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1.
Arnold SN  Steneck RS 《PloS one》2011,6(12):e28681
Free space is necessary for larval recruitment in all marine benthic communities. Settling corals, with limited energy to invest in competitive interactions, are particularly vulnerable during settlement into well-developed coral reef communities. This situation may be exacerbated for corals settling into coral-depauperate reefs where succession in nursery microhabitats moves rapidly toward heterotrophic organisms inhospitable to settling corals. To study effects of benthic organisms (at millimeter to centimeter scales) on newly settled corals and their survivorship we deployed terra-cotta coral settlement plates at 10 m depth on the Mesoamerican Barrier Reef in Belize and monitored them for 38 mo. During the second and third years, annual recruitment rates declined by over 50% from the previous year. Invertebrate crusts (primarily sponges) were absent at the start of the experiment but increased in abundance annually from 39, 60, to 73% of the plate undersides by year three. Subsequently, substrates hospitable to coral recruitment, including crustose coralline algae, biofilmed terra-cotta and polychaete tubes, declined. With succession, substrates upon which spat settled shifted toward organisms inimical to survivorship. Over 50% of spat mortality was due to overgrowth by sponges alone. This result suggests that when a disturbance creates primary substrate a "recruitment window" for settling corals exists from approximately 9 to 14 mo following the disturbance. During the window, early-succession, facilitating species are most abundant. The window closes as organisms hostile to coral settlement and survivorship overgrow nursery microhabitats.  相似文献   

2.
 Recruitment of scleractinian corals on settlement plates at Heron Island, Great Barrier Reef, was examined over four years (September 1991–September 1995) to quantify spatio-temporal patterns at different scales and to assess post settlement mortality. Recruitment was dominated by pocilloporid corals which accounted for 80.1% of the 8627 spat counted, whereas non-isoporan acroporids represented only 16.4%. Poritids, faviids and isoporan acroporids rarely recruited to the plates (3.5%), despite their obvious abundance as adults on the reef. Recruitment patterns on the plates indicate strong space-time interactions as evidenced by patchy recruitment of both pocilloporid and acroporid spat. Interactions were found between space (on the scale of 102 m, i.e. sites within zones, and 101 m, i.e. racks within sites) and time (on the scale of years) for pocilloporids and between space (on the scale of 103 m, i.e. zones, and 102 m) and time (on the scale of years) for acroporids. Post-recruitment mortality of acroporid spat in the period 3–10 months after their major spawning was dependent on their initial recruitment density, but pocilloporid mortality was either independent of initial recruitment density or, more likely, obscured by additional recruitment of pocilloporids to plates between late February and September. High rates of recruitment and growth by other sessile organisms, particularly bryozoans and oysters, appear to result in increased post-recruitment mortality and limit recruitment of scleractinian corals on settlement plates. The work reinforces an emerging picture that coral recruitment patterns are determined by mechanisms that manifest over a large range of spatial scales. Accepted: 1 September 1997  相似文献   

3.
Shedding light on the ability of benthic artificial reef (AR) communities to resemble those of a natural reef (NR) is of great importance if we are to harness ARs as tools for rehabilitation and restoration of degraded marine habitats. Studying recruitment processes to experimental settlement plates attached to ARs and NRs reveal the factors that shape community structure at the two reef types, and determine the ability of an AR to support communities similar to those found in adjacent natural habitats. In this study, conducted in Eilat (Red Sea), we used settlement plates to test the hypothesis that differences in benthic communities between ARs and NRs are derived from differential recruitment processes. A monitoring period of 18 months revealed great differences in the recruitment of corals and other benthic communities between the studied ARs and adjacent NRs. The ARs were either made of PVC or metal and 10-17 years old when the study commenced. The recruitment of soft corals reflected the species assemblage found in the area, consisting mainly of the family Nephtheidae and Xeniidae, species, while that of stony corals was mostly determined by the life history traits of the recruited taxa, e.g., the opportunistic nature of the family Pocilloporidae. Benthic organisms, mainly filter feeders like bryozoans, bivalves, sponges and tunicates, were more abundant at the ARs than at the NRs, mainly on the underside of the plates. We suggest that this differential recruitment resulted from a synergistic effect of abiotic and biotic factors, including current regime, sedimentation load and larval settlement preferences, which subsequently differentiated the composition of the benthic communities at the ARs and NRs. Thus, in order to construct an AR for restoration purposes, it must offer similar structural features to those found in the natural surrounding, leading to recruitment of local taxa. However, if the AR and NR will differ structurally, the composition of recruits will also differ and eventually the communities at the two reef types will become distinct, hereby increasing the species diversity in the area.  相似文献   

4.
Tropical reefs are dynamic ecosystems that host diverse coral assemblages with different life-history strategies. Here, we quantified how juvenile (<50 mm) coral demographics influenced benthic coral structure in reef flat and reef slope habitats on the southern Great Barrier Reef, Australia. Permanent plots and settlement tiles were monitored every six months for three years in each habitat. These environments exhibited profound differences: the reef slope was characterised by 95% less macroalgal cover, and twice the amount of available settlement substrata and rates of coral settlement than the reef flat. Consequently, post-settlement coral survival in the reef slope was substantially higher than that of the reef flat, and resulted in a rapid increase in coral cover from 7 to 31% in 2.5 years. In contrast, coral cover on the reef flat remained low (~10%), whereas macroalgal cover increased from 23 to 45%. A positive stock-recruitment relationship was found in brooding corals in both habitats; however, brooding corals were not directly responsible for the observed changes in coral cover. Rather, the rapid increase on the reef slope resulted from high abundances of broadcast spawning Acropora recruits. Incorporating our results into transition matrix models demonstrated that most corals escape mortality once they exceed 50 mm, but for smaller corals mortality in brooders was double those of spawners (i.e. acroporids and massive corals). For corals on the reef flat, sensitivity analysis demonstrated that growth and mortality of larger juveniles (21–50 mm) highly influenced population dynamics; whereas the recruitment, growth and mortality of smaller corals (<20 mm) had the highest influence on reef slope population dynamics. Our results provide insight into the population dynamics and recovery trajectories in disparate reef habitats, and highlight the importance of acroporid recruitment in driving rapid increases in coral cover following large-scale perturbation in reef slope environments.  相似文献   

5.
Studies of the distribution of scleractinian coral recruits within stacks of settlement plates at Orpheus Island and Lizard Island revealed that corals are recruited almost exclusively on the lower surfaces of the settlement plates. The absence of recruitment on the upper surfaces of the plates is attributed to the observed high levels of sediment on these surfaces. All hermatypic coral spat were found near the edge of the settlement plates, whereas ahermatypic corals were located towards the center. Growth and survivorship of hermatypic corals were dependent on the position of attachment, decreasing with distance from the edges of the plates. Survivorship of ahermatypic corals was uniformly high. Light levels on the lower surfaces of the plates within the stacks were shown to decay asymptotically as a function of distance from the plate edge. There was a highly significant non-linear relationship between relative light intensity on the settlement plates and the positions of coral spat. In the absence of evidence for selective hydrodynamic deposition of spat, these results suggest that light intensity is the most important factor in determining the position of attachment of coral spat on the underside of these plates.  相似文献   

6.
During a study of the spatial and temporal patterns of desmosponge (Porifera, Demospongiae) recruitment on rocky and coral reef habitats of Santa Marta, Colombian Caribbean Sea, preliminary attempts were made to estimate actual settlement rates from short-term (1 to a few days) recruitment censuses. Short-term recruitment rates on black, acrylic plastic plates attached to open, non-cryptic substratum by anchor screws were low and variable (0–5 recruits/plate in 1–2 days, sets of n=5–10 plates), but reflected the depth and seasonal trends found using mid-term (1 to a few months) censusing intervals. Moreover, mortality of recruits during 1–2 day intervals was low (0–12%). Thus, short-term censusing intervals can be used to estimate actual settlement rates. To be able to make statistical comparisons, however, it is necessary to increase the number of recruits per census by pooling data of n plates per set, and to have more than one set per site or treatment.  相似文献   

7.
Artificial reefs provide shelter and can be an important source of food for fish depending on the epibenthic community on the structure. The growth and diversity of this community is influenced by the substratum material and the surface orientation of the reef. Settlement plates of four materials (Perspex, sandstone, wood and steel) were deployed in three orientations (upwards, downwards and vertical) at a depth of 33 m on a designed artificial reef (DAR) off the coast of Sydney, Australia. After three months, the steel surfaces had lower invertebrate species richness, total abundance and diversity compared to other surfaces. Steel was not an ideal material for the initial recruitment and growth of epibenthic invertebrates. A longer duration would be required to develop a mature epibenthic community. Surface orientation had species-specific impacts. Surface material and orientation are important factors for developing epibenthic assemblages, and are thus likely to affect the broader artificial reef assemblage, including fish.  相似文献   

8.
Recruitment is widely acknowledged as one of the most important processes in the maintenance of coral reef systems, particularly in their recovery and replenishment following disturbances. In this study variation in coral recruitment was monitored for 3 years at Sodwana Bay, South Africa, located in the Greater St Lucia Wetland Park (GSLWP), a world heritage site. Due to the latitude (27–28°S) and physical conditions, this area is considered marginal for coral reef growth. Recruitment of corals to ceramic tiles peaked between March and May each year, with broadcast spawners dominating plates in March and brooders more abundant in May. Pocilloporid corals were the most abundant during all periods except March 2001, when acroporids comprised 72% of the total number of spat. Total recruitment was the highest during this period, with a mean of 13.2 ± 15.73 (mean ± SD) corals tile−1. Recruitment rates varied widely between and within different reefs monitored. Patterns of variation were inconsistent between seasons, but some reefs had persistently low rates of settlement.  相似文献   

9.
The density of recruits of scleractinian corals on settlement plates at Lord Howe Island, a small isolated sub-tropical island 630 km off the Australian coastline, was within the range of values reported for comparable studies on the Great Barrier Reef. However, there was a difference in the relative abundance of taxonomic groups, with recruitment at Lord Howe Island during the summer of 1990/91 dominated by corals from the Family Pocilloporidae, Family Poritidae, and sub-genus Acropora (Isopora) (in order of abundance). By contrast, on the Great Barrier Reef, recruits are generally predominantly species from the Family Acroporidae (other than the Acropora (Isopora) group). Both the recruits and the established coral communities at Lord Howe Island are dominanted by corals which release brooded planulae, as opposed to the pattern of mass-spawning with external fertilisation more typical of Great Barrier Reef corals. I hypothesise that the release of brooded planulae would be advantageous in an isolated reef community because (a) brooded larvae can travel large distances and survive the journey to the isolated reef and/or (b) brooded larvae have a shorter period before they are competent to settle and are therefore more likely to be retained on the parental reef once a population has been established.  相似文献   

10.
Processes occurring early in the life stages of corals can greatly influence the demography of coral populations, and successful settlement of coral larvae that leads to recruitment is a critical life history stage for coral reef ecosystems. Although corals in Singapore persist in one the world’s most anthropogenically impacted reef systems, our understanding of the role of coral settlement in the persistence of coral communities in Singapore remains limited. Spatial and temporal patterns of coral settlement were examined at 7 sites in the southern islands of Singapore, using settlement tiles deployed and collected every 3 months from 2011 to 2013. Settlement occurred year round, but varied significantly across time and space. Annual coral settlement was low (~54.72 spat m-2 yr-1) relative to other equatorial regions, but there was evidence of temporal variation in settlement rates. Peak settlement occurred between March–May and September–November, coinciding with annual coral spawning periods (March–April and October), while the lowest settlement occurred from December–February during the northeast monsoon. A period of high settlement was also observed between June and August in the first year (2011/12), possibly due to some species spawning outside predicted spawning periods, larvae settling from other locations or extended larval settlement competency periods. Settlement rates varied significantly among sites, but spatial variation was relatively consistent between years, suggesting the strong effects of local coral assemblages or environmental conditions. Pocilloporidae were the most abundant coral spat (83.6%), while Poritidae comprised only 6% of the spat, and Acroporidae <1%. Other, unidentifiable families represented 10% of the coral spat. These results indicate that current settlement patterns are reinforcing the local adult assemblage structure (‘others’; i.e. sediment-tolerant coral taxa) in Singapore, but that the replenishment capacity of Singapore’s reefs appears relatively constrained, which could lead to less resilient reefs.  相似文献   

11.
Restoration of degraded coral reef communities is dependent on successful recruitment and survival of new coral planulae. Degraded reefs are often characterized by high cover of fleshy algae and high microbial densities, complemented by low abundance of coral and coral recruits. Here, we investigated how the presence and abundance of macroalgae and microbes affected recruitment success of a common Hawaiian coral. We found that the presence of algae reduced survivorship and settlement success of planulae. With the addition of the broad-spectrum antibiotic, ampicillin, these negative effects were reversed, suggesting that algae indirectly cause planular mortality by enhancing microbial concentrations or by weakening the coral’s resistance to microbial infections. Algae further reduced recruitment success of corals as planulae preferentially settled on algal surfaces, but later suffered 100% mortality. In contrast to survival, settlement was unsuccessful in treatments containing antibiotics, suggesting that benthic microbes may be necessary to induce settlement. These experiments highlight potential complex interactions that govern the relationships between microbes, algae and corals and emphasize the importance of microbial dynamics in coral reef ecology and restoration. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.  相似文献   

12.
The influence of fucalean canopy species and dominant understory macroalgae on algal colonization was investigated to evaluate whether layering contributes to patterns in algal diversity. Patterns in recruitment were compared among total-clearing, understory-removal, canopy-removal, and undisturbed plots (plot area = 0.25 m2), using a randomized block design in depths <10 m and 10–20 m at Woody Island, Western Australia. To evaluate if propagules were available in the water column above the canopy layer, settlement plates (plate area = 0.04 m2) were deployed in depths <10 m, 10–20 m, and >20 m. A total of 198 macroalgal species was recorded. Biomass of the understory species Osmundaria prolifera Lamouroux and Botryocladia sonderi Silva was similar between canopy-removal and undisturbed plots. Diversity of macroalgae was similar in the presence and absence of a canopy layer. Taxa found in the canopy showed different patterns in recruitment. Cystoseiraceae recruited predominantly in total-clearings in both depth strata. Sargassaceae recruited most abundantly in depths <10 m. Density of canopy taxa on settlement plates was similar with depth (20–30 juveniles per plate), and juveniles were mainly Cystoseiraceae. In contrast to kelp beds or forests, patterns in algal colonization appeared to be maintained by environmental factors or processes other than the direct effects of layering in the subtidal fucoid-dominated assemblages at Woody Island. Handling editor: K. Martens  相似文献   

13.
Sponges mediate consolidation of Porites furcata rubble on shallow Caribbean reefs by quickly adhering to rubble and stabilizing it until carbonate secreting organisms can grow and consolidate it to the reef. Experimental investigations demonstrate that the entire cycle from (1) temporary binding of rubble by sponges, through (2) rubble consolidation by encrusting coralline algae, to (3) colonization of consolidated rubble by corals, can be completed within 10 months. Bound rubble both adds to vertical reef growth and also provides stable substrata for colonization by corals. Corals that colonize stabilized rubble are damaged less and survive better than on unstable rubble. Rubble that is not temporarily stabilized by sponges does not become bound to the reef, because continuous movement disturbs the consolidation process, and does not provide suitable substrata for settlement and growth of corals. Sponge-mediated consolidation of rubble may increase rates of reef growth and enhance reef recovery after damage. This new role for sponges in reef growth is not obvious from examination of the internal fabric of a reef frame. Spongemediated consolidation may help to explain geographic and temporal differences in growth and morphology among shallow reefs of ramose corals.  相似文献   

14.
Although the global decline in coral reef health is likely to have profound effects on reef associated fishes, these effects are poorly understood. While declining coral cover can reduce the abundance of reef fishes through direct effects on recruitment and/or mortality, recent evidence suggests that individuals may survive in disturbed habitats, but may experience sublethal reductions in their condition. This study examined the response of 2 coral associated damselfishes (Pomacentridae), Chrysiptera parasema and Dascyllus melanurus, to varying levels of live coral cover. Growth, persistence, and the condition of individuals were quantified on replicate coral colonies in 3 coral treatments: 100% live coral (control), 50% live coral (partial) and 0% live coral (dead). The growth rates of both species were directly related to the percentage live coral cover, with individuals associated with dead corals exhibiting the slowest growth, and highest growth on control corals. Such differences in individual growth between treatments were apparent after 29 d. There was no significant difference in the numbers of fishes persisting or the physiological condition of individuals between different treatments on this time-scale. Slower growth in disturbed habitats will delay the onset of maturity, reduce lifetime fecundity and increase individual's vulnerability to gape-limited predation. Hence, immediate effects on recruitment and survival may underestimate the longer-term impacts of declining coral on the structure and diversity of coral-associated reef fish communities.  相似文献   

15.
In this study, coral recruitment was measured on a kilometer-wide scale over two years on shallow (5-6 m depth) fringing reefs in St. John, US Virgin Islands, with the objective of determining the extent to which variation in recruitment was affected by biophysical coupling involving temperature and flow. Coral recruitment was measured using settlement tiles deployed at 10 sites along 10 km of shore. The tiles were first deployed in August 2006, and thereafter replaced every ≈ 6 months to sample from either August to January, or January to August over 2 years. Seawater temperature was recorded at the 10 sites using logging thermistors, and flow was quantified using drogues. Overall, corals recruited at a rate equivalent to 76 corals m− 2 6 months− 1, and were represented mostly by poritids (43% of recruits), agaricids (29%), faviids (17%) and siderastreids (7%). Although the density of recruits differed among sites in a pattern that varied among periods and years, there was a consistent trend for mean density to decline from ≈ 4 corals tile− 1 at eastern sites, to ≤ 1 coral tile− 1 at western sites. One aspect of seawater temperature - the daily range - differed among sites and was greater at western compared to eastern sites, and while it was related inversely to recruitment over one of the sampling periods, it was equivocal as a physical process affecting recruitment. Instead, our results are consistent with biophysical coupling involving patch depletion and downstream filtering, whereby patches of coral larvae are delivered to the south shore of St. John and depleted of larvae through settlement as the water progresses westward.  相似文献   

16.
A novel method for the observation of recruitment of fluorescent taxa was developed combining fluorescence census techniques with conventional microscopic examination. The new technique was used to observe coral recruitment on natural limestone plates over a period of 4 months on Meras reef, North Sulawesi, Indonesia. During this period, fluorescence photos were taken of each plate on a weekly basis. This allowed for the detailed observation of fluorescent coral recruitment in-situ. After specific time periods, the plates were sampled and the detected recruits were categorized to family level using their skeletal structure. The fluorescence census technique detected 97.6% of all coral recruits bigger than one millimetre in diameter. The diameter measurements of the recruit's skeletons were used to estimate growth rates at family level. Using the photo series method, the time of settlement of fluorescent recruits on the plates was determined. The results showed roughly linear diameter growth of recruits in the first two months. Acroporidae-polyps were 1.14 mm in diameter when they settled and grew at rate of approximately 0.18 mm in diameter per week. Pocilloporidae-polyps settled at approximately 1.14 mm and showed a growth rate of approximately 0.23 mm in diameter per week. Recruits of three families were observed in detail over time on the artificial plates using this method. While recruitment was fairly continuous, an increase in pocilloporid recruits was observed during the month of September. Similar increases were observed for acroporid recruits during the middle of November and the second quarter of December. This method identifies the limits of detection for the fluorescence census technique used and represents a useful method for the temporal fine scale observation of recruitment in situ.  相似文献   

17.
To determine what happens to scleractinian corals that have been killed by black band disease (BBD), massive corals with BBD were monitored for 11 years on a shallow reef (<10 m depth) in St. John, US Virgin Islands. Small quadrats (0.039 m2) were used to compare the rates of scleractinian recruitment to the skeletons of corals killed by either BBD or physical disturbance (Hurricane Hugo 1989). Coral recruitment was also quantified on the adjacent fringing reef using larger quadrats (0.25 m2) to detect possible biases associated with using small, permanent quadrats to assess recruitment to BBD-killed corals. Of 28 tagged colonies with BBD in 1988, 43% were lost to Hurricane Hugo in 1989, 7% were lost to unknown causes between 1991 and 1992, and 14 were monitored annually for 11 years; of these, 71% were dead and still in their original growth position in 1998. Between 1988 and 1997, corals recruited to the BBD-killed surfaces at a rate of 1.1 ± 0.3 recruits · 0.039 m−2 · decade−1 (mean ± SE, n = 14), although mortality reduced the density to 0.3 ± 0.2 recruits · 0.039 m−2 by 1997. The rate of recruitment and the taxonomic composition of the coral recruits to BBD-killed corals were indistinguishable statistically from those to corals killed by Hurricane Hugo. This demonstrates that BBD creates space that is functionally the same as other dead coral surfaces in providing a substratum for coral recruitment. However, because coral recruits are dispersed widely, clumped in distribution and temporally variable in density on the fringing reef as a whole, it is unlikely that they will be found on monitored coral colonies that have been killed by BBD. While this hypothesis is consistent with the higher density of recruits on the fringing reef compared with BBD-killed corals, further studies are required to investigate alternative explanations such as the role of substratum age in favoring recruitment to surfaces other than those killed recently by BBD. Accepted: 26 August 1999  相似文献   

18.
The global degradation of coral reefs is having profound effects on the structure and species richness of associated reef fish assemblages. Historically, variation in the composition of fish communities has largely been attributed to factors affecting settlement of reef fish larvae. However, the mechanisms that determine how fish settlers respond to different stages of coral stress and the extent of coral loss on fish settlement are poorly understood. Here, we examined the effects of habitat degradation on fish settlement using a two-stage experimental approach. First, we employed laboratory choice experiments to test how settlers responded to early and terminal stages of coral degradation. We then quantified the settlement response of the whole reef fish assemblage in a field perturbation experiment. The laboratory choice experiments tested how juveniles from nine common Indo-Pacific fishes chose among live colonies, partially degraded colonies, and dead colonies with recent algal growth. Many species did not distinguish between live and partially degraded colonies, suggesting settlement patterns are resilient to the early stages of declining coral health. Several species preferred live or degraded corals, and none preferred to associate with dead, algal-covered colonies. In the field experiment, fish recruitment to coral colonies was monitored before and after the introduction of a coral predator (the crown-of-thorns starfish) and compared with undisturbed control colonies. Starfish reduced live coral cover by 95–100%, causing persistent negative effects on the recruitment of coral-associated fishes. Rapid reductions in new recruit abundance, greater numbers of unoccupied colonies and a shift in the recruit community structure from one dominated by coral-associated fishes before degradation to one predominantly composed of algal-associated fish species were observed. Our results suggest that while resistant to coral stress, coral death alters the process of replenishment of coral reef fish communities.  相似文献   

19.
Studies on coral communities have typically been conducted in shallow waters (∼5 m). However, in the face of climate change, and as shallow coral communities become degraded, a greater understanding of deeper coral communities is needed as they become the main reef remnants, playing a central role in the future of coral reefs. To understand the dynamics of deeper coral assemblages, the recruitment and taxonomic composition of different life-stages at 5 and 15 m depths were compared at three locations in Lyudao, southeastern Taiwan in 2010. Coral recruits (<1 cm diameter, <4 months old) were examined using settlement plates. Juvenile corals (1–5 cm, several years old) were examined with quadrats, and adult corals (>5 cm, several years to decades old) were examined using transect lines. Pocilloporid and poritid corals had similar and higher numbers of recruits at 5 m compared to 15 m, whereas acroporid recruits were more abundant at 15 m. The primary cause for the former may be larval behavior, such that they position themselves in shallow waters, while that for the latter may be the dominance of brooding acroporid species (Isopora spp.) at 15 m. The taxonomic composition, especially between recruits and juveniles/adults, was more similar at 15 m than at 5 m. These results suggest a change in the relative importance of pre- and post-settlement processes in assemblage determinants with depth; coral assemblages in shallow habitats (more disturbed) are more influenced by post-settlement processes (mortality events), while those in deeper habitats (more protected) are more influenced by pre-settlement processes (larval supply).  相似文献   

20.
A rapid benthic line-transect survey method for use by non-specialist observers is described. At both Davies Reef (mid-continental shelf) and Myrmidon Reef (outer-continental shelf) in the central Great Barrier Reef a set of 6 sites of varying depths on the reef flat, crest and slope were sampled using this method. At least 10 contiguous 10 m transects were made at each site. Benthic organisms were recorded as life forms with categories based on both high level taxa and morphologies, and including scleractinian corals, alcyonarians, sponges, algae and others. Percentage cover data for 19 benthic categories are presented for all sites. Coral cover on both reefs is high on the crest and slope but low on the reef flat. At all sites the cover of soft corals and sponges is much less than cover of hard corals and algae. Abundances of soft corals and sponges increase with depth. Analysis of gaps between hard corals show that many colonies grow close to each other (<1 cm)even when total coral cover is low.  相似文献   

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