper habitats, and diving/wading predators, which are more effective in shallow habitats) in contributing to the bigger fish — deeper habitat pattern in streams.     

Habitat selection by two predators in an urban area: The stone marten and red fox in Wrocław (SW Poland)

We analyzed the habitat use of stone martens and red foxes based on incidental observations within the urbanized zone of Wrocław, SW Poland. We compared proportional habitat use at observation sites with randomly selected points and evaluated differences in distance to the water sources and to urban boundaries. Habitat use by both species was different from what we had expected from random points. Stone martens used high-density housing more frequently than red foxes and that expected from random points and avoided open and industrial areas, whereas red foxes used housing estates significantly more often than expected and avoided high-density housing. Both species used the other habitats according to their availability. Stone martens often selected habitats located closer to the city centre, whereas red foxes often selected habitats closer to urban borders than expected. The distribution of red foxes and stone martens is influenced by several factors including the availability of shelter and food, as well as the opportunity to move around undetected. Interspecific competition may also play an important role in habitat selection. Stone martens seem to be better adapted to urbanized areas than red foxes.     

Influences of wind-wave exposure on the distribution and density of recruit reef fishes at Kure and Pearl and Hermes Atolls,Northwestern Hawaiian Islands

This paper describes the results of a field survey designed to test the prediction that the density of benthic juveniles of shallow-reef fishes is greater on wind-wave “exposed” sectors of a pair of isolated oceanic atolls (Kure, Pearl and Hermes) at the far northwestern end of the Hawaiian Islands, an archipelago in which east-northeasterly trade winds dominate onshore water flow and transport by surface currents. The densities of recruits (juveniles ≤5 cm total length) were higher overall on windward versus leeward sectors of carbonate rock-rubble back reefs at both atolls, and the pattern was stronger for smaller (likely younger, more recently settled) recruits of four of the five most abundant species and the remainder pooled as an “Other” taxon. The windward-leeward disparity was four-fold greater at Pearl Hermes (the atoll with a three-fold longer perimeter) than at Kure. Resident predator biomass also was correlated with recruit densities, but habitat (benthic substratum) effects were generally weak. The distribution and abundance of recruits and juveniles of the primarily endemic reef fishes on shallow back reefs at these atolls appear partly influenced by relative rates of water flow over windward vs. leeward sectors of barrier reef and by the size, shape, and orientation of habitat parcels that filter out postlarval fishes with relatively weak swimming capabilities like labroids. Whole-reef geomorphology as well as fine-scale habitat heterogeneity and rugosity should be considered among the suite of many factors used to interpret observed spatial patterns of post-settlement juvenile fish distribution at atolls and perhaps some other tropical reefs. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Epibenthic colonization of concrete and steel pilings in a cold-temperate embayment: a field experiment

With large-scale development of offshore wind farms, vertical structures are becoming more common in open water areas. To examine how vertical structures of different materials may be colonized by epibenthic organisms, an experiment was carried out using steel and concrete pilings constructed to resemble those commonly used in wind farm constructions as well as in bridges, jetties and oil platforms. The early recruitment and succession of the epibenthic communities were sampled once a month for the first 5 months and then again after 1 year. Further, the fish assemblages associated with the pillars were sampled and compared to natural areas. The main epibenthic species groups, in terms of coverage, differed between the two materials at five out of six sampling occasions. Dominant organisms on steel pillars were the barnacle Balanus improvisus, the calcareous tubeworm Pomatoceros triqueter and the tunicate Ciona intestinalis. On the concrete pillars, the hydroid Laomedea sp. and the tunicates Corella parallelogramma and Ascidiella spp. dominated. However, there was no different in coverage at different heights on the pillars or in biomass and species abundance at different directions (north-east or south-west) 5 months after submergence. Fish showed overall higher abundances and species numbers on the pillars (but no difference between steel and concrete) compared to the surrounding soft bottom habitats but not compared to natural vertical rock walls. Two species were attracted to the pillars, indicating a reef effect; Gobiusculus flavescens and Ctenolabrus rupestris. The bottom-dwelling gobies, Pomatoschistus spp., did not show such preferences. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Spawning ascent durations of pelagic spawning reef fishes

Predation risks have been hypothesized to influence spawning behaviors of coral reef fishes that broadcast gametes pelagically.The duration of spawning ascents of 13 species were measured from video footage at a single spawning site for multiple coral reef fishes to investigate if this behavior was influenced by varying risks of predation.Fishes that spawned in pairs had ascents of longer duration than group-spawning species.Duration of spawning ascents did not vary between fishes spawning at daytime and dusk,nor between group-spawning species with specific anti-predatory morphological adaptations.These results indicate that risk of predation may not significantly influence the duration of spawning ascents of pair spawning reef fishes at our study site,while group-spawning behaviors are influenced by predation.Avoidance of egg predation by benthic organisms and female mate choice are more likely to influence the pelagic spawning behaviors of all fishes observed [Current Zoology 58 ( 1 ):95-102,2012].     

Morphological structure in a reef fish assemblage

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. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Using experimental ecology to understand stock enhancement: Comparisons of habitat-related predation on wild and hatchery-reared Penaeus plebejus Hess

Marine stock enhancement is often characterized by poor survival of hatchery-reared individuals due to deficiencies in their fitness, such as a diminished capacity to avoid predators. Field experiments were used to examine predation on Penaeus plebejus, a current candidate for stock enhancement in Australia. We compared overall survival of, and rates of predation on, wild P. plebejus juveniles, naïve hatchery-reared juveniles (which represented the state of individuals intended for stock enhancement) and experienced hatchery-reared juveniles (which had been exposed to natural predatory stimuli). Predation was examined in the presence of an ambush predator (Centropogon australis White, 1790) and an active-pursuit predator (Metapenaeus macleayi Haswell) within both complex (artificial macrophyte) and simple (bare sand and mud) habitats. Overall survival was lower and rates of predation were higher in simple habitats compared to complex habitats in the presence of C. australis. However, the three categories of juveniles survived at similar proportions and suffered similar rates of predation within each individual habitat. No differences in survival and rates of predation were detected among habitats or the categories of juveniles when M. macleayi was used as a predator. These results indicate that wild and hatchery-reared P. plebejus juveniles are equally capable of avoiding predators. Furthermore, exposure of hatchery-reared juveniles to wild conditions does not increase their ability to avoid predators, suggesting an innate rather than learned anti-predator response. The lower predation by C. australis in complex habitats was attributed to a reduction in this ambush predator's foraging efficiency due to the presence of structure. Ecological experiments comparing wild and hatchery-reared individuals should precede all stock enhancement programs because they may identify deficits in hatchery-reared animals that could be mitigated to optimize survival. Such studies can also identify weaknesses in wild animals, relative to hatchery-reared individuals, that may lead to the loss of resident populations.