Feeding guilds and food resource partitioning in a lake fish assemblage: an ontogenetic approach

Ontogenetic diet patterns and trophic guild structure of a 15 species temperate lake fish assemblage were analysed over wide size intervals (up to seven orders of magnitude in body mass), representing practically the whole life span in most species. A two-step objective clustering technique supplemented with other multivariate statistical tools proved that size-related diet changes clearly played an important role in structuring trophic organization of fishes inhabiting Lake Balaton. As many as 13 out of the 15 fish species showed marked size-related dietary changes with two to four ontogenetic feeding stages. At the assemblage level, 11 trophic guilds were separated. Guild membership was size-dependent in 11 fish species that participated in two to four trophic guilds during their life span. The most complex trophic ontogeny was observed in roach Rutilus rutilus and asp Aspius aspius with four guild memberships. This study showed that trophic status of fishes may be very size-sensitive and thus a universal classification of fish species to general trophic guilds, such as 'planktivore', 'benthivore', 'piscivore' or 'herbivore', should be applied very carefully even in environmental monitoring and fisheries management applications, unless it is supported by relevant results of life span diet analyses.     

Morphological integration and developmental progress during fish ontogeny in two contrasting habitats

SUMMARY Morphological integration can respond to environmental conditions, a response that may be dynamic through ontogeny. Among fishes, brook charrs ( Salvelinus fontinalis ) display a trophic polymorphism that makes it a good species for analyzing the ontogeny of morphological integration. To better understand the processes regulating variation and integration, we assess the ontogenetic dynamics of covariances and developmental progress for populations of S. fontinalis from two habitats that differ in water velocity; lake and stream. Geometric morphometrics and developmental progress were evaluated on 751 and 198 specimens, respectively. In both habitats, most ossification events occur before the transition from alevin to juvenile. This threshold defines two distinct periods. During the first period representing free-embryos and alevins, there are important shape changes and rapid ossification, integration tends to be relatively low and decreasing and the variance of shape drastically decreases. During the juvenile period, the rate of shape change decreases and the onset of ossification is nearly complete, plus integration increases and shape variance stabilizes. While we find two distinct developmental periods, we nonetheless find a notable stability underlying the ontogenetic dynamics of variability as well as gradual change in the structure of covariation within each habitat. Our results imply that the variability of juvenile body shape does not seem to retain signals of variability determined early in ontogeny and warrants caution in using juvenile as guides to the earlier causes of variability. Overall, this study highlights the difficulty of inferring causes of integration from studies of static covariance.     

Testing the evolutionary constraints of metamorphosis: The ontogeny of head shape in Triturus newts

In vertebrates with complex, biphasic, life cycles, larvae have a distinct morphology and ecological preferences compared to metamorphosed juveniles and adults. In amphibians, abrupt and rapid metamorphic changes transform aquatic larvae to terrestrial juveniles. The main aim of this study is to test whether, relative to larval stages, metamorphosis (1) resets the pattern of variation between ontogenetic stages and species, (2) constrains intraspecific morphological variability, and (3) similar to the “hour‐glass” model reduces morphological disparity. We explore postembryonic ontogenetic trajectories of head shape (from hatching to completed metamorphosis) of two well‐defined, morphologically distinct Triturus newts species and their F1 hybrids. Variation in head shape is quantified and compared on two levels: dynamic (across ontogenetic stages) and static (at a particular stage). Our results show that the ontogenetic trajectories diverge early during development and continue to diverge throughout larval stages and metamorphosis. The high within‐group variance and the largest disparity level (between‐group variance) characterize the metamorphosed stage. Hence, our results indicate that metamorphosis does not canalize head shape variation generated during larval development and that metamorphosed phenotype is not more constrained relative to larval ones. Therefore, metamorphosis cannot be regarded as a developmental constraint, at least not for salamander head shape.     

Ontogeny of the hominoid scapula: The influence of locomotion on morphology

Primate shoulder morphology has been linked with locomotor habits, oftentimes irrespective of phylogenetic heritage. Among hominoids, juvenile African apes are known to climb more frequently than adults, while orangutans and gibbons maintain an arboreal lifestyle throughout ontogeny. This study examined if these ontogenetic locomotor differences carry a morphological signal, which should be evident in the scapulae of chimpanzees and gorillas but absent in taxa that do not display ontogenetic behavioral shifts. The scapular morphology of five hominoid primates and one catarrhine outgroup was examined throughout ontogeny to evaluate if scapular traits linked with arboreal activities are modified in response to ontogenetic behavioral shifts away from climbing. Specifically, the following questions were addressed: 1) which scapular characteristics distinguish taxa with different locomotor habits; and 2) do these traits show associated changes during development in taxa known to modify their behavioral patterns? Several traits characterized suspensory taxa from nonsuspensory forms, such as cranially oriented glenohumeral joints, obliquely oriented scapular spines, relatively narrow infraspinous fossae, and inferolaterally expanded subscapularis fossae. The relative shape of the dorsal scapular fossae changed in Pan, Gorilla, and also Macaca in line with predictions based on reported ontogenetic changes in locomotor behavior. These morphological changes were mostly distinct from those seen in Pongo, Hylobates, and Homo and imply a unique developmental pattern, possibly related to ontogenetic locomotor shifts. Accordingly, features that sorted taxa by locomotor habits and changed in concert with ontogenetic behavioral patterns should be particularly useful for reconstructing the locomotor habits of fossil forms. Am J Phys Anthropol 152:239–260, 2013. © 2013 Wiley Periodicals, Inc.     

Evolution of adaptation through allometric shifts in a marine snail

Variation in ontogenetic development among individuals may be a major contributor to morphological variation within species. Evolution of different growth trajectories might, for example, evolve as a response to varying ecological contexts of individuals living in different environments, or by life-stage or gender differences. The intertidal periwinkle Littorina saxatilis is strongly polymorphic in shell shape. We compared ontogenetic trajectories between life stages, local populations, and sexes to understand how different morphological end points are reached during ontogeny and what might cause these differences. Applying landmark-based geometric morphometrics, we captured shell shape variation for four Swedish populations of this species. We also derived a method to visualize ontogenetic trajectories described by the relationship of size to the multivariate shape space. We found that growth trajectories differed between individuals living in different habitats, as well as between sexes and maturity stages. Males living on rocky cliffs grew isometrically throughout life, whereas females from the same habitat switched from isometric growth as juveniles to allometric growth as adults. In contrast, males and females living on boulders grew allometrically as juveniles but changed to isometric growth at sexual maturity. Thus, in this species, ontogenetic growth seems influenced by habitat-associated selection as well as by gender and age-specific selection. These differing selection regimes result in ontogenetic shifts in allometry in three of the four groups examined.     

Diets and trophic‐guild structure of a diverse fish assemblage in Chesapeake Bay,U.S.A.

Dietary habits and trophic‐guild structure were examined in a fish assemblage (47 species) of the Chesapeake Bay estuary, U.S.A., using 10 years of data from >25 000 fish stomachs. The assemblage was comprised of 10 statistically significant trophic guilds that were principally differentiated by the relative amounts of Mysida, Bivalvia, Polychaeta, Teleostei and other Crustacea in the diets. These guilds were broadly aggregated into five trophic categories: piscivores, zooplanktivores, benthivores, crustacivores and miscellaneous consumers. Food web structure was largely dictated by gradients in habitat (benthic to pelagic) and prey size. Size classes within piscivorous species were more likely to be classified into different guilds, reflecting stronger dietary changes through ontogeny relative to benthivores and other guilds. Relative to predator species and predator size, the month of sampling had negligible effects on dietary differences within the assemblage. A majority of sampled fishes derived most of their nutrition from non‐pelagic prey sources, suggesting a strong coupling of fish production to benthic and demersal food resources. Mysida (predominantly the opossum shrimp Neomysis americana) contributed substantially to the diets of over 25% of the sampled predator groups, indicating that this species is a critical, but underappreciated, node in the Chesapeake Bay food web.     

Habitat fragmentation and body‐shape variation of African hind Cephalopholis taeniops (Valenciennes) in an archipelago system (Cape Verde,eastern Atlantic Ocean)

Morphological variations of the serranid fish Cephalopholis taeniops were studied in relation to habitat fragmentation in the Cape Verde Archipelago. While a significant allometric effect existed (11·5% of total body‐shape variation), differences in morphology associated with sex and ontogeny were not significant. MANOVA followed by CVA showed that each island presented a particular allometric pattern. Average body shape for all islands was well discriminated with CVA models. Pair‐wise comparisons of the ontogeny of morphological change between islands revealed that northern islands (Santo Antão, São Vicente and Santa Luzia) along with Boavista Island showed a similar direction in shape ontogeny, while all other paired comparisons indicated different ontogenetic patterns. When comparing directions of inter‐population shape changes, individuals from Fogo Island, the southernmost locality, departed far from the orthogonal relation, suggesting that they were undergoing independent body‐shape trajectories. Physical isolation by geographic distance and depth was positively correlated with morphological divergence among populations from different islands. This finding supports the hypothesis that habitat fragmentation in the Cape Verde Archipelago can be interpreted in terms of marine population structure.     

Seasonal and spatial variations of fish trophic guilds in a shallow,semi-enclosed tropical estuarine bay

A study on seasonal and spatial variations of feeding habits and trophic guilds of dominant fish species in Pattani Bay during March 2003 to February 2004 was aimed at classifying diet composition, identifying dominant food components of each species, categorizing trophic guilds of the community and evaluating effects of habitat characteristics and seasonality on guild organization. Most fishes showed high food intake, fed on a diverse range of food items but relied heavily on calanoid copepods and shrimps. All species, with the exception of Epinephelus coioides, were classified as specialist feeders. Four main dietary guilds were classified. Three of these were classified as the guilds dominated by at least two major food items. Significant variations in trophic guilds of 28 fish species based on habitat types and seasons were also identified. They could be divided into three seasonal groups and three site-groups and a single site. Trophic organization for each season and habitat ranged from two to four groups. This information identifies groups of fishes that seasonally and spatially utilize different food resources within a semi-enclosed estuarine bay ecosystem.     

Linking stream ecology with morphological variability in a native freshwater fish from semi‐arid Australia

Environmental variation is a potent force affecting phenotypic expression. While freshwater fishes have provided a compelling example of the link between the environment and phenotypic diversity, few studies have been conducted with arid‐zone fishes, particularly those that occur in geographically isolated regions where species typically inhabit intermittent and ephemeral creeks. We investigated morphological variation of a freshwater fish (the western rainbowfish, Melanotaenia australis) inhabiting creeks in the Pilbara region of northwest Australia to determine whether body shape variation correlated with local environmental characteristics, including water velocity, habitat complexity, predator presence, and food availability. We expected that the geographic isolation of creeks within this arid region would result in habitat‐specific morphological specializations. We used landmark‐based geometric morphometrics to quantify the level of morphological variability in fish captured from 14 locations within three distinct subcatchments of a major river system. Western rainbowfish exhibited a range of morphologies, with variation in body depth accounting for a significant proportion (>42%) of the total variance in shape. Sexual dimorphism was also apparent, with males displaying deeper bodies than females. While the measured local habitat characteristics explained little of the observed morphological variation, fish displayed significant morphological differentiation at the level of the subcatchment. Local adaptation may partly explain the geographic patterns of body shape variation, but fine‐scale genetic studies are required to disentangle the effects of genetic differentiation from environmentally determined phenotypic plasticity in body shape. Developing a better understanding of environment–phenotype relationships in species from arid regions will provide important insights into ecological and evolutionary processes in these unique and understudied habitats.     

The ontogeny of feeding behaviour in four species of centrarchid fish

Relatively little work has been done on the ontogeny of feeding behaviour in fish. One of the aims of this study was to compare the ontogeny of feeding behaviour in four species of centrarchid fish of diverse feeding guilds. A second aim was to test Mayr's hypothesis that the locomotory acts involved in the ontogeny of feeding behaviour would be stereotyped. All four species initially displayed five feeding acts, with three of the acts disappearing from the repertoire at various intervals. The disappearance of the acts appeared to be related to the size and not to the age of the fry. The acts decreased in frequency as the fry aged and differences among the species in frequencies were minimal. The data supported Mayr's hypothesis but a modifying corollary was added to account for changes due to maturation.     

Interdecadal trends in composition,density, size,and mean trophic level of fish species and guilds before and after coastal development in the Mexican Caribbean

This study explores the possible influence of human coastal development (before and after) and protected area status (within and outside a marine protected area, MPA) on composition, density, and maximum size of fish species and guilds, including mean trophic level of the fish community, in four localities of the Yucatan Peninsula, Mexican Caribbean. Reef fish density, maximum length, species composition, and trophic guilds were recorded by SCUBA belt transects and stationary points in fore reef and lagoon reef areas at decadal intervals (1995–1998, 2006–2010, 2014–2015). Mean density of most species and guilds decreased significantly through the years, as also did mean trophic level of the fish community. Some fish species increased in length. Fish density for many species was larger outside than inside the MPA in 1995–1998; however, the difference tended to disappear in the more recent decades, which reflects either a positive effect of the MPA, or a detrimental effect of coastal development in the non-protected area. Nevertheless, the overall negative trends suggest a regional or global rather than a local cause.     

Different cranial ontogeny in Europeans and Southern Africans

Modern human populations differ in developmental processes and in several phenotypic traits. However, the link between ontogenetic variation and human diversification has not been frequently addressed. Here, we analysed craniofacial ontogenies by means of geometric-morphometrics of Europeans and Southern Africans, according to dental and chronological ages. Results suggest that different adult cranial morphologies between Southern Africans and Europeans arise by a combination of processes that involve traits modified during the prenatal life and others that diverge during early postnatal ontogeny. Main craniofacial changes indicate that Europeans differ from Southern Africans by increasing facial developmental rates and extending the attainment of adult size and shape. Since other studies have suggested that native subsaharan populations attain adulthood earlier than Europeans, it is probable that facial ontogeny is linked with other developmental mechanisms that control the timing of maturation in other variables. Southern Africans appear as retaining young features in adulthood. Facial ontogeny in Europeans produces taller and narrower noses, which seems as an adaptation to colder environments. The lack of these morphological traits in Neanderthals, who lived in cold environments, seems a paradox, but it is probably the consequence of a warm-adapted faces together with precocious maturation. When modern Homo sapiens migrated into Asia and Europe, colder environments might establish pressures that constrained facial growth and development in order to depart from the warm-adapted morphology. Our results provide some answers about how cranial growth and development occur in two human populations and when developmental shifts take place providing a better adaptation to environmental constraints.     

Community structure affects trophic ontogeny in a predatory fish

While most studies have focused on the timing and nature of ontogenetic niche shifts, information is scarce about the effects of community structure on trophic ontogeny of top predators. We investigated how community structure affects ontogenetic niche shifts (i.e., relationships between body length, trophic position, and individual dietary specialization) of a predatory fish, brown trout (Salmo trutta). We used stable isotope and stomach content analyses to test how functional characteristics of lake fish community compositions (competition and prey availability) modulate niche shifts in terms of (i) piscivorous behavior, (ii) trophic position, and (iii) individual dietary specialization. Northern Scandinavian freshwater fish communities were used as a study system, including nine subarctic lakes with contrasting fish community configurations: (i) trout‐only systems, (ii) two‐species systems (brown trout and Arctic charr [Salvelinus alpinus] coexisting), and (iii) three‐species systems (brown trout, Arctic charr, and three‐spined sticklebacks [Gasterosteus aculeatus] coexisting). We expected that the presence of profitable small prey (stickleback) and mixed competitor–prey fish species (charr) supports early piscivory and high individual dietary specialization among trout in multispecies communities, whereas minor ontogenetic shifts were expected in trout‐only systems. From logistic regression models, the presence of a suitable prey fish species (stickleback) emerged as the principal variable determining the size at ontogenetic niche shifts. Generalized additive mixed models indicated that fish community structure shaped ontogenetic niche shifts in trout, with the strongest positive relationships between body length, trophic position, and individual dietary specialization being observed in three‐species communities. Our findings revealed that the presence of a small‐sized prey fish species (stickleback) rather than a mixed competitor–prey fish species (charr) was an important factor affecting the ontogenetic niche‐shift processes of trout. The study demonstrates that community structure may modulate the ontogenetic diet trajectories of and individual niche specialization within a top predator.     

Quantifying rapid spatial and temporal variations of CO<Subscript>2</Subscript> fluxes from small,lowland freshwater ponds

In Lake Tanganyika, several lineages of cichlids have diversified into 200 species. Tribe Tropheini contains the greatest richness of herbivorous species that are classified into four ecomorphs: grazers, browsers, suckers, and pickers. All of these ecomorphs coexist on littoral rocky shores. To reveal the differences among species within ecomorphs, we analysed fish shape and functional morphology related to feeding habits, using geometric morphometrics, and measured intestine length. Furthermore, we tested the relationship between genetic and morphological distances. As a result, diversities of functional morphology among ecomorphs, and among species within ecomorphs, were revealed. In grazers, morphological diversity was greatest in jaws and the opening direction of mouth, suggesting that these species have adapted to feed on various substrata. In browsers, intestine length varied among species, indicating that this ecomorph consists of species with various specializations in herbivory. Morphological divergence was found only in the mandible and occlusal facet in grazers. No clear relation was found between morphological and phylogenetic distances in browsers, which are a polyphyletic group, indicating that their traits were acquired by convergence in multiple lineages. Our data suggest that the observed morphological variation among species enable sympatric species to separate their feeding niches.     

The influence of habitat homogenization on the trophic structure of fish fauna in tropical streams

Habitat homogenization is one of the most important drivers of change in riverine fauna. Therefore, the aim of this study was to determine whether habitat homogenization influences the trophic structure of fish assemblages in tropical streams. We sampled 78 streams located in pasture and crop lands to examine habitat variables and fish. Principal coordinates analysis, canonical analysis of principal coordinates, and a distance-based test for homogeneity of multivariate dispersions revealed two groups of streams, designated homogeneous and heterogeneous, based on the habitat variables. We determined trophic guilds according to the frequency and biovolume of food items. Seven guilds were identified: aquatic insectivores, terrestrial insectivores, detritivores, herbivores, omnivores, algivores, and detritivores–algivores. Homogeneous streams showed higher abundance and biomass of aquatic insectivores, detritivores, and algivores. Heterogeneous streams showed greater diversity of trophic guilds and higher abundance and biomass of terrestrial insectivores and herbivores than homogeneous streams. Our results demonstrate that trophic structure is influenced by habitat condition. Additionally, the riparian canopy and nearshore vegetation have a modulating role in the trophic structure of stream fishes due to their influence on resource supply and promotion of the physical heterogeneity of the channel.     

Effects of habitat and food resources on morphology and ontogenetic growth trajectories in perch

Studies on resource polymorphism have mainly been considered at the end stage of ontogeny, whereas many species undergo diet changes as they grow. We conducted a field survey to analyze the role of adaptive variation during ontogeny in Eurasian perch (Perca fluviatilis). We caught perch from the littoral and pelagic zones of a lake to investigate whether perch differ in morphology and diet between these habitats. We also investigated whether there were any differences in morphological trajectories during the ontogeny of perch collected from the two habitats. We found that perch caught in the littoral habitat, independently of size, had a deeper body, larger head and mouth and longer fins than perch caught in the pelagic zone. Macroinvertebrates and fish dominated the diet of littoral perch, whereas the diet of the pelagic perch consisted mainly of zooplankton and to some extent fish. Independently of size, the more streamlined individuals had a larger proportion of zooplankton and a smaller proportion of macroinvertebrates in their diet than the deeper-bodied individuals, indicating a relation between diet and morphology. Some morphological characters followed different ontogenetic trajectories in the two habitats; e.g. the changes to a deeper body and a larger head were faster in the littoral than in the pelagic perch. The relationship between the length of perch and the size of the mouth and fins also differed between perch from the two habitats, where the increase in the length of the pelvic fin and the area of the mouth increased faster with size in the littoral perch. Our findings show that variation in morphology between habitats differs during ontogeny in a way that corresponds to functional expectations for fish species that occupy these habitats.     

Influence of lateral gradients of hydrologic connectivity on trophic positions of fishes in the Upper Mississippi River

1. Riverscapes consist of the main channel and lateral slackwater habitats along a gradient of hydrological connectivity from maximum connection in main channel habitats to minimum connection in backwaters. Spatiotemporal differences in water currents along this gradient produce dynamic habitat conditions that influence species diversity, population densities and trophic interactions of fishes. 2. We examined the importance of lateral connectivity gradients for food web dynamics in the Upper Mississippi River during spring (high flow, moderately low temperatures) and summer (low flow, higher temperatures). We used literature information and gut contents analyses to determine feeding guilds and stable isotope analysis to estimate mean trophic position of local fish assemblages. During June and August 2006, we collected over 1000 tissue samples from four habitats (main channel, secondary channels, tertiary channels and backwaters) distributed within four hydrologic connectivity gradients. 3. Mean trophic position differed among feeding guilds and seasons, with highest values in spring. Mean trophic position of fish assemblages, variability in trophic position and food chain length (maximum trophic position) of the two dominant piscivore species (Micropterus salmoides and M. dolomieu) in both seasons were significantly associated with habitat along the lateral connectivity gradient. Food chain length peaked in tertiary channels in both seasons, probably due to higher species diversity of prey at these habitats. We infer that food chain length and trophic position of fish assemblages were lower in backwater habitats in the summer mainly because of the use of alternative food sources in these habitats. 4. A greater number of conspecifics exhibited significant among‐habitat variation in trophic position during the summer, indicating that low river stages can constrain fish movements in the Upper Mississippi River. 5. Results of this study should provide a better understanding of the fundamental structure of large river ecosystems and an improved basis for river rehabilitation and management through knowledge of the importance of lateral complexity in rivers.     

Ontogeny and sex alter the effect of predation on body shape in a livebearing fish: sexual dimorphism, parallelism, and costs of reproduction

Predation can cause morphological divergence among populations, while ontogeny and sex often determine much of morphological diversity among individuals. We used geometric morphometrics to characterize body shape in the livebearing fish Brachyrhaphis rhabdophora to test for interactions between these three major shape-determining factors. We assessed shape variation between juveniles and adults of both sexes, and among adults for populations from high- and low-predation areas. Shape differed significantly between predation regimes for all juveniles regardless of sex. As males grew and matured into adults, ontogenetic shape trajectories were parallel, thus maintaining shape differences in adult males between predation environments. However, shape of adult females between predation environments followed a different pattern. As females grew and matured, ontogenetic shape trajectories converged so that shape differences were less pronounced between mature females in predator and nonpredator environments. Convergence in female body shape may indicate a trade-off between optimal shape for predator evasion versus shape required for the livebearing mode of reproduction.     

Patterns of cranial ontogeny in lacertid lizards: morphological and allometric disparity

We explored the ontogenetic dynamics of the morphological and allometric disparity in the cranium shapes of twelve lacertid lizard species. The analysed species (Darevskia praticola, Dinarolacerta mosorensis, Iberolacerta horvathi, Lacerta agilis, L. trilineata, L. viridis, Podarcis erhardii, P. melisellensis, P. muralis, P. sicula, P. taurica and Zootoca vivipara) can be classified into different ecomorphs: terrestrial lizards that inhabit vegetated habitats (habitats with lush or sparse vegetation), saxicolous and shrub‐climbing lizards. We observed that there was an overall increase in the morphological disparity (MD) during the ontogeny of the lacertid lizards. The ventral cranium, which is involved in the mechanics of jaw movement and feeding, showed higher levels of MD, an ontogenetic shift in the morphospace planes and more variable allometric patterns than more conserved dorsal crania. With respect to ecology, the allometric trajectories of the shrub‐climbing species tended to cluster together, whereas the allometric trajectories of the saxicolous species were highly dispersed. Our results indicate that the ontogenetic patterns of morphological and allometric disparity in the lacertid lizards are modified by ecology and functional constraints and that the identical mechanisms that lead to intraspecific morphological variation also produce morphological divergence at higher taxonomic levels.     

Effects of alternate reef states on coral reef fish habitat associations

The present study describes ontogenetic shifts in habitat use for 15 species of coral reef fish at Rangiroa Atoll, French Polynesia. The distribution of fish in different habitats at three ontogenetic stages (new settler, juvenile, and adult) was investigated in coral-dominated and algal-dominated sites at two reefs (fringing reef and inner reef of motu). Three main ontogenetic patterns in habitat use were identified: (1) species that did not change habitats between new settler and juvenile life stages (60% of species) or between juvenile and adult stages (55% of species—no ontogenetic shift); (2) species that changed habitats at different ontogenetic stages (for the transition “new settler to juvenile stage”: 15% of species; for the transition “juvenile to adult stage”: 20% of species); and (3) species that increased the number of habitats they used over ontogeny (for the transition “new settler to juvenile stage”: 25% of species; for the transition “juvenile to adult stage”: 25% of species). Moreover, the majority of studied species (53%) showed a spatial variability in their ontogenetic pattern of habitat use according to alternate reef states (coral reef vs algal reef), suggesting that reef state can influence the dynamics of habitat associations in coral reef fish.