Micro-topography mediates interactions between corals,algae, and herbivorous fishes on coral reefs

Processes occurring during the early life stages of corals are important for the replenishment of coral assemblages and the resilience of coral reefs. However, the factors influencing early life stages of corals are not well understood, and the role of micro-topographic complexity for habitat associations of juvenile corals is largely unexplored. This study investigated the microhabitat distribution patterns of early life stages of corals and a potential macroalgal competitor (Turbinaria ornata) across two reef zones (reef crest and outer reef flat) on Lizard Island, Great Barrier Reef. In both reef zones, both corals and T. ornata were significantly more abundant in concealed microhabitats than in semi-concealed or open microhabitats (GLMM: P < 0.001). The prevalence of juvenile corals and T. ornata within concealed environments suggests that they might be effective refuges from grazing by herbivorous fishes. The density of juvenile corals was positively related, and density of T. ornata negatively related to the abundance of two groups of herbivorous fishes, pairing rabbitfishes, and surgeonfishes in the genus Zebrasoma (BEST ENV-BIO: r _s  = 0.72, P < 0.01), which feed in concealed microhabitats. This correlative evidence suggests that crevices may be important for early life stages of both coral and macroalgae, and that a specific suite of crevice-feeding fishes may influence benthic community dynamics in these microhabitats.     

The role of turtles as coral reef macroherbivores

Herbivory is widely accepted as a vital function on coral reefs. To date, the majority of studies examining herbivory in coral reef environments have focused on the roles of fishes and/or urchins, with relatively few studies considering the potential role of macroherbivores in reef processes. Here, we introduce evidence that highlights the potential role of marine turtles as herbivores on coral reefs. While conducting experimental habitat manipulations to assess the roles of herbivorous reef fishes we observed green turtles (Chelonia mydas) and hawksbill turtles (Eretmochelys imbricata) showing responses that were remarkably similar to those of herbivorous fishes. Reducing the sediment load of the epilithic algal matrix on a coral reef resulted in a forty-fold increase in grazing by green turtles. Hawksbill turtles were also observed to browse transplanted thalli of the macroalga Sargassum swartzii in a coral reef environment. These responses not only show strong parallels to herbivorous reef fishes, but also highlight that marine turtles actively, and intentionally, remove algae from coral reefs. When considering the size and potential historical abundance of marine turtles we suggest that these potentially valuable herbivores may have been lost from many coral reefs before their true importance was understood.     

Direct evaluation of macroalgal removal by herbivorous coral reef fishes

Few studies have examined the relative functional impacts of individual herbivorous fish species on coral reef ecosystem processes in the Indo-Pacific. This study assessed the potential grazing impact of individual species within an inshore herbivorous reef fish assemblage on the central Great Barrier Reef (GBR), by determining which fish species were able to remove particular macroalgal species. Transplanted multiple-choice algal assays and remote stationary underwater digital video cameras were used to quantify the impact of local herbivorous reef fish species on 12 species of macroalgae. Macroalgal removal by the fishes was rapid. Within 3 h of exposure to herbivorous reef fishes there was significant evidence of intense grazing. After 12 h of exposure, 10 of the 12 macroalgal species had decreased to less than 15% of their original mass. Chlorodesmis fastigiata (Chlorophyta) and Galaxaura sp. (Rhodophyta) showed significantly less susceptibility to herbivorous reef fish grazing than all other macroalgae, even after 24 h exposure. Six herbivorous and/or nominally herbivorous reef fish species were identified as the dominant grazers of macroalgae: Siganus doliatus, Siganus canaliculatus, Chlorurus microrhinos, Hipposcarus longiceps, Scarus rivulatus and Pomacanthus sexstriatus. The siganid S. doliatus fed heavily on Hypnea sp., while S. canaliculatus fed intensively on Sargassum sp. Variation in macroalgal susceptibility was not clearly correlated with morphological and/or chemical defenses that have been previously suggested as deterrents against herbivory. Nevertheless, the results stress the potential importance of individual herbivorous reef fish species in removing macroalgae from coral reefs.     

Indirect consequences of fishing: reduction of coralline algae suppresses juvenile coral abundance

Removing predatory fishes has effects that cascade through ecosystems via interactions between species and functional groups. In Kenyan reef lagoons, fishing-induced trophic cascades produce sea urchin-dominated grazing communities that greatly reduce the overall cover of crustose coralline algae (CCA). Certain species of CCA enhance coral recruitment by chemically inducing coral settlement. If sea urchin grazing reduces cover of settlement-inducing CCA, coral recruitment and hence juvenile coral abundance may also decline on fished reefs. To determine whether fishing-induced changes in CCA influence coral recruitment and abundance, we compared (1) CCA taxonomic compositions and (2) taxon-specific associations between CCA and juvenile corals under three fisheries management systems: closed, gear-restricted, and open-access. On fished reefs (gear-restricted and open-access), abundances of two species of settlement-inducing CCA, Hydrolithon reinboldii and H. onkodes, were half those on closed reefs. On both closed and fished reefs, juveniles of four common coral families (Poritidae, Pocilloporidae, Agariciidae, and Faviidae) were more abundant on Hydrolithon than on any other settlement substrate. Coral densities were positively correlated with Hydrolithon spp. cover and were significantly lower on fished than on closed reefs, suggesting that fishing indirectly reduces coral recruitment or juvenile success over large spatial scales via reduction in settlement-inducing CCA. Therefore, managing reefs for higher cover of settlement-inducing CCA may enhance coral recruitment or juvenile survival and help to maintain the ecological and structural stability of reefs.     

Climate-driven coral reorganisation influences aggressive behaviour in juvenile coral-reef fishes

Globally, habitat degradation is altering the abundance and diversity of species in a variety of ecosystems. This study aimed to determine how habitat degradation, in terms of changing coral composition under climate change, affected abundance, species richness and aggressive behaviour of juveniles of three damselfishes (Pomacentrus moluccensis, P. amboinensis and Dischistodus perspicillatus, in order of decreasing reliance on coral). Patch reefs were constructed to simulate two types of reefs: present-day reefs that are vulnerable to climate-induced coral bleaching, and reefs with more bleaching-robust coral taxa, thereby simulating the likely future of coral reefs under a warming climate. Fish communities were allowed to establish naturally on the reefs during the summer recruitment period. Climate-robust reefs had lower total species richness of coral-reef fishes than climate-vulnerable reefs, but total fish abundance was not significantly different between reef types (pooled across all species and life-history stages). The nature of aggressive interactions, measured as the number of aggressive chases, varied according to coral composition; on climate-robust reefs, juveniles used the substratum less often to avoid aggression from competitors, and interspecific aggression became relatively more frequent than intraspecific aggression for juveniles of the coral-obligate P. moluccensis. This study highlights the importance of coral composition as a determinant of behaviour and diversity of coral-reef fishes.     

Density and height of Sargassum influence rabbitfish (f. Siganidae) settlement on inshore reef flats of the Great Barrier Reef

Macroalgal beds have been suggested to be an important settlement habitat for a diversity of reef fishes, yet few studies have considered how the composition or structure of macroalgal beds may influence fish settlement. The aim of this study was to investigate how the physical characteristics of Sargassum beds, a common macroalga on inshore coral reefs, influence the abundance of recently-settled rabbitfishes (Siganidae) on Orpheus Island, Great Barrier Reef. The abundance of recently-settled rabbitfish (< 3 cm total length), the density and height of Sargassum thalli, and benthic composition were quantified within replicate 1-m² quadrats across 15 mid-reef flat sites. A total of 419 recently-settled rabbitfish from three species (Siganus doliatus, S. lineatus and S. canaliculatus) were recorded across 150 quadrats (range 0–16 individuals m⁻²), with S. doliatus accounting for the majority (85.2%) of individuals recorded. The abundance of S. doliatus and S. lineatus was greatest at moderate Sargassum densities (ca. 20–30 holdfasts m⁻²) and generally increased with Sargassum height and the cover of ‘other’ macroalgae. These findings demonstrate the potential importance of the physical characteristics of macroalgal beds to the settlement of rabbitfishes on inshore reef flats.

     

Long-term effects of the cleaner fish Labroides dimidiatus on coral reef fish communities

Cleaning behaviour is deemed a mutualism, however the benefit of cleaning interactions to client individuals is unknown. Furthermore, mechanisms that may shift fish community structure in the presence of cleaning organisms are unclear. Here we show that on patch reefs (61–285 m²) which had all cleaner wrasse Labroides dimidiatus (Labridae) experimentally removed (1–5 adults reef⁻¹) and which were then maintained cleaner-fish free over 8.5 years, individuals of two site-attached (resident) client damselfishes (Pomacentridae) were smaller compared to those on control reefs. Furthermore, resident fishes were 37% less abundant and 23% less species rich per reef, compared to control reefs. Such changes in site-attached fish may reflect lower fish growth rates and/or survivorship. Additionally, juveniles of visitors (fish likely to move between reefs) were 65% less abundant on removal reefs suggesting cleaners may also affect recruitment. This may, in part, explain the 23% lower abundance and 33% lower species richness of visitor fishes, and 66% lower abundance of visitor herbivores (Acanthuridae) on removal reefs that we also observed. This is the first study to demonstrate a benefit of cleaning behaviour to client individuals, in the form of increased size, and to elucidate potential mechanisms leading to community-wide effects on the fish population. Many of the fish groups affected may also indirectly affect other reef organisms, thus further impacting the reef community. The large-scale effect of the presence of the relatively small and uncommon fish, Labroides dimidiadus, on other fishes is unparalleled on coral reefs.     

Abrolhos bank reef health evaluated by means of water quality, microbial diversity, benthic cover, and fish biomass data

The health of the coral reefs of the Abrolhos Bank (Southwestern Atlantic) was characterized with a holistic approach using measurements of four ecosystem components: (i) inorganic and organic nutrient concentrations, [1] fish biomass, [1] macroalgal and coral cover and (iv) microbial community composition and abundance. The possible benefits of protection from fishing were particularly evaluated by comparing sites with varying levels of protection. Two reefs within the well-enforced no-take area of the National Marine Park of Abrolhos (Parcel dos Abrolhos and California) were compared with two unprotected coastal reefs (Sebasti?o Gomes and Pedra de Leste) and one legally protected but poorly enforced coastal reef (the "paper park" of Timbebas Reef). The fish biomass was lower and the fleshy macroalgal cover was higher in the unprotected reefs compared with the protected areas. The unprotected and protected reefs had similar seawater chemistry. Lower vibrio CFU counts were observed in the fully protected area of California Reef. Metagenome analysis showed that the unprotected reefs had a higher abundance of archaeal and viral sequences and more bacterial pathogens, while the protected reefs had a higher abundance of genes related to photosynthesis. Similar to other reef systems in the world, there was evidence that reductions in the biomass of herbivorous fishes and the consequent increase in macroalgal cover in the Abrolhos Bank may be affecting microbial diversity and abundance. Through the integration of different types of ecological data, the present study showed that protection from fishing may lead to greater reef health. The data presented herein suggest that protected coral reefs have higher microbial diversity, with the most degraded reef (Sebasti?o Gomes) showing a marked reduction in microbial species richness. It is concluded that ecological conditions in unprotected reefs may promote the growth and rapid evolution of opportunistic microbial pathogens.     

Branching coral as a macroalgal refuge in a marginal coral reef system

Marginal coral reef systems may provide valuable insights into the nature of ecosystem processes in systems on the trajectory towards a phase shift to an alternate ecosystem state. This study investigates the process of herbivory in a marginal coral reef system in the Keppel Islands at the southern end of the Great Barrier Reef. Branching Acropora coral and the brown macroalga Lobophora variegata occupied up to 95% of the reef crest substratum at the three surveyed reefs. Feeding rates of herbivorous fishes and removal rates of Lobophora were directly quantified within areas of branching Acropora and on planar surfaces. Feeding rates by herbivorous fishes were habitat dependent with the highest bite rates being found in planar habitats for both Lobophora and the epilithic algal matrix (EAM) by 1–2 orders of magnitude, respectively. Feeding rates on Lobophora were, however, much lower than rates on the EAM. The low rates of Lobophora removal and significantly lower rates of herbivory in branching habitats were consistent with the high biomass of this brown alga throughout the Keppel Islands and with its distribution on reef crests, where Lobophora biomass was 20 times greater in branching than in planar habitats. This lack of feeding by herbivorous fishes within branching coral habitats in the Keppel Islands contrasts with the typical role of coral and topographic complexity on herbivores on coral reefs and highlights the potential for complex interactions between algae, corals and fishes on coral reefs. On marginal systems, herbivory may modify algal distributions but may be unable to contain the proliferation of algae such as Lobophora.     

A 3000 year record of Caribbean reef urchin communities reveals causes and consequences of long‐term decline in Diadema antillarum

Urchins are the last abundant grazers of macroalgae on most Caribbean reefs following the historical overexploitation of herbivorous fishes. The long‐spined urchin Diadema antillarum was particularly effective at controlling macroalgae and facilitating coral dominance on Caribbean reefs until its ecological extinction from a catastrophic disease epidemic in the early 1980s. Despite their important role in the structure and functioning of Caribbean reef ecosystems, the natural dynamics of Caribbean reef urchin communities are poorly known due to the paucity of ecological survey data prior to large‐scale human disturbances and the Diadema dieoff. To help resolve the baseline abundances and ecological roles of common urchin taxa, we track changes in urchin abundance and composition over the past 3000 yr from analysis of subfossil urchin spines preserved in reef matrix cores collected in Caribbean Panama. Echinometra consistently dominated the subfossil spine assemblage, while Diadema was consistently rare in the subfossil record in this region. Rather than increasing during a period of heightened human exploitation of their fish competitors and predators, Diadema began declining over a millennium ago. Convergent cross mapping (CCM) causality analyses reveal that Diadema abundance is causally related to coral community composition. Diadema is negatively affected by Acropora cervicornis dominance, likely due to the tight association between this coral and the threespot damselfish, an effective Diadema competitor. Conversely, Diadema positively affects the abundance of the coral Madracis mirabilis, possibly via its control of macroalgae. Causal relationships were not detected among abundances of individual urchin taxa, indicating that inter‐specific echinoid competition is not a factor limiting Diadema recovery. Our detailed record of prehistorical and historical urchin community dynamics suggests that the failure of Diadema to recover over 30 yr after its mass mortality event may be due in part to the prey release of damselfish following the long‐term overfishing of piscivorous fishes.     

Multiscale habitat associations of detrivorous blennies (Blenniidae: Salariini)

The distribution and habitat associations of detrivorous blennies on a tropical coral reef were investigated at several spatial scales and compared with other fish that feed on the epilithic algal matrix to assess density and biomass contributions of small detrivorous fishes to these assemblages. At broad spatial scales total blenny abundance and biomass were highest on the tops of reefs exposed to prevailing winds. On the finer scale of microhabitat use, all species showed a preference for non-living corals, although the type of coral utilised differed between species. The high abundance of blennies on reef tops and non-living corals may be partially related to the quality and availability of detritus in these habitats. Comparisons of total blenny abundance and biomass with other territorial detrivores found that blennies accounted for approximately 60% of this functional group's density and 21% of their biomass on exposed reef tops. Overall, territorial detrivores were found to constitute approximately 37% of the density and 26% of the biomass of the detrivorous/herbivorous fish assemblage on exposed reef tops. Small detrivorous fish therefore represent a substantial proportion of fish assemblages that feed on epilithic algae and associated detritus on coral reefs.     

The role of the reef flat in coral reef trophodynamics: Past,present, and future

The reef flat is one of the largest and most distinctive habitats on coral reefs, yet its role in reef trophodynamics is poorly understood. Evolutionary evidence suggests that reef flat colonization by grazing fishes was a major innovation that permitted the exploitation of new space and trophic resources. However, the reef flat is hydrodynamically challenging, subject to high predation risks and covered with sediments that inhibit feeding by grazers. To explore these opposing influences, we examine the Great Barrier Reef (GBR) as a model system. We focus on grazing herbivores that directly access algal primary productivity in the epilithic algal matrix (EAM). By assessing abundance, biomass, and potential fish productivity, we explore the potential of the reef flat to support key ecosystem processes and its ability to maintain fisheries yields. On the GBR, the reef flat is, by far, the most important habitat for turf‐grazing fishes, supporting an estimated 79% of individuals and 58% of the total biomass of grazing surgeonfishes, parrotfishes, and rabbitfishes. Approximately 59% of all (reef‐wide) turf algal productivity is removed by reef flat grazers. The flat also supports approximately 75% of all grazer biomass growth. Our results highlight the evolutionary and ecological benefits of occupying shallow‐water habitats (permitting a ninefold population increase). The acquisition of key locomotor and feeding traits has enabled fishes to access the trophic benefits of the reef flat, outweighing the costs imposed by water movement, predation, and sediments. Benthic assemblages on reefs in the future may increasingly resemble those seen on reef flats today, with low coral cover, limited topographic complexity, and extensive EAM. Reef flat grazing fishes may therefore play an increasingly important role in key ecosystem processes and in sustaining future fisheries yields.     

A unified model explains commonness and rarity on coral reefs

Abundance patterns in ecological communities have important implications for biodiversity maintenance and ecosystem functioning. However, ecological theory has been largely unsuccessful at capturing multiple macroecological abundance patterns simultaneously. Here, we propose a parsimonious model that unifies widespread ecological relationships involving local aggregation, species‐abundance distributions, and species associations, and we test this model against the metacommunity structure of reef‐building corals and coral reef fishes across the western and central Pacific. For both corals and fishes, the unified model simultaneously captures extremely well local species‐abundance distributions, interspecific variation in the strength of spatial aggregation, patterns of community similarity, species accumulation, and regional species richness, performing far better than alternative models also examined here and in previous work on coral reefs. Our approach contributes to the development of synthetic theory for large‐scale patterns of community structure in nature, and to addressing ongoing challenges in biodiversity conservation at macroecological scales.     

Otolith growth of Springer's demoiselle, Chrysiptera springeri (Pomacentridae, Allen & Lubbock), on a protected and non-protected coral reef

The structural complexity of coral reefs is important for their function as shelter and feeding habitats for coral reef fishes, but physical disturbance by human activities often reduce complexity of the reefs by selectively destroying fragile and more complex coral species. The damselfish Springer's demoiselle Chrysiptera springeri primarily utilize complex coral heads for shelter and are hence vulnerable to human disturbance. In order to evaluate the potential effect of habitat degradation on juvenile fish growth, coral reef cover, fish age at settling and otolith growth, juvenile Springer's demoiselle was investigated on a protected and non‐protected coral reef in Darvel Bay, Borneo. The protected reef had higher coverage of complex branching corals and exhibited a more complex 3‐dimensional structure than the non‐protected reef. Springer's demoiselle settled at the same age on non‐protected and protected reefs. The growth rates of the otoliths from Springer's demoiselle were similar during the pre‐settlement period on the two reefs (manova , P > 0.05), but from age 20 to 48 days (post‐settlement period) the otolith growth rate of juveniles on the non‐protected reef was reduced compared to those from the protected reef (manova , P = 0.017). However, the differences in the otolith size, and by inference, fish size, after 48 days were small. The small effect of habitat degradation on growth is likely related to the fact that the Springer's demoiselles collected on the non‐protected reef were associated with the few remaining complex coral heads. Increased foraging‐predation tradeoffs on the non‐protected reef may decrease food intake and growth of juvenile Springer's demoiselle, but the main effect of habitat degradation on their abundance is likely to be related to lack of suitable shelter, and consequently reduced carrying capacity, on disturbed reefs.     

Evaluation of algal regulation by herbivorous fishes on Caribbean coral reefs

The role of herbivorous fishes in maintaining low macroalgal cover was evaluated on coral reefs on several reef sites from Guadeloupe, either protected or not. Grazing by herbivorous fishes was assessed on different algal facies using fish-bite counts. Algal consumption by fish was estimated as well as algal production. Bite counts revealed that herbivorous fishes feed preferentially on algal turf and avoid brown macroalgae. The algal consumption varied between 0.4 and 2.8 g m⁻² days⁻¹ and was higher inside marine protected areas than outside. Comparison with algal production revealed that herbivorous fishes did not succeed in regulating algal growth. The insufficient number of grazers may lead to the dominance of stable assemblages of macroalgae on coral reefs, preventing the recovery of reef into previous coral-dominated ecosystems.     

Diet and cross-shelf distribution of rabbitfishes (f. Siganidae) on the northern Great Barrier Reef: implications for ecosystem function

Herbivorous fishes are a critical functional group on coral reefs, and there is a clear need to understand the role and relative importance of individual species in reef processes. While numerous studies have quantified the roles of parrotfishes and surgeonfishes on coral reefs, the rabbitfishes (f. Siganidae) have been largely overlooked. Consequently, they are typically viewed as a uniform group of grazing or browsing fishes. Here, we quantify the diet and distribution of rabbitfish assemblages on six reefs spanning the continental shelf in the northern Great Barrier Reef. Our results revealed marked variation in the diet and distribution of rabbitfish species. Analysis of stomach contents identified four distinct groups: browsers of leathery brown macroalgae (Siganus canaliculatus, S. javus), croppers of red and green macroalgae (S. argenteus, S. corallinus, S. doliatus, S. spinus) and mixed feeders of diverse algal material, cyanobacteria, detritus and sediment (S. lineatus, S. punctatissimus, S. punctatus, S. vulpinus). Surprisingly, the diet of the fourth group (S. puellus) contained very little algal material (22.5 %) and was instead dominated by sponges (69.1 %). Together with this variation in diet, the distribution of rabbitfishes displayed clear cross-shelf variation. Biomass was greatest on inner-shelf reefs (112.7 ± 18.2 kg.ha^?1), decreasing markedly on mid- (37.8 ± 4.6 kg.ha^?1) and outer-shelf reefs (9.7 ± 2.2 kg.ha^?1). This pattern was largely driven by the browsing S. canaliculatus that accounted for 50 % of the biomass on inner-shelf reefs, but was absent in mid- and outer-shelf reefs. Mixed feeders, although primarily restricted to the reef slope and back reef habitats, also decreased in abundance and biomass from inshore to offshore, while algal cropping taxa were the dominant group on mid-shelf reefs. These results clearly demonstrate the extent to which diet and distribution vary within the Siganidae and emphasise the importance of examining function on a species-by-species basis.     

Seascape-scale trophic links for fish on inshore coral reefs

It is increasingly accepted that coastal habitats such as inshore coral reefs do not function in isolation but rather as part of a larger habitat network. In the Caribbean, trophic subsidies from habitats adjacent to coral reefs support the diet of reef fishes, but it is not known whether similar trophic links occur on reefs in the Indo-Pacific. Here, we test whether reef fishes in inshore coral, mangrove, and seagrass habitats are supported by trophic links. We used carbon stable isotopes and mathematical mixing models to determine the minimum proportion of resources from mangrove or seagrass habitats in the diet of five fish species from coral reefs at varying distances (0–2,200 m) from these habitats in Moreton Bay, Queensland, eastern Australia. Of the fish species that are more abundant on reefs near to mangroves, Lutjanus russelli and Acanthopagrus australis showed no minimum use of diet sources from mangrove habitat. Siganus fuscescens utilized a minimum of 25–44 % mangrove sources and this contribution increased with the proximity of reefs to mangroves (R ² = 0.91). Seagrass or reef flat sources contributed a minimum of 14–78 % to the diet of Diagramma labiosum, a species found in higher abundance on reefs near seagrass beds, but variation in diet among reefs was unrelated to seascape structure. Seagrass or reef flat sources also contributed a minimum of 8–55 % to a fish species found only on reefs (Pseudolabrus guentheri), indicating that detrital subsidies from these habitats may subsidize fish diet on reefs. These results suggest that carbon sources from multiple habitats contribute to the functioning of inshore coral reef ecosystems and that trophic connectivity between reefs and mangroves may enhance production of a functionally important herbivore.     

Ontogenetic changes in habitat selection during settlement in a coral reef fish: ecological determinants and sensory mechanisms

The behavior of marine larvae during and after settlement can help shape the distribution and abundance of benthic juveniles and therefore the intensity of ecological interactions on reefs. Several laboratory choice-chamber experiments were conducted to explore sensory capabilities and behavioral responses to ecological stimuli to better understand habitat selection by “pre-metamorphic” (larval) and “post-metamorphic” (juvenile) stages of a coral reef fish (Thalassoma hardwicke). T. hardwicke larvae were attracted to benthic macroalgae (Turbinaria ornata and Sargassum mangarevasae), while slightly older post-metamorphosed juveniles chose to occupy live coral colonies (Pocillopora damicornis). Habitat choices of larvae were primarily based upon visual cues and were not influenced by the presence of older conspecifics. In contrast, juveniles selected live coral colonies and preferred those occupied by older conspecifics; choices made by juveniles were based upon both visual and olfactory cues from conspecifics. Overall, the laboratory experiments suggest that early life-history stages of T. hardwicke use a range of sensory modalities that vary through ontogeny, to effectively detect and possibly discriminate among different microhabitats for settlement and later occupation. Habitat selection, based upon cues provided by environmental features and/or by conspecifics, might have important consequences for subsequent competitive interactions.