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The abundance and distribution of demersal fishes rely on larvae successfully settling from the pelagic environment to a benthic habitat and their subsequent survival. With high mortality rates during this life stage, settling to a habitat that maximizes survival is critical. However, relationships between settlement choices and subsequent survival are poorly understood and may vary among species with different habitat preferences. To test this, we focused on five congeneric (Pomacentrus) damselfish species that are known to differ in their habitat choices and explored whether habitat associations at settlement influenced survival. Newly settled individuals were tagged and monitored daily for two weeks to estimate natural mortality rates. Morphological attributes of fish and characteristics of settlement habitats, including depth, rugosity, benthic substrata, and local fish assemblages, were used to predict mortality. We found that some species displayed stronger associations with specific benthic substrata at settlement, but contrary to expectations, these selected habitat characteristics were relatively weak predictors of survivorship. Our survival analysis revealed that the best predictors of survivorship were rugosity (P. adelus and P. amboinensis) and two morphological traits, body depth and ocellus size (P. chrysurus and P. adelus). Interestingly, we found that P. moluccensis that settled in areas of high coverage of mounding coral experienced increased mortality. Of the remaining substrata, analysis showed that instead of associating with habitat characteristics enhancing survivorship, individuals tended not to associate with habitats characteristics that increased mortality (e.g., Turbinaria and sand). This study highlights the species-specific drivers of early post-settlement mortality in coral reef fishes. [ABSTRACT FROM AUTHOR]

Coral reef fishes are known to respond to chemical cues in the selection of appropriate microhabitats at settlement. Coral- and non-coral-associated species are likely to respond to different stimuli and the cues may change as larvae settle and become familiar with the reef environment. Here, the chemosensory responses of both late-stage larvae and newly settled juvenile damselfishes to microhabitat odours were tested in Kimbe Bay (PNG), including four obligate coral-dwelling species (Dascyllus melanurus, D. reticulatus, Chrysiptera arnazae and Pomacentrus moluccensis) and four rubble-dwelling species (Pomacentrus adelus, P. simsiang, Chrysiptera cyanea and C. rollandi). Damselfishes were subjected to a series of pair-wise chemosensory choice trials using a two-channel choice flume. The responses by late-stage larvae (pre-settled) to chemical cues from their preferred microhabitat type mirrored those exhibited by recently settled juveniles. All four rubble-dwelling damselfish species exhibited a significant aversion toward chemical cues derived from coral microhabitats, preferring to remain either in the water seeded with chemical cues derived from rubble, or the unseeded (control) water. The obligate coral-dwelling damselfish species tended to avoid rubble and select coral cues when tested against seawater, but unexpectedly, exhibited a neutral response when given a choice between coral and rubble odours. These results indicate that coral and rubble-dwelling damselfish differ in the strength of their attraction to preferred or avoidance of non-preferred microhabitats. Both factors are likely to play a role in settlement choices. Our results indicated that for some species, newly settled juveniles could act as suitable substitutes for testing larval behavioural traits. [ABSTRACT FROM AUTHOR]

The size and structure of social groups of animals can be governed by a range of ecological factors and behavioral interactions. In small, highly site-attached coral reef fishes, group size is often constrained by the size of the habitat patch they are restricted to. However, group size may also be influenced by changes in abundance along important environmental gradients, such as depth or distance offshore. In addition, the body size and sex structure within social groups can be determined by the size of the habitat patch and the dominance relationships among group members. Here we examined the roles of ecological factors and behavioral interactions in governing group size and structure in the orange clownfish, Amphiprion percula, on inshore reefs in Kimbe Bay, Papua New Guinea. We quantified relationships between ecological variables (anemone size, depth, and distance from shore) and social group variables (group size, and total body length of the three largest individuals (ranks 1, 2, and 3)). Anemone size explained the greatest amount of variation in group variables, with strong, positive relationships between anemone surface area and group size, and total length of individuals ranked 1, 2, and 3. Group structure was also weakly correlated with increasing depth and distance from shore, most likely through the indirect effects of these environmental gradients on anemone size. Variation in group size and the lengths of ranks 2 and 3 were all closely related to the length of rank 1. Path analysis indicated that anemone size has a strong direct effect on the length of rank 1. In turn, the length of rank 1 directly affects the size of the subordinate individuals and indirectly affects the group size through its influence on subordinates. Hence, anemone size directly and indirectly controls social group size and structure in this space-limited fish species. It is also likely that anemonefish have feedback effects on anemone size, although this could not be differentiated in the path analysis. [ABSTRACT FROM AUTHOR]

Coral degradation is a major threat towards the biodiversity of coral-reef ecosystems, either through the physical effects of environmental change, or biological agents such as crown-of-thorns (Acanthaster planci). Coral loss is leading to significant declines in reef-fish assemblages, particularly those dependent on live coral as settlement sites. Most reef fishes use olfactory stimuli at settlement; however, their ability to detect chemical stimuli from degraded corals or A. planci is unknown. Here, olfactory responses of juvenile reef fishes to the presence of stressed corals and A. planci were tested. Juveniles of eight common coral-associated species were subjected to a series of pair-wise choice tests, where the period of time spent in two differing water sources was noted. All species demonstrated a significant attraction towards healthy coral (≥76%), avoiding cues emitted by stressed coral colonies. When given the choice between a control water (untreated reef water) and water containing chemical cues from A. planci, most species elicited no response. Finally, when given the choice between chemical cues derived from feeding A. planci or the control, all species avoided A. planci (≥70%). Our results indicated that juvenile reef fish are capable of distinguishing the state of coral health, but not directly from disturbance agents. [ABSTRACT FROM AUTHOR]

Disturbance can result in the fragmentation and/or loss of suitable habitat, both of which can have important consequences for survival, species interactions, and resulting patterns of local diversity. However, effects of habitat loss and fragmentation are typically confounded during disturbance events, and previous attempts to determine their relative significance have proved ineffective. Here we experimentally manipulated live coral habitats to examine the potential independent and interactive effects of habitat loss and fragmentation on survival, abundance, and species richness of recruitment-stage, coral-associated reef fishes. Loss of 75% of live coral from experimental reefs resulted in low survival of a coral-associated damselfish and low abundance and richness of other recruits 16 weeks after habitat manipulations. In contrast, fragmentation had positive effects on damselfish survival and resulted in greater abundance and species richness of other recruits. We hypothesize that spacing of habitat through fragmentation weakens competition within and among species. Comparison of effect sizes over the course of the study period revealed that, in the first six weeks following habitat manipulations, the positive effects of fragmentation were at least four times stronger than the effects of habitat loss. This initial positive effect of fragmentation attenuated considerably after 16 weeks, whereas the negative effects of habitat loss increased in strength over time. There was little indication that the amount of habitat influenced the magnitude of the habitat fragmentation effect. Numerous studies have reported dramatic declines in coral reef fish abundance and diversity in response to disturbances that cause the loss and fragmentation of coral habitats. Our results suggest that these declines occur as a result of habitat loss, not habitat fragmentation. Positive fragmentation effects may actually buffer against the negative effects of habitat loss and contribute to the resistance of reef fish populations to declines in coral cover. [ABSTRACT FROM AUTHOR]

The relative roles of competition and predation in demographic density dependence are poorly known. A tractable experimental design to determine such effects and their interactions for demersal (seafloor oriented) fishes and similar sedentary species is cross-factoring multiple densities of new recruits with the presence and absence of predators. This design allows one to distinguish between density-dependent mortality due to competition alone, predation alone, or an interaction between the two, especially when supplemental field observations are available. To date, 14 species of marine fish have been examined with some variant of this design, and for 12 species predation was demonstrated to be the sole or major cause of density dependence. However, as competition may be slow acting relative to predation, the importance of competition can be underestimated in short-term experiments. On the Great Barrier Reef, we conducted a long-term field experiment in which multiple densities of new recruits of a planktivorous damselfish were cross-factored with the presence or absence of resident piscivorous fish on patch reefs. During the first 10 months, no density-dependent mortality was detected, regardless of whether resident predators were present or absent. By the end of the experiment at 17 months, per capita mortality was strongly density dependent and highly compensatory in both predator treatments; all reefs ultimately. supported nearly the same adult density regardless of experimental treatment. Examination of treatment effect sizes suggested that competition was the main source of density-dependent mortality, with predation being merely a proximate agent of death. We hypothesize that predators were ineffective in this system compared, with similar studies elsewhere because prey density was low relative to ample prey refuges provided by highly complex corals. Combined with previous studies, these findings indicate that density-dependent mortality in demersal marine fishes is often caused by interplay of predation and competition, whose roles may be altered by variation in habitat complexity and larval supply. These conclusions are relevant to marine fisheries models, which typically assume that density dependence is due solely to intraspecific competition. [ABSTRACT FROM AUTHOR]

Herbivorous fishes have been attributed a central role in structuring benthic communities on coral reefs. However, the relative importance of different behavioural groups of herbivores may differ and their interactions may be complex. This study focuses on an experiment that discriminates between two groups of herbivorous fish: (1) “Foragers” (relatively mobile, schooling grazers, including parrotfishes and surgeonfishes) and (2) “Farmers” (highly site-attached, territorial species, primarily damselfishes). Preliminary observations at Kimbe Bay (Papua New Guinea) showed that both groups were common, and that farmers defended areas from foragers and maintained algal communities that were distinct from the surrounding undefended substratum. An orthogonal combination of a farmer removal treatment and a forager exclusion treatment was applied to isolate their separate effects on algae and corals, and to determine whether farmer territory composition results from forager exclusion or algal cultivation. The experiment showed that foragers had quantitatively greater and qualitatively different effects on sessile benthic community structure than farmers. Where foragers were excluded, there were substantial increases in the cover and biomass of macro-algae and a decline in some corals, regardless of the presence of farmers. Where farmers were removed there was a moderate decline in the cover of some food algal species, regardless of whether foragers had access. No effect of the exclusion of foragers by farmers could be detected. Our results support prevailing views that foragers have a major impact on coral reefs and farmers cultivate selected algae, but challenge the hypothesis that damselfish influence habitat structure by moderating forager disturbance. [ABSTRACT FROM AUTHOR]

The evolution of different colour morphs and how they are maintained in animal populations is poorly understood. We investigated the mechanisms maintaining yellow and brown morphs of a coral-reef fish, Pseudochromis fuscus, at Lizard Island, on the Great Barrier Reef. Histological examination of the gonads revealed that colour morphs were not sex-limited, therefore sexual selection does not appear to promote dichromatism in this species. The field distributions of the two colour morphs were spatially segregated, limiting the opportunity for negative frequency-dependent selection to operate. Our results support another ecological mechanism of coexistence. The yellow morph occurred in deeper areas, usually close to the reef edge, where there was a proportionally high cover of live branching corals. In contrast, the brown morph occurred in shallower areas, more distant from the reef edge, that were proportionally low in live branching corals. Within these habitats, each colour morph of P. fuscus displayed a close association with similar coloured damselfishes from the genus Pomacentrus. The yellow morph was associated with predominantly yellow damselfishes (P. moluccensis and P. amboinensis) and the brown morph with darker coloured species (P. adelus and P. chrysurus). Multiple-choice experiments in the laboratory revealed that: (1) each colour morph of P. fuscus preferentially selected habitat patches occupied by damselfishes with the same colouration; and (2) differences in microhabitat use between the two colour morphs of P. fuscus were due to the presence of different coloured damselfishes in these microhabitats. P. fuscus is a predator of newly recruited damselfishes and the striking resemblance between each morph of P. fuscus and the damselfish with which it was associated, suggests that aggressive mimicry may promote coexistence of P. fuscus colour morphs. [ABSTRACT FROM AUTHOR]

Increasing disturbances on coral reefs threaten fish species with close microhabitat associations in shallow waters, but deep reefs may provide refuge habitats. Assessing this potential requires a comprehensive understanding of how versatility in microhabitat use, preference, and selectivity interact with changes in habitat composition along depth gradients. We examined six damselfish species categorized by versatility of shallow-water microhabitat association (‘complex-coral-specialists’, ‘coral-associates’, and ‘generalists’), along a depth gradient from 10 to 30 m in Kimbe Bay, Papua New Guinea, and tested four important hypotheses. (1) We examined associations with hard-coral and complex-coral microhabitats. Hard-coral association declined with depth more among generalists than coral-associates but complex-coral microhabitat association declined for all species except one complex-coral-specialist. (2) We studied whether microhabitat selectivity declines with depth. Unexpectedly, selectivity increased with depth among both generalist and specialist species. (3) Within species, we tested for positive relationships between fish abundance and preferred microhabitat availability at each depth. However, relationships were stochastic across depths for all but one complex-coral-specialist. (4) Finally, we tested for positive relationships between the number of microhabitats selected by a species and the species’ abundance at each depth, finding that species’ abundances were not consistently related to microhabitat versatility. Our results suggest that several species currently utilize deep coral microhabitats (≤ 30 m), including specialists that strongly associate with vulnerable coral habitats in shallow water. Considerable microhabitat plasticity occurred along the gradient, even amidst stable preferences, and versatile species were not habitat limited, though restricted versatility may limit refuge potential for some species. [ABSTRACT FROM AUTHOR]

Aquatic animals commonly rely on chemical cues to provide information regarding their surroundings. They can respond either by being attracted to (potential mates, preferred habitats) or avoiding (predators, competitors) the source of the stimuli. Coral reef fishes use chemical cues to detect habitats, avoid predators and recognise conspecifics. However, the extent to which chemical cues are used to detect and respond to potential competitors, has received little attention. Here we test olfactory preferences for conspecifics and heterospecifics in newly settled juvenile coral reef fishes. Juveniles of 4 common coral-associated damselfish species: Dascyllus melanurus , Dascyllus reticulatus , Chrysiptera arnazae and Pomacentrus moluccensis, were subjected to olfactory choice tests. Three of the 4 species (excluding P. moluccensis ) demonstrated preferences for waterborne conspecifics odours. All species exhibited an avoidance towards heterospecific odours; this aversion was consistently greatest towards P. moluccensis . A neutral response toward heterospecifics was only evident in two instances (1) between the two congeneric Dascyllus species, with D. melanurus toward D. reticulatus, and (2) with C. arnazae toward D. melanurus . While it is already known that the presence of conspecifics plays a vital role in settlement site selection, we show here that the presence of heterospecifics may also be key in determining the spatial distributions of juveniles across areas of coral reef. [ABSTRACT FROM AUTHOR]

Abstract: Coral reef fish use olfaction to respond to a variety of chemical cues that allow them to detect predators, the presence of resident conspecifics and the ‘smell’ of reef water. However, whether olfaction is utilised by juvenile reef fish for discerning among the multiple microhabitat types present on the reef is not known. Additionally the relative importance of chemical cues emitted from habitat types versus those from resident individuals to juveniles has not been assessed. The ability to distinguish and use such cues could be critical in determining spatial distributions of juveniles across coral reef areas. Here we test olfactory preferences for conspecifics, coral health and different branching coral species in newly recruited juveniles of coral reef fishes. Newly settled juveniles of three common coral associated species in Kimbe Bay, Papua New Guinea, Dascyllus melanurus, Chrysiptera parasema and Chaetodon octofasciatus, were subjected to a series of pairwise olfactory choice tests using two-channel choice flume. All three species exhibited a significant preference for the scent of conspecifics (60–70%) and live coral (60–70%) over water with no associated reef cues. All species strongly avoided the odour of degraded coral (≥80%). Only one species, D. melanurus showed olfactory discrimination among branching coral species, favouring Pocillopora damicornis over Acropora spp. Our results indicate that olfactory preferences for both conspecifics and live coral may combine to determine juvenile and recruitment distribution patterns in these species and local population declines in response to coral degradation but do not necessarily contribute to specific coral habitat associations. [Copyright &y& Elsevier]

Abstract: Although the global decline in coral reef health is likely to have profound effects on reef associated fishes, these effects are poorly understood. While declining coral cover can reduce the abundance of reef fishes through direct effects on recruitment and/or mortality, recent evidence suggests that individuals may survive in disturbed habitats, but may experience sublethal reductions in their condition. This study examined the response of 2 coral associated damselfishes (Pomacentridae), Chrysiptera parasema and Dascyllus melanurus, to varying levels of live coral cover. Growth, persistence, and the condition of individuals were quantified on replicate coral colonies in 3 coral treatments: 100% live coral (control), 50% live coral (partial) and 0% live coral (dead). The growth rates of both species were directly related to the percentage live coral cover, with individuals associated with dead corals exhibiting the slowest growth, and highest growth on control corals. Such differences in individual growth between treatments were apparent after 29 d. There was no significant difference in the numbers of fishes persisting or the physiological condition of individuals between different treatments on this time-scale. Slower growth in disturbed habitats will delay the onset of maturity, reduce lifetime fecundity and increase individual''s vulnerability to gape-limited predation. Hence, immediate effects on recruitment and survival may underestimate the longer-term impacts of declining coral on the structure and diversity of coral-associated reef fish communities. [Copyright &y& Elsevier]

Roving herbivorous fishes play an important role in coral reef communities by removing turf-algae, which can facilitate the settlement of coral larvae. Territorial damselfishes can influence the foraging patterns of roving herbivores by excluding them from their territories, altering the benthic assemblage. However, the impacts depend on the intensity of aggression and which taxonomic groups of roving herbivores are being excluded. Here we document the foraging activity of roving herbivores (Acanthuridae, Scaridae, Siganidae) and the extent to which they are subject to aggression by Pomacentrus adelus , the most abundant territorial damselfish in Kimbe Bay, Papua New Guinea. We then conducted experimental removals (220 m 2 plots on the reef flat) of P. adelus to examine its impact on roving herbivores and the benthic community structure. We hypothesized that the removal of P. adelus would lead to an increase in roving herbivore abundance and foraging activity and a decline in algal cover. The relative abundance (MaxN) and foraging activity (bite rate) of each taxa were examined pre and post-removal using video quadrats. The overall relative abundance of roving herbivores was not influenced by the removal of P. adelus . No changes in foraging patterns were observed for parrotfish, the family that received the highest rate of agonistic interactions, and rabbitfish. The removal of P. adelus resulted in a significant decrease in surgeonfish feeding, suggesting P. adelus alters foraging patterns indirectly through territorial maintenance and not aggression. The only measurable benthic impact of the P. adelus removal was an increase in sediment, while all other substratum types remained constant. These results indicate that P. adelus does not have a negative impact on all roving herbivores and instead may contribute to surgeonfish foraging indirectly through the removal of sediment. The generalisation that territorial damselfish reduce foraging rates of roving herbivores may not be applicable in all systems or for all species. [ABSTRACT FROM AUTHOR]

Abstract: Coral reef ecosystems are experiencing a global decline in coral cover, with direct effects on reef fishes. A decline in habitat may lead to crowding of live coral specialists into remnant habitat patches, intensifying intraspecific competition. Increased local densities of conspecifics are known to negatively affect key demographic processes, but the magnitude of density effects and the role of crowding in response to habitat loss are poorly understood. In this study we examined habitat use and relationships between habitat availability and population density in a coral-dwelling damselfish — Chrysiptera parasema. First, we conducted habitat use and availability surveys to establish the level of habitat selectivity. We then investigated the evidence for crowding due to habitat loss by comparing densities within juvenile aggregations on natural reefs with high and low cover of the preferred habitat. Finally, we used a manipulative patch-reef experiment to measure the potential effects of crowding on mortality of juvenile C. parasema. Surveys revealed that 97% of juvenile C. parasema were associated with Acropora corals. Furthermore, C. parasema densities were closely related to the cover of bottlebrush Acropora, the preferred growth form. Contrary to predictions, there was no evidence of crowding on natural reefs with low coral cover, but rather, reefs with abundant Acropora cover supported larger aggregations with double the density of juveniles. We hypothesized that low densities of C. parasema on natural reefs with low coral cover could be explained by intense intraspecific competition. Experimental manipulations showed that juvenile mortality was density-dependent, with mortality 20% higher on high-density experimental patch-reefs compared to low-density reefs. Behavioural observations on the patch-reefs revealed that the frequency of agonistic interactions and distance to shelter were both unrelated to conspecific densities, highlighting the need for further research into mechanisms underpinning density dependent mortality. These results suggest that intraspecific competition may play an important role in reducing reef fish abundance as a consequence of habitat loss. Given that coral reef systems are currently under threat, with a global decline in coral cover, this study adds to the growing body of knowledge of how disturbances to habitat may affect reef fish communities. [Copyright &y& Elsevier]