Localization of sensory mechanisms utilized by coral planulae to detect settlement cues

Coral planulae are induced to settle and metamorphose by contact with either crustose coralline algae or marine bacterial biofilms. Larvae of two coral species, Pocillopora damicornis and Montipora capitata, which respond to different metamorphic cues, were utilized to investigate the sensory mechanisms used to detect metamorphic cues. Because the aboral pole of the coral planula is the point of attachment to the substratum, we predicted that it is also the point of detection for cues. To determine where sensory cells for cues are localized along the body, individual larvae were transversely cut into oral and aboral portions at various levels along the oral–aboral axis, and exposed to settlement‐inducing substrata. Aboral ends of M. capitata metamorphosed, while oral ends continued to swim. However, in larvae of P. damicornis, ¾ oral ends (i.e., lacking the aboral pole) were also able to metamorphose, indicating that the cells that detect cues may be distributed along the sides of the body. These cells do not correspond to FMRFamide‐immunoreactive cells that are present throughout the body. Cesium ions induced both aboral and oral ends of larvae of both species to settle, suggesting that oral ends have not lost their capacity to metamorphose, despite lacking sensory cells to detect natural cues. To determine whether sensory cells in larvae of P. damicornis are restricted to one side of the body, swimming behavior over substrata was observed in larvae labeled with diI, a red fluorescent lipophilic membrane stain. The larvae were found to rotate around the oral–aboral axis, with their surface against the substratum, not favoring a particular side for detecting cues. While clarifying the regions of the larval body important for settlement and metamorphosis in coral planulae, we conclude that significant differences between coral species may be due to differences in the distribution of sensory structures in relation to different planular sizes.     

Coral settlement onto Halimeda opuntia: a fatal attraction to an ephemeral substrate?

Degraded reefs with a high abundance of macroalgae usually also have low densities of coral recruits. Few studies, however, have examined whether these algae affect coral larval settlement. This study demonstrates, experimentally, that larvae of the Caribbean coral Favia fragrum can settle on the green alga Halimeda opuntia even when another substrate more suitable for settlement is present. Larval settlement onto experimental substrates was quantified under three treatments: rubble only, rubble plus plastic algal mimic, and rubble plus live H. opuntia. Similar total larval settlement was observed in all treatments. No larvae settled on the algal mimic, but total settlement was similar on the rubble in the first two treatments, showing that the rubble alone offered sufficient substrate for high settlement success. About half the larvae in the live algal treatment settled on H. opuntia instead of on the rubble, showing that larvae did not reject this substrate as they did the algal mimic. This result raises the possibility that corals will settle on some macroalgae when their abundance is high. Most macroalgae, including H. opuntia, are ephemeral substrates unsuitable for post-settlement survival. Such unexpected settlement may therefore have significant consequences for coral recruitment success on algal-dominated reefs.     

Substratum preferences and planulae settling of two red sea alcyonaceans: Xenia macrospiculata Gohar and Parerythropodium fulvum fulvum (Forskl)

The settling behaviour and substratum preferences of the planulae of the Red Sea soft corals Xenia macrospiculata Gohar and Parerythropodium fulvum fulvum (Forskl) were examined in the laboratory. The planulae of the two species have a short pelagic phase and they tend to settle immediately upon leaving the parent colonies. Mucous secretion is used by the larvae for crawling and adhering to the substratum. They exhibit an aggregated pattern of settlement. The developing polyps are found in depressions or pits of the substratum. The planulae preferentially settle on rough substrata and avoid smooth surfaces. They search for substrata covered with an organic coating, composed of turf or crustose coralline algae. Such substrata create better conditions for larval settlement and metamorphosis. The planulae of P. f. fulvum exhibit a striking preference for upside-down attachment on undersides of the substrata, while Xenia macrospiculata utilizes both substratum faces for settlement. Light intensity seems insignificant in determining attachment sites. The findings of the experiments correspond well with the distributional patterns of juveniles of the two species as found in the natural environment. The specific requirements for settling of both species increase their chances of successful development and thus enhance their survival.     

Reproductive seasonality,settlement, and post-settlement mortality of Pocillopora damicornis (Linnaeus), at Lizard Island,Great Barrier Reef

Pocillopora damicornis (Linnaeus) has seasonal gametogenesis and planula release at Lizard Island, Great Barrier Reef, in contrast with several previous reports on the species at other locations. The number of planulae released and gonad development varied considerably among colonies sampled at the same time, but reproductive activity occurred predominantly in winter. P. damicornis planulae settled preferentially on algal-covered substrata, rather than bare coral substrata, but showed subsequent mortality inversely related to this settlement preference. Competition with algae and biological disturbance contribute to spat mortality at different stages of settlement and growth.     

Reef waters stimulate substratum exploration in planulae from brooding Caribbean corals

This study tested the hypothesis that waters surrounding reefs with healthy coral populations are more likely than degraded sites to induce planulae to navigate downward and begin benthic probing. In the laboratory, larvae from two brooding Caribbean coral species, Agaricia tenuifolia and Porites astreoides, were introduced to seawater collected at (1) 1 m above shallow, healthy reef with high-coral cover, (2) 1 m above shallow, degraded reef with high-macroalgal cover, and (3) ~400 m ocean-ward of the reef in deep, blue water. Counter to the hypothesis, water from both the healthy and degraded reef caused the larvae to swim downward and begin benthic probing. These results suggest that substances carried in reef waters may contribute to macro-scale habitat selection by planulae and that understanding how these waterborne cues mesh with other stimuli used by planulae to select a settlement site may be valuable for deciphering a site’s recruitment potential for corals.     

Larval settlement preferences and post-settlement survival of the threatened Caribbean corals Acropora palmata and A. cervicornis

The settlement specificity of two threatened Caribbean corals, Acropora palmata and A. cervicornis, was tested by measuring their rates of larval metamorphosis in response to crustose coralline algae (CCA) and other substrata. In the no-choice experiments, the coral larvae were placed in six treatments: filtered seawater (FSW), a fragment of biofilmed dead skeleton of A. palmata, or a fragment of one of four species of CCA (Hydrolithon boergesenii, Porolithon pachydermum, Paragoniolithon solubile, and Titanoderma prototypum). Within each CCA treatment, there were three different substrata on which to settle and metamorphose: (1) the CCA surface, (2) the rock under the CCA, or (3) the plastic dish. The 5-day-old larvae of both A. palmata and A. cervicornis had similar rates of total metamorphosis (all substrata combined) in every treatment (excluding FSW) even in the absence of CCA. However, there were differences in larval behavior among the CCA species since the larvae settled and metamorphosed on different substrata in the presence of different CCA species. In the no-choice experiments the larvae of both corals had higher rates of metamorphosis on the top surfaces of H. boergesenii and/or T. prototypum than on P. pachydermum. In the choice experiments, the coral larvae were offered two species of CCA in the same dish. When given a choice, both species of coral larvae had more settlement and metamorphosis on the surface of H. boergesenii or T. prototypum or clean rock than onto the surface of P. solubile. After 6 weeks in the field, transplanted A. palmata recruits had approximately 15% survival on both T. prototypum and H. boergesenii, but A. cervicornis recruits only survived on T. prototypum (13%). Some, but not all, CCA species facilitated the larval settlement and post-settlement survival of these two threatened corals, highlighting the importance of benthic community composition for successful coral recruitment.     

The settlement behaviour of planulae larvae of the hermatypic coral Favia fragum (Esper)

Experiments on the settlement behaviour of planulae larvae of the reef coral Favia fragum (Esper) are described. The larvae are positively phototaxic upon release but reverse this and become attracted to dark surfaces, corners, crevices, and the undersides of objects on the bottom. Clean glass surfaces were preferred to surfaces covered with biological slime but there was no preference for rough against smooth surfaces. There was clear evidence of gregarious settlement behaviour, the planulae being able to recognize both adult colonies and previously settled juveniles. A distinction was made between crawling and swimming larvae, and the consequences of their differences in behaviour on the spatial distribution of Favia on reef are discussed. Settlement behaviour of Favia is similar in many respects to that of the Pacific reef coral Pocillopora damicornis (Dana) but is distinguished by gregarious settlement and by a preference for clean surfaces over surfaces covered with biological slime.     

Short-term and latent post-settlement effects associated with elevated temperature and oxidative stress on larvae from the coral Porites astreoides

Coral reefs across the Caribbean are undergoing unprecedented rates of decline in coral cover during the last three decades, and coral recruitment is one potential process that could aid the recovery of coral populations. To better understand the effects of climate change on coral larval ecology, the larvae of Porites astreoides were studied to determine the immediate and post-settlement effects of elevated temperature and associated oxidative stress. Larvae of Porites astreoides were exposed to 27 °C (ambient) and +3.0 °C (elevated temperature) seawater for a short duration of 24 h; then, a suite of physiological parameters were measured to determine the extent of sublethal stress. Following the +3.0 °C treatment, larvae did not show a significant difference in maximum quantum yield of PSII (F _v/F _m) or respiratory demand when compared to controls maintained at 27 °C. The addition of micromolar concentrations of hydrogen peroxide did not impact respiration or photochemical efficiency. Catalase activity in the larvae increased (>60 %) following exposure to elevated temperature when compared to the controls. Short-term larval survival and settlement and metamorphosis were not affected by increased temperature or the H₂O₂ treatment. However, the settled spat that were exposed to elevated temperature underwent a 99 % reduction in survival compared to 90 % reduction for the control spat when examined 24 days following the deployment of 4-day-old settled spat on settlement tiles in the field. These results show that short-term exposure to some stressors might have small impacts on coral physiology, and no effects on larval survival, settlement and metamorphosis. However, due to post-settlement mortality, these stressors can cause a significant reduction in coral recruitment.     

The chemical cue tetrabromopyrrole from a biofilm bacterium induces settlement of multiple Caribbean corals

Microbial biofilms induce larval settlement for some invertebrates, including corals; however, the chemical cues involved have rarely been identified. Here, we demonstrate the role of microbial biofilms in inducing larval settlement with the Caribbean coral Porites astreoides and report the first instance of a chemical cue isolated from a marine biofilm bacterium that induces complete settlement (attachment and metamorphosis) of Caribbean coral larvae. Larvae settled in response to natural biofilms, and the response was eliminated when biofilms were treated with antibiotics. A similar settlement response was elicited by monospecific biofilms of a single bacterial strain, Pseudoalteromonas sp. PS5, isolated from the surface biofilm of a crustose coralline alga. The activity of Pseudoalteromonas sp. PS5 was attributed to the production of a single compound, tetrabromopyrrole (TBP), which has been shown previously to induce metamorphosis without attachment in Pacific acroporid corals. In addition to inducing settlement of brooded larvae (P. astreoides), TBP also induced larval settlement for two broadcast-spawning species, Orbicella (formerly Montastraea) franksi and Acropora palmata, indicating that this compound may have widespread importance among Caribbean coral species.     

Diversity and dynamics of bacterial communities in early life stages of the Caribbean coral Porites astreoides

In this study, we examine microbial communities of early developmental stages of the coral Porites astreoides by sequence analysis of cloned 16S rRNA genes, terminal restriction fragment length polymorphism (TRFLP), and fluorescence in situ hybridization (FISH) imaging. Bacteria are associated with the ectoderm layer in newly released planula larvae, in 4-day-old planulae, and on the newly forming mesenteries surrounding developing septa in juvenile polyps after settlement. Roseobacter clade-associated (RCA) bacteria and Marinobacter sp. are consistently detected in specimens of P. astreoides spanning three early developmental stages, two locations in the Caribbean and 3 years of collection. Multi-response permutation procedures analysis on the TRFLP results do not support significant variation in the bacterial communities associated with P. astreoides larvae across collection location, collection year or developmental stage. The results are the first evidence of vertical transmission (from parent to offspring) of bacteria in corals. The results also show that at least two groups of bacterial taxa, the RCA bacteria and Marinobacter, are consistently associated with juvenile P. astreoides against a complex background of microbial associations, indicating that some components of the microbial community are long-term associates of the corals and may impact host health and survival.     

Adding coral rubble to substrata enhances settlement of Pocillopora?damicornis larvae

Settlement preferences of Pocillopora damicornis larvae were examined on artificial substrata. Planulation of P. damicornis followed a lunar cycle and the release of larvae occurred after new moon. P. damicornis larvae had the highest rates of settlement within 3 days of being presented settlement substrata. Cumulative settlement gradually increased from 3 to 8 days, and post-settlement mortality was most frequent after 8 days. Settlement experiments showed greatest settlement preference to cement tiles containing 10% coral rubble. This study suggests that physical cues are important in the settlement process, which may be useful for coral reef rehabilitation projects.     

Depth-Independent Reproduction in the Reef Coral Porites astreoides from Shallow to Mesophotic Zones

Mesophotic coral ecosystems between 30–150 m may be important refugia habitat for coral reefs and associated benthic communities from climate change and coastal development. However, reduced light at mesophotic depths may present an energetic challenge to the successful reproduction of light-dependent coral organisms, and limit this refugia potential. Here, the relationship of depth and fecundity was investigated in a brooding depth-generalist scleractinian coral, Porites astreoides from 5–37 m in the U.S. Virgin Islands (USVI) using paraffin tissue histology. Despite a trend of increasing planulae production with depth, no significant differences were found in mean peak planulae density between shallow, mid-depth and mesophotic sites. Differential planulae production over depth is thus controlled by P. astreoides coral cover, which peaks at 10 m and ~35 m in the USVI. These results suggest that mesophotic ecosystems are reproductive refuge for P. astreoides in the USVI, and may behave as refugia for P. astreoides metapopulations providing that vertical larval exchanges are viable.     

Temporal settlement patterns of larvae of the broadcast spawning reef coral Favites chinensis and the broadcast spawning and brooding reef coral Goniastrea aspera from Okinawa, Japan

The faviid corals, Favites chinensis and Goniastrea aspera are widely distributed in the Indo-Pacific region. Both corals are hermaphroditic broadcast spawners, but G. aspera is also known to brood planula larvae in Okinawa. This study investigated the temporal settlement patterns of planula larvae of the scleractinian corals F. chinensis and G. aspera that developed from spawned gametes, and planula release and settlement of brooded larvae of G. aspera from Okinawa, Japan. Some of the broadcast-spawned larvae of F. chinensis and G. aspera had very short pre-competency periods of 1–2 and 2–3 days after spawning, and relatively long maximum settlement-competency periods of 56–63 and 63–70 days after spawning, respectively. These pre-competency periods are among the shortest reported for larvae of broadcast spawning coral species, and appear to be negatively correlated with seawater temperature. F. chinensis larvae tended to settle rapidly with 34–39% of larvae settling in the first week after spawning, while broadcast-spawned G. aspera larvae had a slower settlement pattern with 11–15% of larvae settling in the first week after spawning. Brooded larvae of G. aspera settled more rapidly, with settlement rates of 27–31% within the first 24 h and 45–65% within the first week after the start of the experiment. The production of planula larvae with rapid settlement capabilities may enable F. chinensis and G. aspera to establish and maintain populations in shallow reef sites at Okinawa. The release of the brooded planulae for up to 2 months may explain why G. aspera is locally more dominant on shallow reefs in Okinawa than F. chinensis. On a broader scale, the longer settlement competency periods of some of the broadcast-spawned larvae of these species increase their potential for longer-distance dispersal and may partly explain the wide biogeographic distribution of these species in the Indo-Pacific region.     

A corrosive concoction: The combined effects of ocean warming and acidification on the early growth of a stony coral are multiplicative

Survival of coral planulae, and the successful settlement and healthy growth of primary polyps are critical for the dispersal of scleractinian corals and hence the recovery of degraded coral reefs. It is therefore important to explore how the warmer and more acidic oceanic conditions predicted for the future could affect these processes. This study used controlled culture to investigate the effects of a 1 °C increase in temperature and a 0.2-0.25 unit decrease in pH on the settlement and survival of planulae and the growth of primary polyps in the Tropical Eastern Pacific coral Porites panamensis. We found that primary polyp growth was reduced only marginally by more acidic seawater but the combined effect of high temperature and lowered pH caused a significant reduction in growth of primary polyps by almost a third. Elevated temperature was found to significantly reduce the amount of zooxanthellae in primary polyps, and when combined with lowered pH resulted in a significant reduction in biomass of primary polyps. However, survival and settlement of planula larvae were unaffected by increased temperature, lowered acidity or the combination of both. These results indicate that in future scenarios of increased temperature and oceanic acidity coral planulae will be able to disperse and settle successfully but primary polyp growth may be hampered. The recovery of reefs may therefore be impeded by global change even if local stressors are curbed and sufficient sources of planulae are available.     

Crustose coralline algal species host distinct bacterial assemblages on their surfaces

Crustose coralline algae (CCA) are important components of many marine ecosystems. They aid in reef accretion and stabilization, create habitat for other organisms, contribute to carbon sequestration and are important settlement substrata for a number of marine invertebrates. Despite their ecological importance, little is known about the bacterial communities associated with CCA or whether differences in bacterial assemblages may have ecological implications. This study examined the bacterial communities on four different species of CCA collected in Belize using bacterial tag-encoded FLX amplicon pyrosequencing of the V1–V3 region of the 16S rDNA. CCA were dominated by Alphaproteobacteria, Gammaproteobacteria and Actinomycetes. At the operational taxonomic unit (OTU) level, each CCA species had a unique bacterial community that was significantly different from all other CCA species. Hydrolithon boergesenii and Titanoderma prototypum, CCA species that facilitate larval settlement in multiple corals, had higher abundances of OTUs related to bacteria that inhibit the growth and/or biofilm formation of coral pathogens. Fewer coral larvae settle on the surfaces of Paragoniolithon solubile and Porolithon pachydermum. These CCA species had higher abundances of OTUs related to known coral pathogens and cyanobacteria. Coral larvae may be able to use the observed differences in bacterial community composition on CCA species to assess the suitability of these substrata for settlement and selectively settle on CCA species that contain beneficial bacteria.     

Calcium localisation by X-ray microanalysis and fluorescence microscopy in larvae of zooxanthellate and azooxanthellate corals

X-ray microanalysis and fluorescence microscopy (Calcium Orangetrade mark) was used to determine the distribution of intracellular calcium (I(Ca)), in the form of total and ionic calcium respectively, in planulae and settled larvae of a zooxanthellate coral. The distribution of total calcium only was determined in larvae of an azooxanthellate coral. In azooxanthellate planulae and settled larvae, total I(Ca) concentration in the oral ectoderm was high and similar to that in seawater (SW). Calcium concentration did not vary (P > 0.05) between planulae and settled larvae. However, settled larvae accumulated large amounts of calcium in gastrodermal lipid-containing cells. In contrast, zooxanthellate planulae possessed significantly (P < 0.01) lower concentrations of total I(Ca) within ectodermal cells in comparison to settled larvae. In addition, in settled zooxanthellate larvae total calcium concentration in the mesogloea and coelenteron was significantly (P < 0.05) higher than in the oral ectodermal and gastrodermal cells, respectively. Total I(Ca) concentrations in the oral ectoderm of settled larvae were also significantly (P < 0.01) lower than that of the calicoblastic ectoderm. In zooxanthellate settled larvae, ionic I(Ca) levels in the aboral epithelium surrounding rapidly growing septa were high. These levels increased significantly (P < 0.05) within the tissue surrounding growing septa after incubation in high-calcium SW.     

Specificity of larval settlement of the Caribbean Orange Icing Sponge,Mycale laevis

The Caribbean sponge Mycale laevis is often found growing in close proximity to living scleractinian corals. This commonly observed sponge–coral association has been considered a mutualism, with the coral providing substratum for the sponge, and the sponge protecting the coral skeleton from boring organisms. We examined the specificity of sponge recruitment to live corals, expecting a positive and specific settlement response if a mutualism exists. Benthic surveys conducted off Key Largo, Florida, and Bocas del Toro, Panama, revealed that individuals of M. laevis grew on substrata that included dead coral and other species of sponges. Selectivity analysis indicated that at three of the four survey sites, M. laevis was not randomly distributed, but associated with live corals more frequently than expected from proportional coral cover. However, settlement assays demonstrated that larvae of M. laevis did not preferentially respond to the presence of live coral. We have previously demonstrated that adults of M. laevis are chemically undefended and readily eaten by spongivorous fishes unless protected by adjacent substrata such as live corals. In overfished areas, where spongivore density is low, the sponge is not selectively distributed near corals. Initial results of settlement experiments with different substrata suggested that larvae of M. laevis responded positively to the presence of the chemically defended sponge Amphimedon compressa, perhaps indicating an associational defense. Further experiments revealed that larvae were reacting to artificially high concentrations of exudates from cut surfaces of Am. compressa; settlement was not enhanced in response to healed pieces of Am. compressa. In addition, the larvae of M. laevis did not selectively respond to live coral or to chemically defended heterospecifics. These results indicate that the commonly observed proximity of M. laevis to live corals is not driven by larval settlement behavior, but instead by post‐settlement mortality due to predation.     

Rapid settlement in broadcast spawning corals: implications for larval dispersal

Broadcast spawning of gametes with planktonic development of larvae is the most common reproductive mode in tropical corals, and is generally thought to optimize the dispersal potential of larvae. To this end, many previous studies of coral larval dispersal have focused on the maximum time larvae can remain competent to settle and consequently how far they might disperse. However, dispersal ability of broadcast-spawned coral larvae will be linked, at least in part, to the minimum time to settlement competency as well as the length of the planktonic period—although estimates of minimum time to competency remain largely anecdotal, with few rigorous studies of the pre-competent period. To determine the minimum time to larval settlement in two species of broadcast-spawning coral ( Platygyra daedalea and Goniastrea favulus), we monitored larval settlement rates in aquaria every 6 h from the time larvae commenced swimming (i.e. were ciliated, fully developed larvae) for a period of approximately 10 days. For P. daedalea, peak settlement occurred between 60 and 66 h following fertilization (2.5 and 2.75 days), which is markedly earlier than the 4- to 6-day time period commonly cited as the minimum time before broadcast-spawned coral larvae are competent to settle. Surprisingly, it was also clear from our experimental results that settlement in P. daedalea occurred as a distinct pulse during the 60- to 66-h period, rather than continuously throughout the study period. G. favulus larvae also appear to be able to settle quickly (from 54 h following fertilization). We argue, on the basis of these short competency times and apparently rapid settlement, that dispersal in broadcast-spawning coral larvae may not be as great as has previously been assumed.     

Effects of pressure on swimming behavior in planula larvae of the coral Porites astreoides (Cnidaria, Scleractinia)

Mechanisms governing the behavior of coral planulae are not well understood, particularly those manifesting themselves between the time when the larvae are released and when they settle. Larvae from the hermatypic coral Porites astreoides Lamarck were exposed to different levels of hydrostatic pressure—103.4, 206.9, 310.3, 413.8, and 517.1 kPa (including ambient pressure). Data were collected at stops of the above pressures for 15 min each, respectively. This was done in both an increasing sequence and a decreasing one. When exposed to increases in pressure from 103.4 kPa, larvae swam upward (negative barotaxis) in a spiraling motion. Upon exposure to decreasing pressure from 517.1 kPa, larvae moved downward (positive barotaxis), but the magnitude of the vertical movement was much less than in the case of increasing pressure. This suggests that these larvae are more sensitive to increased pressure than decreasing pressure. High variance was also observed in the responses of these larvae at both the intra- and inter-colony levels. Thus, this behavioral trait is variable within the population. The trait may be genetically based, and thus may be susceptible to alteration by natural selection, although this remains to be demonstrated. This study is the first to document these behavioral mechanisms in coral larvae.     

Depth distributions of coral reef fishes: the influence of microhabitat structure,settlement, and post-settlement processes

Many coral reef fishes have restricted depth ranges that are established at settlement or soon after, but the factors limiting these distributions are largely unknown. This study examines whether the availability of microhabitats (reef substrata) explains depth limits, and evaluates whether juvenile growth and survival are lower beyond these limits. Depth-stratified surveys of reef fishes at Kimbe Bay (Papua New Guinea) showed that the abundance of new settlers and the cover of coral substrata differed significantly among depths. A field experiment investigated whether settling coral reef fishes preferred particular depths, and whether these depth preferences were dependent on microhabitat. Small patch reefs composed of identical coral substrata were set up at five depths (3, 6, 10, 15 and 20 m), and settlement patterns were compared to those on unmanipulated reef habitat at the same five depths. For all species, settlement on patch reefs differed significantly among depths despite uniform substratum composition. For four of the six species tested, depth-related settlement patterns on unmanipulated habitat and on patch reefs did not differ, while for the other two, depth ranges were greater on the patch reefs than on unmanipulated habitat. A second experiment examined whether depth preferences reflected variation in growth and survival when microhabitat was similar. Newly settled individuals of Chrysiptera parasema and Dascyllus melanurus were placed, separately, on patch reefs at five depths (as above) and their survival and growth monitored. D. melanurus, which is restricted to shallow depths, had highest survival and growth at the shallowest depth. Depth did not affect either survival or growth of C. parasema, which has a broader depth range than D. melanurus (between 6 and 15 m). This suggests that the fitness costs potentially incurred by settling outside a preferred depth range may depend on the strength of the depth preference.