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Reproduction in regenerating colonies of the coral Stylophora pistillata

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... Ward (1995) found increased mortality and reduced growth in Pocillopora damicornis exposed to trawling, off the Western Australian coast. Gametogenesis can also be adversely affected by stress (Brown & Howard, 1985;Rinkevich & Loya, 1989), and so recovery rates can be slow. Sedimentation rates can also have a strong influence on the recruitment rates of Scleractinians, with excess sediment in an area preventing settling (Rogers et al., 1984;. ...
... It is hypothesised that the constant trawling in this area may keep the L. pertusa colonies at a size below that necessary for gametogenesis to occur. Rinkevich and Loya (1989) observed that removal of even 23% of a colony of Stylophora pistillata causes sterility for up to a year. There is also the possibility of polyp suffocation from increased sediment suspension. ...
... Pollution, coastal development, detrimental fishing practices and climate change have all had their effect on the reef ecosystem. Reduction of colony size through damage has been shown to raise mortality rates (Loya & Rinkevich, 1979;Rinkevich & Loya, 1989;Ward, 1995), increase disease (Rogers et al., 1988) and reduce reproductive output (Loya & Rinkevich, 1980;Szmant-Froelich, 1985;Zakai et al., 2000). ...
Thesis
p>Deep water corals and coral reefs have gained considerable attention recently, both in the public and scientific domain. These corals inhabit depths from relatively shallow at a few tens of metres, to over 6000m, and most have cosmopolitan distributions. These corals can form complex frameworks that attract a variety of invertebrate and vertebrate fauna. Many commercially important species have been observed using these reefs for protection, procreation and feeding, and so have been the target of recent demersal trawling. This thesis considers the reproduction of eleven species of deep-water scleractinian from the NE Atlantic and Antarctica. Shallow-water reef and solitary coral reproduction has been extensively reported, but basic ecological information on deep-water species is lacking. The gametogenesis and reproductive biology of these eleven species was explored by dissection, histological techniques, and scanning electron microscopy. Reproductive data obtained indicate that, in common with shallow water scleractinia, there is no strict pattern to their reproductive habit and a variety of modes were observed. The species examined in this study ranged from hermaphroditic species that spawn gametes ( Caryophyllia ambrosia, C. seguenzae, C. cornuformis ); gonochoric species that spawn gametes seasonally ( Lophelia pertusa, Madrepora oculata, Flabellum angulare ); gonochoric species that spawn quasi-continuously ( Fungiacyathus marenzelleri, Flabellum alabastrum ) and brooding species ( Flabellum thouarsii, F. curvatum, F. impensum ). Oocyte size appears to increase as depth increases, and fecundity reduces with depth. The population dynamics of C. seguenzae was also examined. There appears to be a large juvenile component to this species population, with stability shown through the three years examined.</p
... The energy resources available to an organism are often Limited and are differentially exploited among a variety of biological demands, such as maintenance, somatic growth, reproduction and regeneration (Kozlowski & Wiegert 1986). Sexual reproduction, growth and regeneration in corals are regarded as energy dependent processes (Loya 1976, Rinkevich & Loya 1989, Harrison & Wallace 1990 and several field studies have already documented that damaged corals have lower fecundity during regeneration as compared to intact colonies (Rinkevich & Loya 1989, Harrison & Wallace 1990, Meesters et al. 1994, Van Veghel & Bak 1994, Ward 1995. Such a notion of energetic constraint between regeneration and reproduction has been suggested for several coral species, including Stylophora pistillata (Loya 1976, Rinkevich & Loya 1989, Fungia granulosa (Chadwick & Loya 1990), Montastrea annulaii's an Vegliei & 8dic i394 j, Fociiiopora damicornis (Ward 1995), Acropora hyacinthus, A. gemmifera and Goniastrea retiformis (Hall in press). ...
... The energy resources available to an organism are often Limited and are differentially exploited among a variety of biological demands, such as maintenance, somatic growth, reproduction and regeneration (Kozlowski & Wiegert 1986). Sexual reproduction, growth and regeneration in corals are regarded as energy dependent processes (Loya 1976, Rinkevich & Loya 1989, Harrison & Wallace 1990 and several field studies have already documented that damaged corals have lower fecundity during regeneration as compared to intact colonies (Rinkevich & Loya 1989, Harrison & Wallace 1990, Meesters et al. 1994, Van Veghel & Bak 1994, Ward 1995. Such a notion of energetic constraint between regeneration and reproduction has been suggested for several coral species, including Stylophora pistillata (Loya 1976, Rinkevich & Loya 1989, Fungia granulosa (Chadwick & Loya 1990), Montastrea annulaii's an Vegliei & 8dic i394 j, Fociiiopora damicornis (Ward 1995), Acropora hyacinthus, A. gemmifera and Goniastrea retiformis (Hall in press). ...
... Sexual reproduction, growth and regeneration in corals are regarded as energy dependent processes (Loya 1976, Rinkevich & Loya 1989, Harrison & Wallace 1990 and several field studies have already documented that damaged corals have lower fecundity during regeneration as compared to intact colonies (Rinkevich & Loya 1989, Harrison & Wallace 1990, Meesters et al. 1994, Van Veghel & Bak 1994, Ward 1995. Such a notion of energetic constraint between regeneration and reproduction has been suggested for several coral species, including Stylophora pistillata (Loya 1976, Rinkevich & Loya 1989, Fungia granulosa (Chadwick & Loya 1990), Montastrea annulaii's an Vegliei & 8dic i394 j, Fociiiopora damicornis (Ward 1995), Acropora hyacinthus, A. gemmifera and Goniastrea retiformis (Hall in press). ...
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Here cve document for the first tlme an oriented intra-colonial translocation of photosyn-thetic products to~rards regenerating areas in 2 scleractinian corals, Favia favus (n = 5) and Platygyra lamelljna (n = 3) in Eilat (Red Sea). I4C bicarbonate was injected into round stainless-steel cylinders (10 h daylight incubation period) enabling labeling of a restr~cted tissue area of 20 cm2 in each colony center. Three tissue lesion sizes (small, intermediate and large) were inflicted simultaneously on each colony at a distance of 10 cm from and at different angles to the labeled area. After 3 wk, tissue and skeletal samples were taken from various locations on the coral colonies, and ''C activity was determined. In F. favus a significant labeling of tissues was recorded In areas bordenng the recuperating large lesions and along the axis connecting these lesions with the labeled centers. This pattern of labeling was not found in the smaller les~ons. In P. lamelljna I4C incorporation was recorded In the tlssues bordering large as well as intermediate sized lesions. The skeletal samples from F. favus showed significant 'v deposition in areas bordering the large lesions, indicating the use of translocated I4C materials for skeletogenesis in large lesion regeneration. Our results indicate that I4C labeled materials originating in distant areas of the coral colony are translocated towards large regenerating les~ons. Furthermore, the size of the coral lesion is shown to affect the magnitude of this translocation.
... In hermatypic corals, photosynthetic products are continuously translocated from the symbiotic zooxanthellae to the host tissue and thereby contribute to a variety of nutritional requirements such as maintenance, synthesis of new cells, skeletal matrix and mucus, deposition of calcium carbonate, and storage of energy rich compounds for coral reproduction (Muscatine & Cernichiari 1969, Crossland et al. 1980a, b, Muscatine et al. 1981, Kellogg & Patton 1983. Rinkevich & Loya 1983, Stimson 1987, Rinkevich 1989). These daily fluxes of photosynthates may also contribute to specific biochemical-physiological needs away from sites of algae-coral translocation (Rinkevich 1989). ...
... Rinkevich & Loya 1983, Stimson 1987, Rinkevich 1989). These daily fluxes of photosynthates may also contribute to specific biochemical-physiological needs away from sites of algae-coral translocation (Rinkevich 1989). In a pioneering study on coral-algae symbiosis,Pearse & Muscatine (1971)documented in Acropora cervicornis that organic products (mainly in the forms of lipids, glycerol and glucose) a r e translocated from the base of branches to the tips, contributing to coral calcification. ...
... The energy resources available to an organism are often Limited and are differentially exploited among a variety of biological demands, such as maintenance, somatic growth, reproduction and regeneration (Kozlowski & Wiegert 1986). Sexual reproduction, growth and regeneration in corals are regarded as energy dependent processes (Loya 1976, Rinkevich & Loya 1989, Harrison & Wallace 1990) and several field studies have already documented that damaged corals have lower fecundity during regeneration as compared to intact colonies (Rinkevich & Loya 1989, Harrison & Wallace 1990, Meesters et al. 1994, Van Veghel & Bak 1994, Ward 1995). Such a notion of energetic constraint between regeneration and reproduction has been suggested for several coral species, including Stylophora pistillata (Loya 1976, Rinkevich & Loya 1989), Fungia granulosa (Chadwick & Loya 1990To date no studies have experimentally evaluated the possible allocation of energy resources towards regenerating parts in corals. ...
... However, because recovery from partial mortality represents a major expenditure of energy for the coral, in an effort to maintain colony integrity, it may interrupt other vital processes that require energy reserves such as reproduction and growth (Oren et al., 1997(Oren et al., , 2001. Many studies have determined the effects of partial mortality on coral energy reserves focusing on reproduction (Kojis and Quinn, 1985;Rinkevich and Loya, 1989;Van Veghel and Bak, 1994;Oren et al., 2001;Graham and van Woesik, 2013;Riegl and Purkis, 2015), and growth (Bak, 1983;Fang et al., 1989;Guzmán et al., 1994;Meesters et al., 1994;Lirman, 2000;Alvarado and Acosta, 2009) while regeneration is active. To our knowledge, no studies have documented the effects, if any, once regeneration has stopped and partial mortality is permanent. ...
... All studies to date, on the effects of partial mortality on reproduction and growth have only focused on the first stages of the injury or lesion when regeneration tends to be accelerated. Reported results differ widely ranging from a reduction in reproductive activity in Stylophora pistillata (Rinkevich and Loya, 1989) and massive Porites spp. (Welsh et al., 2015) and fecundity in Goniastrea favulus (Kojis and Quinn, 1985), to cases where the effects are localized and only affected up to a 15 cm radius around injured areas in Favia favus (Oren et al., 2001). ...
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Partial mortality (PM) is increasingly common in the Elkhorn coral Acropora palmata and, depending on the causative agent, is potentially lethal. The effects of PM on growth, reproduction and total lipid content in A. palmata were studied by sampling apparently healthy (AH) colonies in comparison with colonies showing signs of PM. Branch growth rates and lesion regeneration rates were estimated using monthly photographs over a four-month period prior to the summer spawning season. No differences were found in the growth rates of colonies with PM compared to AH colonies. The areas affected by PM did not regenerate during the period of the study. Colonization of the lesions by competing species and sediment cover were documented and did not show major changes. During the spawning season, percent fertilization, egg volume and embryonic development were evaluated for comparison between AH colonies and those with PM. Total lipids were also quantified in tissues from three branches per colony. Percentage fertilization was similar in both AH colonies and those with PM. Embryonic development was normal, regardless of proximity to the lesion borders. However, egg volume was significantly lower in PM colonies than in AH colonies. Lower lipid concentrations were found at the edges of the lesions and similar to those found at the growing edges of the branches. The lack of regeneration may be explained by the low lipid concentration, because the polyps adjacent to the lesion do not have an adequate energy budget as a result of the damage. This would also affect their ability to compete against organisms that colonize the site of the lesion, a distinct situation to the rapid regeneration rates characteristic of lesions due to physical injury of the colony. Therefore, we conclude that partial mortality in A. palmata affects the colony, inducing energetic stress due to both competition and decreased egg quality.
... As a general rule, regeneration is believed to have the highest priority among life functions since natural selection should favor regenerative processes above other requirements (Karlson 1988). This has been demonstrated by several studies on corals reporting resource trade-offs, with injury causing a reduction in fecundity (Rinkevich and Loya 1989;Van Veghel and Bak 1994;Rinkevich 1996) and growth (Meesters et al. 1994; but see Denis et al. 2013). ...
... Apparent trade-offs between major life history processes and regeneration is a well-studied area in corals (Henry and Hart 2005). Injuries may invoke a decline in reproduction (Rinkevich and Loya 1989;Hall 1998) and growth (Meesters et al. 1994) on a colony-wide scale in some species, although this is not always the case. Occasionally, trade-offs in favor of regeneration are avoided, and other biological processes such as growth (Denis et al. 2013) are enhanced. ...
Article
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Ocean acidification (OA) from rising atmospheric carbon dioxide (CO2) is threatening the future of coral reef ecosystems. Mounting experimental evidence suggests that OA negatively impacts fundamental life functions of scleractinian corals, including growth and sexual reproduction. Although regeneration is regarded as a chief life function in scleractinian corals and essential to maintain the colony’s integrity, the effect of OA on regeneration processes has not yet been investigated. To evaluate the effects of OA on regeneration, the common Indo-Pacific corals Porites sp., Favia favus, Acropora eurystoma, and Stylophora pistillata were inflicted with lesions (314–350 mm2, depending on species) and incubated in different pCO2: (1) ambient seawater (400 µatm, pH 8.1), (2) intermediate (1,800 µatm, pH 7.6), and (3) high (4,000 µatm, pH 7.3) for extended periods of time (60–120 d). While all coral species after 60 d had significantly higher tissue regeneration in ambient conditions as compared to the intermediate and high treatments, reduction in regeneration rate was more pronounced in the slow-growing massive Porites sp. and F. favus than the relatively fast-growing, branching S. pistillata and A. eurystoma. This coincided with reduced tissue biomass of Porites sp., F. favus, and A. eurystoma in higher pCO2, but not in S. pistillata. Porites sp., F. favus, and S. pistillata also experienced a decrease in Symbiodinium density in higher pCO2, while in A. eurystoma there was no change. We hypothesize that a lowered regenerative capacity under elevated pCO2 may be related to resource trade-offs, energy cost of acid/base regulation, and/or decrease in total energy budget. This is the first study to demonstrate that elevated pCO2 could have a compounding influence on coral regeneration following injury, potentially affecting the capacity of reef corals to recover following physical disturbance.
... While there are no studies correlating internal pH at the site of gametogenesis and environmental pH, organisms that regulate their internal environment may have increased metabolic demand in low pH conditions, and less energy available for reproductive processes (Sokolova et al., 2012). In corals, tissue damage caused a reduction in coral fecundity (Rinkevich and Loya, 1989;Van Veghel and Bak, 1994;Rinkevich, 1996) and tissue regenerated at a lower rate under OA in various hexacorals (Horwitz and Fine, 2014). When exposed to acidified conditions, the intracellular fluid of the crab Chionoecetes bairdi did not change in low pH. ...
Article
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Sexual reproduction is a fundamental process essential for species persistence, evolution, and diversity. However, unprecedented oceanographic shifts due to climate change can impact physiological processes, with important implications for sexual reproduction. Identifying bottlenecks and vulnerable stages in reproductive cycles will enable better prediction of the organism, population, community, and global-level consequences of ocean change. This article reviews how ocean acidification impacts sexual reproductive processes in marine invertebrates and highlights current research gaps. We focus on five economically and ecologically important taxonomic groups: cnidarians, crustaceans, echinoderms, molluscs and ascidians. We discuss the spatial and temporal variability of experimental designs, identify trends of performance in acidified conditions in the context of early reproductive traits (gametogenesis, fertilization, and reproductive resource allocation), and provide a quantitative meta-analysis of the published literature to assess the effects of low pH on fertilization rates across taxa. A total of 129 published studies investigated the effects of ocean acidification on 122 species in selected taxa. The impact of ocean acidification is dependent on taxa, the specific reproductive process examined, and study location. Our meta-analysis reveals that fertilization rate decreases as pH decreases, but effects are taxa-specific. Echinoderm fertilization appears more sensitive than molluscs to pH changes, and while data are limited, fertilization in cnidarians may be the most sensitive. Studies with echinoderms and bivalve molluscs are prevalent, while crustaceans and cephalopods are among the least studied species even though they constitute some of the largest fisheries worldwide. This lack of information has important implications for commercial aquaculture, wild fisheries, and conservation and restoration of wild populations. We recommend that studies expose organisms to different ocean acidification levels during the entire gametogenic cycle, and not only during the final stages before gametes or larvae are released. We argue for increased focus on fundamental reproductive processes and associated molecular mechanisms that may be vulnerable to shifts in ocean chemistry. Our recommendations for future research will allow for a better understanding of how reproduction in invertebrates will be affected in the context of a rapidly changing environment.
... Tissue damage resulting from these events can negatively affect colony fitness (Rotjan and Lewis 2008;Jayewardene et al. 2009). Further, broken branches present a potential path for pathogens to enter the colony (Ben-Haim et al. 2003;Rodríguez-Villalobos et al. 2015) and for colonization by fouling organisms on the exposed skeleton (Henry and Hart 2005), both of which can affect critical life history processes such as growth rate (Meesters et al. 1994) and reproductive output (Rinkevich and Loya 1989;Rinkevich 1994). While the ability of coral colonies to heal lesions has been found to vary in response to colony parameters and environmental variables (Fong and Lirman 1995;Henry and Hart 2005;Edmunds and Yarid 2017), the relative importance of these drivers and the spatial scale of variation in the wound healing process have not been as well studied. ...
Article
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Coral colonies regularly suffer tissue damage from natural and anthropogenic sources. The resultant wounds can decrease colony fitness and act as sources of infection or algal overgrowth. By systematically breaking branches on 54 Pocillopora meandrina colonies and following in situ tissue regeneration (April–August 2017), variation in the wound recovery process was investigated within colonies, among colonies, and across four sites on O‘ahu, Hawai‘i. Ninety-five percent of all wounds healed, with an average healing time of 42 days. Average healing time was not different between initial and subsequent wounds. The relative importance of intrinsic factors, extrinsic factors, and disturbance history for the wound repair process was examined. Previous colony stressors, i.e., percent live coral tissue and bleaching history, were not correlated with wound healing time. These results indicate that wound repair is a priority for P. meandrina. Colony size and depth were significantly correlated with wound healing time: larger colonies healed 14 days faster than smaller colonies, and deeper colonies healed 25 days slower than shallower colonies. These findings support the hypothesis that larger colonies have more energy available for tissue regeneration. The observation of longer healing times for deeper colonies is likely driven by extrinsic factors that vary with depth, including temperature, wave energy, and irradiance. Overall, we show that wound healing in P. meandrina is physiologically resilient to previous stressors, but is affected by both colony size and depth. Understanding drivers of variation in regenerative processes for corals is critical for predicting coral population recovery after disturbances.
... Following severe bleaching on the Great Barrier Reef in 1998, 45% of colonies of the scleractinian coral Acropora hyacinthus were gravid compared to 100% in the year previous to bleaching (Baird & Marshall, 2002). In addition, physical damage causes corals to significantly decrease reproductive output (Rinkevich & Loya, 1989;Van Veghel & Bak, 1994;Zakai, Levy, & Chadwick-Furman, 2000). There therefore appears to be environmental constraints and an energetic trade-off between parental recovery and reproductive output (Rinkevich, 1996). ...
Article
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Abstract Successful reproductive output and recruitment is crucial to coral persistence and recovery following anthropogenic stress. Feeding is known to alter coral physiology and increase resilience to bleaching. The goal of the study was to address the knowledge gap of the influence of feeding on reproductive output and offspring phenotype. Colonies of Stylophora pistillata from the Northern Gulf of Aqaba (Red Sea) were fed an Artemia diet or unfed for 5 months during gametogenesis, fertilization, and brooding. In addition, time to settlement and mortality of planulae were assessed at water temperatures ranging from winter temperature (22°C) to three degrees above average peak summer temperature (31°C). A range of physiological parameters was measured in parents and offspring. In brooding parents, feeding significantly increased protein concentration and more than tripled the number of released planulae. Planulae from unfed colonies had higher chlorophyll per symbiont concentration and concomitantly higher photosynthetic efficiency compared to planulae from fed parents. In settlement assays, planulae showed a similar thermal resistance as known for this Red Sea adult population. Mortality was greater in planulae from unfed parents at ambient and 3°C above ambient temperature despite higher per offspring investment in terms of total fatty acid content. Fatty acid profiles and relative abundances were generally conserved between different fed and unfed colonies but planulae were enriched in monounsaturated fatty acids relative to adults, that is, 16:1, 18:1, 20:1, 22:1, and 24:1 isomers. Ultimately the availability of zooplankton could influence population physiology and recruitment in corals.
... However, fragmentation by corallivores can also have detrimental effects on coral sexual reproduction. For instance, ∼55% of regenerating fragmented Stylophora pistillata colonies resembling pufferfish predation were sterile during the reproductive season and had an order magnitude less larvae released compared to intact colonies (Rinkevich and Loya, 1989). Moreover, parrotfish in Belize selectively preyed on Montastrea annularis polyps containing more gonads, which reduced coral reproductive effort (Rotjan and Lewis, 2008b). ...
Article
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Corallivory is the predation of coral mucus, tissue, and skeleton by fishes and invertebrates, and a source of chronic stress for many reef-building coral species. Corallivores often prey on corals repeatedly, and this predation induces wounds that require extensive cellular resources to heal. The effects of corallivory on coral growth, reproduction, and community dynamics are well-documented, and often result in reduced growth rates and fitness. Given the degree of anthropogenic pressures that threaten coral reefs, it is now imperative to focus on understanding how corallivory interacts with anthropogenic forces to alter coral health and community dynamics. For example, coral bleaching events that stem from global climate change often reduce preferred corals species for many corallivorous fishes. These reductions in preferred prey may result in declines in populations of more specialized corallivores while more generalist corallivores may increase. Corallivory may also make corals more susceptible to thermal stress and exacerbate bleaching. At local scales, overfishing depletes corallivorous fish stocks, reducing fish corallivory and bioerosion, whilst removing invertivorous fishes and allowing population increases in invertebrate corallivores (e.g., urchins, Drupella spp.). Interactive effects of local stressors, such as overfishing and nutrient pollution, can alter the effect of corallivory by increasing coral-algal competition and destabilizing the coral microbiome, subsequently leading to coral disease and mortality. Here, we synthesize recent literature of how global climate change and local stressors affect corallivore populations and shape the patterns and effect of corallivory. Our review indicates that the combined effects of corallivory and anthropogenic pressures may be underappreciated and that these interactions often drive changes in coral reefs on scales from ecosystems to microbes. Understanding the ecology of coral reefs in the Anthropocene will require an increased focus on how anthropogenic forcing alters biotic interactions, such as corallivory, and the resulting cascading effects on corals and reef ecosystems.
... In parallel to the decline in stem cell pools, regeneration rate was also reduced in sexual polyps. Trade-offs and allocation strategies between regeneration and other life-history traits such as sexual reproduction, asexual reproduction, growth or successive regeneration events were reported previously (Campbell, 1967;Gross, 1925;Kanajew, 1926;Martínez, 1996;Rinkevich & Loya, 1989;Tardent, 1963;Tardent & Tardent, 1956;Zattara & Bely, 2013). Regeneration is a somatic function that likely depends on the availability of cellular resources (stem cells). ...
Article
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1.In many basal metazoans both somatic and reproductive functions are performed by cellular derivatives of a single multipotent stem cell population. Reproduction can drain these stem cell pools, imposing a physiological cost with subsequent negative effects on somatic maintenance functions. 2.In the freshwater cnidarian Hydra oligactis both asexual (budding) and sexual reproductive modes (production of resting eggs) are present, and both of these are dependent on a common pool of interstitial stem cells. Resting eggs tolerate harsh abiotic conditions which neither the parental animals, nor asexual offspring can survive (e.g. freezing). Therefore, when facing unfavourable conditions and increased mortality risk, hydra polyps are expected to show higher level of differentiation of interstitial stem cells into germ cells (i.e. sexual reproduction) than other cell types needed for self‐maintenance or asexual reproduction. 3.Here, by comparing sexually and asexually reproducing individuals to non‐reproductives, we studied the physiological costs of reproduction (size of interstitial stem cell pools, their somatic derivatives and regeneration rate, which is dependent on these cell types) in H. oligactis polyps from a free‐living Hungarian population prior to the onset of winter. 4.Sexual individuals were characterized by significantly smaller interstitial stem cell pools, fewer nematoblasts involved in food capture and lower regeneration ability compared to non‐reproductives, but asexuals did not differ from non‐reproductive animals. We also found a negative correlation between germ cell counts and stem cell numbers in males (but not in females). 5.We suggest that the lower numbers of these cell types and lower regenerative ability in sexual individuals reflect a somatic cost of sexual reproduction. Our results also suggest that increased differentiation of stem cells into gametes might limit investment into somatic functions in hydra polyps. Exhaustion of cellular resources (stem cells) could be a major mechanism behind the extreme post‐reproductive senescence observed in this species. This article is protected by copyright. All rights reserved.
... Defending the perimeter requires the allocation of resources. The energy obtained from 96 photosynthesis-carried out by endosymbiotic algae-and heterotrophic feeding (Porter, 1976) 97 is then distributed throughout the colony using the coenosarc tissue (Rinkevich & Loya, 1989;98 Oren et al., 1997;Henry & Hart, 2005;Schweinsberg et al., 2015). As the colony's surface area 99 increases so does its potential for nutrient acquisition and distribution (Oren et al., 2001). ...
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Corals have built reefs on the benthos for millennia, becoming an essential element in marine ecosystems. Climate change and human impact, however, are favoring the invasion of non-calcifying benthic algae and reducing coral coverage. Corals rely on energy derived from photosynthesis and heterotrophic feeding, which depends on their surface area, to defend their outer perimeter. But the relation between geometric properties of corals and the outcome of competitive coral-algal interactions is not well known. To address this, 50 coral colonies interacting with algae were sampled in the Caribbean island of Curacao. 3D and 2D digital models of corals were reconstructed to measure their surface area, perimeter, and polyp sizes. A box counting algorithm was applied to calculate their fractal dimension. The perimeter and surface dimensions were statistically non-fractal, but differences in the mean surface fractal dimension captured relevant features in the structure of corals. The mean fractal dimension and surface area were negatively correlated with the percentage of losing perimeter and positively correlated with the percentage of winning perimeter. The combination of coral perimeter, mean surface fractal dimension, and coral species explained 19% of the variability of losing regions, while the surface area, perimeter, and perimeter-to-surface area ratio explained 27% of the variability of winning regions. Corals with surface fractal dimensions smaller than two and small perimeters displayed the highest percentage of losing perimeter, while corals with large surface areas and low perimeter-to-surface ratios displayed the largest percentage of winning perimeter. This study confirms the importance of fractal surface dimension, surface area, and perimeter of corals in coral-algal interactions. In combination with non-geometrical measurements such as microbial composition, this approach could facilitate environmental conservation and restoration efforts on coral reefs.
... Stoddart singled out the following areas of destruction and regeneration. In the previous recovery of the reef after the devastating hurricane in 1931 it took 10 to 20 years [20]. Stoddart [19] suggested that this time will take at least 20 -25 years. ...
Article
The current research is on fossil and recent coral reef ecosystem by traced processes of recovery after damage of different species. Examples of regeneration of artificially planted fragments of various species of scleractinian and reconstruction of part of the ecosystem of the reef are presented. Particular attention is drawn to the possibility of regeneration after typhoons and anthropogenic impacts.
... Fragmentation and removal of parts of the colony may reduce the reproductive ability of corals [51][52][53] . It is expected that the reduction in colony size due to fragmentation may affect reproductive output 54 . ...
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Present study consists reproductive behaviors of natural and transplanted acroporan corals in Vaan Island of the Gulf of Mannar. Single reproductive season per year and reproductive synchrony have been observed during the study period between 2008 and 2010 among the monitored Acropora species in both natural and transplanted colonies. Studies on the gametogenic cycle in three species, Acropora intermedia, A. nobilis and A. cytherea showed a single annual cycle of gametogenesis. Patterns of oogenesis, spermatogenesis and fecundity were almost similar between the three species and between natural and restored colonies. Multi-specific synchronous spawning was observed among the acroporans from both natural and restored colonies during March in all the study years. It was obvious that restored corals involve in the sexual reproduction effectively in Vaan Island exactly as the natural corals. Hence restoration through fragment transplantation is a successful option to employ in any degraded reef.
... Partial mortality is a constant source of sublethal stress for most colonial, sessile, and modular organisms (Mistri and Ceccherelli, 1995;Harmelin and Marinopoulos, 1994;Meszaros and Bigger, 1999;Jackson and Winston, 1981;Crawley, 1983). The ability of these organisms to repair wounds quickly and successfully can have major consequences for their survival, growth, and reproduction (Meesters et al., 1996;Rinkevich, 1996;Rinkevich and Loya, 1989;Rotjan and Lewis, 2005). Because of the ecological and adaptive importance of wound healing, conservation and monitoring efforts are beginning to explore recovery from artificial wounds as a proxy for organismal status and resilience to a variety of stressors (as in Fisher et al., 2007;Meesters et al., 1992). ...
... Fragmentation and removal of parts of the colony may reduce the reproductive ability of corals [51][52][53] . It is expected that the reduction in colony size due to fragmentation may affect reproductive output 54 . ...
... Paradigma yang diterima para ahli biologi karang yaitu reproduksi, pertumbuhan dan regenerasi pada karang merupakan proses "pembelanjaan energi yang mahal" dan harus dibelanjakan dalam alokasi sumberdaya di antara mereka (Rinkevich & Loya 1989, Harrison & Wallace 1990). Dengan demikian lokalisasi pertumbuhan pada bagian tertentu suatu koloni bertanggung jawab terhadap rendahnya fekunditas yang dikandung pada bagian tersebut atau dengan kata lain, alokasi sumberdaya untuk suatu fungsi biologi tertentu akan mengorbankan fungsi biologi lainnya. ...
... Reproduction in corals is sensitive to stressors that reduce energy available for processes not directly involved in survival over extended periods of time (Jokiel and Guinther 1978;Rinkevich and Loya 1989). Large reductions in the reproductive output of gorgonians suffering long-term infections of the fungus Aspergillus sydowii also support the hypothesis that energetic resources available for investment in gametogenesis are limited in chronically diseased corals . ...
... A majority of researchers believe that the attention of those engaged in the matter should be focused on the size of coral fragments, the season of their transplantation, orientation in the place of transplanting, and selection of substrate. These factors are most important for restoration of reproductive abilities of new coral colonies reared from fragments [27][28][29][30]. ...
... It is known that corals usually delay reproduction until they have achieved sufficient size (Szmant 1991). Furthermore, regeneration of damage lesions has been found to affect colony reproduction and fecundity due to a reallocation of energy resources (Rinkevich and Loya 1989;Van Veghel and Bak 1994). Hence, partial mortality may still have significant consequences for reproduction by reducing fecundity at the polyp level, reducing total colony reproductive output, or reducing colony size to below the reproductive threshold (Weil 2004). ...
Article
In the Florida Keys, increases in disease abundance and decreases in coral cover were documented during the 1990s, raising concern about the contribution of disease to coral decline. The prevalence, severity, and lethality of coral diseases in the Florida Keys was quantified by following the fate of over 500 diseased colonies in 14 stations from 2002-2004, and assessing changes via digital photography. Disease prevalence ranged from 4.0-8.2%, and incidence of new infections fluctuated considerably from year to year. Between 2002-2004, disease lethality was low: 1% of the population died, and 3% suffered partial mortality from disease. Between 2002-2003, tissue loss to disease was small (0.4 m 2), and monitored stations saw no significant changes in coral cover. However, unexpected long-term impacts of disease could be seen because 1) diseases targeted larger sized (more fecund) colonies, and 2) four of the most important reef building species accumulated most of the tissue loss.
... for the 11 most abundant species [34]. High rates of background partial mortality within a population may lead to a decline in population densities through time because they can result in reduced colony growth [32,43], reproductive output [44,45], and reduced colony size [46] of individuals. Cumming [47] showed that recent injury can predict colony fate even more than colony size. ...
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Even in the absence of major disturbances (e.g., cyclones, bleaching), corals are consistently subject to high levels of background mortality, which undermines individual fitness and resilience of coral colonies. Partial mortality may impact coral response to climate change by reducing colony ability to recover between major acute stressors. This study quantified proportion of injured versus uninjured colonies (the prevalence of injuries) and instantaneous measures of areal extent of injuries across individual colonies (the severity of injuries), in four common coral species along the Great Barrier Reef in Australia: massive Porites, encrusting Montipora, Acropora hyacinthus and Pocillopora damicornis. A total of 2,276 adult colonies were surveyed three latitudinal sectors, nine reefs and 27 sites along 1000 km2 on the Great Barrier Reef. The prevalence of injuries was very high, especially for Porites spp (91%) and Montipora encrusting (85%) and varied significantly, but most lay at small spatial scales (e.g., among colonies positioned <10-m apart). Similarly, severity of background partial mortality was surprisingly high (between 5% and 21%) but varied greatly among colonies within the same site and habitat. This study suggests that intraspecific variation in partial mortality between adjacent colonies may be more important than variation between colonies in different latitudinal sectors or reefs. Differences in the prevalence and severity of background partial mortality have significant ramifications for coral capacity to cope with increasing acute disturbances, such as climate-induced coral bleaching. These data are important for understanding coral responses to increasing stressors, and in particular for predicting their capacity to recover between subsequent disturbances.
... Potential demographic consequences of injury include reduced survivorship, growth and reproduction, colony shrinkage and increased fragmentation. Experimental studies have demonstrated reduced growth (Bak 1983, Meesters et al. 1994) and reproduction (Rinkevich & Loya 1989, Hall 1997a). Fragmentation has been linked with injury because areas devoid of live tissue become infested by boring sponges and prone to breakage (Highsmith 1982). ...
Article
Tissue injury, in which the skeleton is stripped of living tissue, is common in reef-building corals and has potentially important demographic consequences. To examine the significance of tissue injury for natural populations, I monitored 1627 colonies in 30 taxa of Indo-Pacific branching corals at 3 to 5 mo intervals over a 2 yr period. Recent injury (inflicted within the few days prior to censusing) was a highly significant predictor of colony fate within 3 to 5 mo, for acroporid corals with small, compact branches (hispidose and corymbose growth forms). In contrast, colony size was not a significant predictor of fate for these corals after recent injury was included in the models. Both recent injury and colony size were significant predictors of fate for pocilloporids (small bushy growth form). Neither were good predictors of fate for arborescent acroporids (large, widely-spaced branches), even though recent injury was up to 3 times more common in these corals. Old injury (inflicted several weeks or more prior to censusing) covering > 5% of the colony was a highly significant predictor of colony death within 3 to 5 mo for corymbose species. Colonies with both old and recent injuries were highly likely to die: 33 and 54% of colonies in separate censuses died within 3 mo. The predictive power of recent injury implies chronic or repetitive tissue loss and prolonged decline, since most recent injuries were small (<30 cm(2)) and did not account for the colony decline per se. Since colony size was not as good a predictor of colony fate as recent injury for small-branched acroporids, size-based population models for these corals may be improved by incorporating tissue injury as an indicator of colony condition.
... In addition, the partial mortality induced by the disease may have further detrimental effects on competitive abilities, growth, reproduction and resistance of the diseased colonies to injuries and to other diseases. For example, regeneration of lesions has been found to affect colony growth (Bak 1983, Meesters et al. 1994) and reproduction (Rinkevich & Loya 1989, Van Veghel & Bak 1994. Szmant-Froelich (1985) found that large colonies had higher reproductive success than small. ...
Article
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In July 1997, a coral disease called plague appeared on the coral reefs of St. Lucia, West Indies. A survey of 6 coral species on 3 reefs in March 1998 revealed an overall disease incidence of 11%. Montastraea faveolata and Colpophyllia natans were the most affected species with 19% and 13% of colonies infected respectively. Disease frequency was independent of coral density and the distribution of the disease among 5 x 5 m plots was not clumped, suggesting that proximity to affected colonies did not increase probability of infection. Among-colony size variation in disease incidence and tissue mortality indicates that small coral colonies are more likely to escape infections than large colonies, but once infection occurs, small colonies will experience faster colony mortality. This suggests that the effect of the disease on coral population structure will be greater total mortality in small colonies relative to large colonies, at least over short-time scales. At the most severely affected site, it was estimated in March 1998 that plague had killed 6.6% of living coral during the preceding 8 months. Despite these losses being small compared to losses from other well-known disturbances on coral reefs, such as hurricanes, the disease selectively impacted 2 major reef frame builders. More than 90% of the estimated tissue loss was accounted for by Montastraea faveolata and Colpophyllia natans combined. In addition, infected colonies continued to lose living tissue from February 1998 to October 1998, with no new coral recruitment observed on the substrate opened up by the disease. Of the colonies infected in February 1998, 28% still showed signs of disease in October 1998, Over the long term, if losses are sustained at such rates, plague could cause complete mortality in large colonies and could progressively deplete two of the most important reef frame builders in some of the richest and most visited coral reefs of St. Lucia.
... Such complexity and lack of data on reproductive energy demand in relation to Me history aspects, environmental variability or colony structure are reasons for our inability to make generalizations. Data available on energetic relationships between reproduction and Life history aspects include: growth rate (Chornesky & Peters 1987, Richmond 1987 ), regeneration and survival (Babcock 1988, Rinkevich & Loya 1989, Van Veghel & Bak 1994), and recruitment success (Richmond 1981, Sammarco & Andrews 1989). Studies on environmental conditions and coral reproduction have reported a negative relationship between fecundity and external stress factors (Kojis & Quinn 1984, Jokiel 1985, Rinckevich & Loya 1987, Tomascik & Sander 1987, Szmant & Gassman 1990, Glynn et al. 1991). ...
... It has been suggested for a variety of solitary and colonial taxa that regeneration and sexual reproduction compete for the available energy resources (Kojis and Quinn 1985, Szmant-Froelich 1985, Chapman et al. 1990, Harrison and Wallace 1990, Babcock 1991, Hall and Hughes 1996, Rinkevich 1996). The negative effect of injury on reproduction has already been demonstrated in many clonal organisms (bivalves, Trevaillion et al. 1970; ectoprocts and sponges, Jackson and Palumbi 1979; zoanthids, Karlson 1983) including corals (Rinkevich and Loya 1989, Van Veghel and, Ward 1995, Hall 1997). However, in contrast to previous studies in corals, which concluded that injury reduces coral fecundity only in the vicinity of the lesion (Wahle 1983, Van Veghel and Bak 1994, Hall 1997), our results show that the negative effect of injury on fecundity in F. favus may extend up to a distance of 15 cm from the injured area (Fig. 5c). ...
Article
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Modular organisms consist of repeated building blocks. An important consequence of modularity may be reflected in the ability of a colony to continually reallocate priority of resource transport among its units in response to stress. Hermatypic corals, the main organisms constructing tropical reefs, are prone to damage by a multitude of agents. Since colonization of lesions bY competitors is a potent threat to colonial organisms, fast recovery is an important component of colony survival. Previous regeneration studies have claimed that the energy requirements of this essential process are fueled only by the polyps directly bordering the injured area. This "localized regeneration hypothesis" rejects the necessity for wide colony integration during regeneration and sees no advantage to large colony size. The objective of the present study was to test an alternative regeneration hypothesis that argues, in contrast, that injury repair (i.e., closure of lesions by newly formed tissues) in corals may require extended colony integration (i.e., internal translocation of resources from sites of acquisition to sites of maximal demand). To test our hypothesis we examined: (1) the relationship between colony size and percentage recovery of lesions differing in size and shape; and (2) the effect of different sized lesions on the fecundity of polyps located at increasing distances from the lesion site. Both experiments were conducted on the common, spherically shaped coral Favia favus in the Red Sea near Eilat, Israel. The relatively small lesions (< 1 cm 2) were the only ones to support the localized regeneration hypothesis, since their recovery was unaffected by colony size. However, the two larger lesion types (approximate sizes of 2 cm 2 and 3 cm 2) confirmed the importance of large colony size for achieving fast recovery. In the second experiment we found that small lesions, repeated monthly, caused only a localized reduction in fecundity, while larger monthly repeated lesions caused significant reductions in fecundity up to a distance of 15 cm away from their site. Both experiments indicate that regeneration from injury may require an extended magnitude of energy integration throughout the colony, and that the extent of this integration is regulated by the colony in accordance with lesion characteristics. It is also concluded that in long-lived organisms such as corals, there is a priority of energy allocation to recovery rather than to reproduction. Our findings reveal the existence of injury thresholds within a colony that determine energy allocation and intra-colonial translocation of energy products toward regions of maximal demand. We suggest that such injury thresholds may characterize many other coral species and that colony integration during stress is a basic life-preserving ability and one of the most important advantages of clonal and colonial organisms.
... 4 in our paper clearly refutes this marginal claim. Overlooking this parameter is particularly surprising, since Rinkevich himself used this very parameter to ascertain reproductive fitness of this coral species in previous publications (Rinkevich and Loya, 1989). Nevertheless, and perhaps not surprisingly, he failed to do so in his recent publication concerning the reproduction of S. pistillata in the vicinity of the fish farms (see BEA). ...
... The experiment was carried out in the breeding season and, hence, tissue regeneration was slow because energy was diverted to reproduction. Rinkevich and Loya (1989) reported that the regeneration of broken branches of Stylophora pistillata also resulted in a significant decrease in fecundity, lasting at least 19 months after regeneration started. ...
... Regeneration plays a fundamental role in colony survival. Because it requires energy, regeneration affects growth (Bak 1983), reproduction (Rinkevich & Loya 1989), and possibly also resistance to diseases (Bak & Criens 1981), as well as competitive ability. The regen-eration of a wound starts by the formation of a new tissue layer that is formed by the surrounding tissue. ...
Article
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The influence of colony tissue regeneration on growth was studied in the reef-building coral Montastrea annularis (Ellis and Solander, 1786) by buoyant weighing. Experimental corals consisted of a series of cores, 2 cores from each of 20 colonies. One core of each pair was artificially damaged by removing approximately 1 cm(2) of tissue and skeleton, the other served as a control for normal growth. Growth was measured as calcification, i.e. mg calcium carbonate production. There was a linear relation between growth and solar radiation. Variation in growth, calculated before the regeneration experiment, was insignificant between cores from the same colony but significantly different between colonies. Growth was reduced in damaged cores when compared to controls. Calcification decreased immediately upon damage and remained reduced during the whole study (56 d). Lesions caused by physical damage did not always close completely. We propose and test a model that describes regeneration in terms of closure of lesions. This model includes an asymptote in an exponentially decreasing function. Calcification remained suppressed after regeneration slowed down and lesions were still not completely closed, probably because of the formation of polyps and skeletal features in the new tissue. We suggest regeneration to be fuelled by polyps and tissue directly bordering the damaged area. Also, successful regeneration depends on the amount of tissue bordering a lesion and not on colony size.
... Energetic trade-offs between reproduction, tissue growth and repair in corals are not fully understood. Tissue injuries may create shifts in energy allocation within a single coral colony (reviewed in Rinkevich & Loya 1989), or alternately in the utilization of circulating stem cells that are available in limited supply (Rinkevich 1996). Between-colony competition likely imposes energetic demands on corals, which in turn cause a trade-off between the allocation of energy to aggressive interactions versus colony growth and reproduction (Tanner 1997). ...
Article
Outcomes of competition between corals vary temporally and spatially, and depend in part on the agonistic mechanism involved. Competition may impact coral growth, reproduction and energy reserves, however few experimental studies have quantified these effects. We conducted a 1 yr laboratory experiment on competition between 2 massive corals, Platygyra daedalea and Favites complanata. Colonies of P. daedalea developed sweeper tentacles and extensively damaged the F. complanata colonies, causing them to loose ca. 55% of their soft tissue and eventually killing 30% of F. complanata colonies. Skeletal accretion rate varied widely among corals within each treatment, and correlated negatively with the percent tissue damaged on competing colonies of F. complanata. On isolated control colonies, sweeper tentacles developed randomly throughout the year, and then reverted back to feeding tentacles. They appeared to serve as probes to detect the approach of competitors. Development of sweeper tentacles is a powerful aggressive/defensive mechanism that may enable brain corals to dominate some reef areas in the Indo-Pacific region.
... Regeneration plays a fundamental role in colony survival. Because it requires energy, regeneration affects growth (Bak 1983), reproduction (Rinkevich & Loya 1989), and probably also resistance to diseases and competitive ability. ...
Article
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We assessed the influence of coral 'bleaching' on the regenerative potential of corals in a series of field experiments with artificial lesions on colony surfaces. These lesions were made during a mass bleaching event in 1990 on normally coloured and bleached colonies of 3 species: Montastrea annularis, the main reef-building coral in the Caribbean, Porites astreoides, a relatively common and opportunistic coral, and Meandrina meandrites, a species common on the deeper reef slope. Regeneration characteristics studied included rate of tissue growth, recovery of lesion area and restoration of tissue colour. Tissue regeneration rates were lower in all species when comparing bleached with unbleached colonies, although there were species-specific differences. Long-term exposure to increased temperature probably affects regeneration, as bleached colonies of P. astreoides in the thermal effluent of a power plant regenerated as rapidly as normal colonies situated upcurrent and downcurrent of the power plant. Colour recovery of bleached colonies was closely associated with seawater temperature. When ambient seawater temperature started to decrease (<30-degrees-C), colonies began to regain their normal colour. Colour restoration was slowest in M annularis. Normal colour of coral colonies in the power plant effluent returned at the same time, but at temperatures 2-degrees-C higher than elsewhere on the reef. Mortality on bleached colonies of M annularis and P. astreoides was higher than on normal colonies 131 d after the start of the regeneration experiments. Large parts of the colony died when the tissue was already regaining its normal colour and lesion regeneration was almost completed. Mortality was most severe for colonies in the effluent. We recorded the sensitivity of coral species to bleaching in surveys over the reef downcurrent and upcurrent of the power plant. These showed that community structure and colony condition on the shallow terrace downcurrent differed noticeably from the upcurrent site. Downcurrent, species diversity was lower mainly because of high densities of Porites astreoides and Diploria strigosa. Bleaching was also more prominent downcurrent. The decreased regeneration rates and increased mortality of bleached colonies (in Montastrea annularis in 30 % of the colonies), during and subsequent to bleaching, clearly indicate the increased vulnerability of coral reefs during periods of thermal stress. Coral reef management should consider limiting activities resulting in small lesions under periods of increased environmental stress.
... 4 in our paper clearly refutes this marginal claim. Overlooking this parameter is particularly surprising, since Rinkevich himself used this very parameter to ascertain reproductive fitness of this coral species in previous publications (Rinkevich and Loya, 1989). Nevertheless, and perhaps not surprisingly, he failed to do so in his recent publication concerning the reproduction of S. pistillata in the vicinity of the fish farms (see BEA). ...
... It has been suggested for a variety of solitary and colonial taxa that regeneration and sexual reproduction compete for the available energy resources (Kojis and Quinn 1985, Szmant-Froelich 1985, Chapman et al. 1990, Harrison and Wallace 1990, Babcock 1991, Hall and Hughes 1996, Rinkevich 1996). The negative effect of injury on reproduction has already been demonstrated in many clonal organisms (bivalves, Trevaillion et al. 1970; ectoprocts and sponges, Jackson and Palumbi 1979; zoanthids, Karlson 1983) including corals (Rinkevich and Loya 1989, Van Veghel and, Ward 1995, Hall 1997). However, in contrast to previous studies in corals, which concluded that injury reduces coral fecundity only in the vicinity of the lesion (Wahle 1983, Van Veghel and Bak 1994, Hall 1997), our results show that the negative effect of injury on fecundity in F. favus may extend up to a distance of 15 cm from the injured area (Fig. 5c). ...
... Asexual reproduction by fragmentation is an important mode of colony propagation for many scleractinians (Highsmith, 1982). However it can be maladaptive (Smith and Hughes, 1999), for instance, when repeated physical disturbances result in energy being diverted away from reproduction into tissue repair (SzmantFroelich, 1985; Kojis and Quinn, 1985; Epstein et al., 2001; Rinkevich and Loya, 1989; Oren et al., 2001). The potential for such negative effects complicate fragmentation/transplantation experiments aimed at examining variation in reproductive success in corals. ...
Article
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Ramets of the coral Goniopora columna (Dana, 1846–1849) were excised from five large colonies at one site south of mainland Singapore and transplanted within the same site and depth (2.3 m); to a more disturbed site closer to the mainland at a similar depth (2.2 m); and to greater depth at both sites (8.9 m and 7.6 m, respectively). Ramets were left for one year, then harvested before the predicted spawning periods. Fecundity (average number of oocytes polyp), reproductive effort (average oocyte diameter), and polyp tissue diameter were compared among transplanted clonal ramets, control ramets, and parent colonies. Typically, fragmentation in corals results in reductions in reproductive output because energy is diverted into tissue repair, but in this case reproductive effort and fecundity did not change significantly when ramets were transplanted within the same site and depth. Clonal fragments transplanted to the more disturbed site had significantly fewer oocytes, smaller oocytes and smaller polyps after one year suggesting that energy was diverted away from normal functions in response to stressors in the new environment. Ramets transplanted to 7–9 m bleached at both sites after acute periods of reduced irradiance, resulting in high mortality and reproductive failure in surviving ramets.
... Corals have limited energy stores that are partitioned among growth, regeneration, and reproduction (Bak 1983, Harrison & Wallace 1990. Energetically expensive reproduction is often compromised in favor of tissue regeneration processes (Szmant-Froelich 1985, Rinkevich & Loya 1989, Harrison & Wallace 1990, Van Veghel & Bak 1994. Van Veghel & Bak (1994) conducted a careful study of Montastraea annularis, M. faveolata, and M. franksi and found that colonies inflicted with artificial lesions ~10 wk prior to spawning showed a reduced reproductive effort (fertility, fecundity, fewer gonads per polyp, and fewer eggs per gonad) in neighboring polyps, compared to polyps located 20 cm away from the lesion area on the same colony. ...
Article
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It is well known that herbivores have numerous and diverse impacts on plant and algal fitness, community structure and ecosystem function. The importance of corallivory as a selective force, however, has been underestimated. Corallivores, or consumers of live coral tissue, employ a wide variety of feeding strategies and can be obligate or facultative coral feeders. Our literature review reveals a complex array of corallivores across the globe, represented by 11 families of fishes and 5 invertebrate phyla and totaling over 160 species known to consume scleractinian corals world- wide. Importantly, although these corallivores span a wide taxonomic range, we found that they have been reported to feed on relatively few genera of hard corals, specifically, on only 28 scleractinian genera worldwide. Damage by corallivores ranges from minor to lethal, but there is a growing body of evidence to support that even limited removal of tissue or skeletal structures has growth and/or fit- ness consequences for a scleractinian coral colony. In light of increasing reef stressors and diminish- ing coral populations, we suggest that the role of corallivores in reef trophodynamics is more complex than appreciated previously.
... While growth and survival are important in the evaluation of restoration efforts, they are not the only criteria by which reef restoration success should be measured. For example, other areas that require further research include the optimal density and arrangement for coral transplantation, the most appropriate species-or combinations of species-for transplantation, the most suitable time of year to transplant and the effect of fragmentation and transplantation on reproduction (Rinkevich & Loya 1989;Dizon & Yap 2005;Okubo et al. 2005;Sleeman et al. 2005). ...
Article
Transplantation of coral fragments is seen as a potential method to rapidly restore coral cover to areas of degraded reef; however, considerable research is still needed to assess the effectiveness of coral transplantation as a viable reef restoration tool. Initially, during restoration efforts, coral transplants are attached artificially. Self-attachment (i.e., growth of coral tissue onto the substrate) provides a more secure and lasting bond, thus knowledge about self-attachment times for corals is of importance to reef restoration. While it is known that coral fragments may generate new tissue and bond to substrata within a few weeks of transplantation, surprisingly little is known about the speed of self-attachment for most species. Two independent experiments were carried out to examine the self-attachment times of 12 scleractinian and one non-scleractinian coral species to a natural calcium carbonate substrate. The first experiment examined times to self-attachment in 11 species of differing morphologies from seven families over approximately 7 months, whereas the second experiment examined three fast-attaching Acropora species over approximately 1 month. In the first experiment, the branching species Acropora muricata had a significantly faster self-attachment time compared to all other species, while Echinopora lamellosa had the slowest self-attachment time. For the second experiment, A. muricata was significantly slower to self-attach than Acropora hyacinthus (tabular) and Acropora digitifera (corymbose-digitate). The results suggest that a combination of factors including growth rates, growth form and life history may determine how quickly fragments of coral species self-attach after fragmentation and transplantation.
... Regeneration frequently proceeds at the expense of sexual reproduction in sponges and corals. Regenerating individuals have reduced fecundity or remain sexually infertile, while others may prematurely shed inviable sexually-derived larvae (REISWIG, 1973;TUNNICLIFFE, 1981;WAHLE, 1983b;KOJIS and QUINN, 1985;RICHMOND, 1987;RINKEVICH and LOYA, 1987;HOPPE, 1988;RINKEVICH and LOYA, 1989;GUZMÁN and HOLST, 1993;VAN VEGHEL and BAK, 1994;SMITH and HUGHES, 1999;LIRMAN, 2000b;OREN et al., 2001;HENRY et al., 2003;HENRY and KENCHINGTON, 2004). HALL and HUGHES (1996) considered that unusually large but sexually immature corals may represent evidence of altered investments in sexual reproduction owing to the energetic demands imposed by previous injuries. ...
Article
The ability of bottom-dwelling marine epifauna to regenerate injured or lost body parts is critical to the survival of individuals from disturbances that inflict wounds. Numerous studies on marine sponges (Phlyum Porifera) and corals (of the orders Scleractinia and Alcyonacea) suggest that regeneration is limited by many intrinsic (individual-dependent) and extrinsic (environment-dependent) factors, and that other fife history processes may compete with regeneration for energetic and cellular resources. We review how intrinsic (size, age, morphology, genotype) and extrinsic (wound characteristics, water temperature, food availability, sedimentation, disturbance history, selection) factors limit regeneration in sponges and corals. We then review the evidence for impaired somatic growth and sexual reproduction, and altered outcomes of interactions (anti-predator defenses, competitive abilities, self- and non-self recognition abilities) with other organisms in regenerating sponges and corals. We demonstrate that smaller, older sponges and corals of decreasing morphological complexities tend to regenerate less well than others, and that regeneration can be modulated by genotype. Large wounds with small perimeters inflicted away from areas where resources are located tend to be regenerated less well than others, as are injuries inflicted when food is limited and when the animal has been previously or recently injured. We also demonstrate that regeneration strongly impairs somatic growth, reduces aspects of sexual reproduction, and decreases the ability for sponges and corals to defend themselves against predators, to compete, and to recognize conspecifics. Effects of limited regeneration and impaired life histories may manifest themselves in higher levels of biological assembly e.g., reduced accretion of epifaunal biomass, reduced recruitment and altered biotic associations, and thus affect marine community and ecosystem recovery from disturbances.
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Abstract: Histological investigations for the reproductive conditions of Acropora hemperchii and Pocillopora verrucosa in the onshore reefs at site 1, which impacted by oil pollution, and site 2 which , impacted by phosphate shipping, and those in the offshore ones at sites 3 and 4 at, respectively; each was impacted by diving activities, were studied. Both of the given species was found to be hermaphrodite broadcast spawners annually. The gonads of P. verrucosa were found on stalks arising from the column wall; but in A. hemperchii, the gonads fill the entire mesenteries. Moreover, there are pronounced differences in the timing and length of gametogenesis and spawning of A. hemperchii and P. verrucosa between different geographic regions. A. hemperchii showed the least productivity at the effected with oil pollution at site 1, represented by: 55 % of mature oocytes and an additional 38 % of immature oocytes. P. verrucosa declined to 46 %of mature oocytes and 50% of immature oocytes. The highest productivity of A. hemperchii was represented by 90% matures,3 % immatures and 7% empty and for Pocillopora verrucosa increased to 92% matures, 2% immature, and 5 % empty at the offshore site 2, the site impacted by eutrophication as a result of phosphate shipment. The spawning season of A. hemperchii in the northern Red Sea at site 1, is two months later than in the southern site 2, where A. hemperchii starts spawning in May and extended to July. However, P. verrucosa begins one month early than A. hemperchii at the same site, and extends for three months also from April to June. Moreover, spawning of P. verrucosa at site 2 is one month early than at its spawning at site 1. While, P. verrucosa at sites 3&4 starts spawning at April and extended for two months only.
Article
Changes in coral abundance are typically used to assess coral mortality rates following major acute disturbances on coral reefs. However, coral abundance metrics do not consider partial mortality (hereafter injury) or background mortality occurring independently of major disturbances. As such we have little understanding about the influence of major versus routine disturbances on coral assemblages. We compared the incidence of recent (in the last 4-6 wk) coral injury (tissue loss affecting 5-99% of the colony) among different genera and size classes (<5, 5-40 and >40 cm diameter) at 16 reefs throughout Australia’s Coral Sea Marine Park before and during a mass bleaching event in 2020. Despite mass bleaching, the overall incidence of recent injury was consistent and low (<5%) across both years. However, there were marked inter-annual changes in the taxonomic hierarchy for incidence of recent injury. Interestingly, massive Porites exhibited higher incidence of recent injury during the 2020 mass bleaching whilst Acropora and Pocillopora exhibited less recent injury during mass bleaching compared to surveys conducted before the bleaching in 2018-2019. Incidence of recent injury increased with colony size, highlighting the vulnerability of large colonies and possibility of shifts in community size structure. Continued assessment of recent injuries as coral communities recover will be critical to understand their vulnerability to future disturbances and changing environmental conditions.
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In many basal metazoans both somatic and reproductive functions are performed by cellular derivatives of a single multipotent stem cell population. Reproduction can drain these stem cell pools, imposing a physiological cost with subsequent negative effects on somatic maintenance functions. In the freshwater cnidarian Hydra oligactis both asexual (budding) and sexual reproductive modes (production of resting eggs) are present, and both of these are dependent on a common pool of interstitial stem cells. Resting eggs tolerate abiotic conditions which neither the parental animals, nor asexual offspring can survive (e.g. freezing). Therefore, when facing unfavorable conditions and increased mortality risk, hydra polyps are expected to show higher differentiation of interstitial stem cells into germ cells (i.e. sexual reproduction), compared to other cell types needed for selfmaintenance or asexual reproduction. Here, by comparing sexually and asexually reproducing individuals to non-reproductives, we studied the physiological costs of reproduction (size of interstitial stem cell pools, their somatic derivatives and regeneration rate, which is dependent on these cell types) in H. oligactis polyps from a free-living Hungarian population prior to the onset of winter. Sexual individuals (but not asexuals) were characterized by significantly smaller interstitial stem cell pools, fewer somatic derivatives (nematoblasts involved in food capture) and lower regeneration ability compared to non-reproductives. We also found a negative correlation between germ cell counts and stem cell numbers in males (but not in females). These results show that the physiological costs of reproduction are higher for sexual individuals. They also suggest that increased differentiation of stem cells into gametes might limit investment into somatic functions in hydra polyps. Exhaustion of cellular resources (stem cells) could be a major mechanism behind the extreme post-reproductive senescence observed in this species.
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As coral reefs continue to decline globally, coral restoration practitioners have explored various approaches to return coral cover and diversity to decimated reefs. While branching coral species have long been the focus of restoration efforts, the recent development of the microfragmentation coral propagation technique has made it possible to incorporate massive coral species into restoration efforts. Microfragmentation (i.e., the process of cutting large donor colonies into small fragments that grow fast) has yielded promising early results. Still, best practices for outplanting fragmented corals of massive morphologies are continuing to be developed and modified to maximize survivorship. Here, we compared outplant success among four species of massive corals (Orbicella faveolata, Montastraea cavernosa, Pseudodiploria clivosa, and P. strigosa) in Southeast Florida, US. Within the first week following coral deployment, predation impacts by fish on the small (<5 cm 2) outplanted colonies resulted in both the complete removal of colonies and significant tissue damage, as evidenced by bite marks. In our study, 8-27% of fragments from four species were removed by fish within one week, with removal rates slowing down over time. Of the corals that remained after one week, over 9% showed signs of fish predation. Our findings showed that predation by corallivorous fish taxa like butterflyfishes (Chaetodontidae), parrotfishes (Scaridae), and damselfishes (Pomacentridae) is a major threat to coral outplants, and that susceptibility varied significantly among coral species and outplanting method. Moreover, we identify factors that reduce predation impacts such as: (1) using cement instead of glue to attach corals, (2) elevating fragments off the substrate, and (3) limiting the amount of skeleton exposed at the time of outplanting. These strategies are essential to maximizing the efficiency of outplanting techniques and enhancing the impact of reef restoration.
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Studies on the population and community dynamics of scleractinian corals typically focus on catastrophic mortality associated with acute disturbances (e.g., coral bleaching and outbreaks of crown-of-thorns starfish), though corals are subject to high levels of background mortality and injuries caused by routine and chronic processes. This study quantified prevalence (proportion of colonies with injuries) and severity (areal extent of injuries on individual colonies) of background mortality and injuries for four common coral taxa (massive Porites, encrusting Montipora, Acropora hyacinthus and branching Pocillopora) on the Great Barrier Reef, Australia. Sampling was conducted over three consecutive years during which there were no major acute disturbances. A total of 2276 adult colonies were surveyed across 27 sites, within nine reefs and three distinct latitudinal sectors. The prevalence of injuries was very high (>83%) across all four taxa, but highest for Porites (91%) and Montipora (85%). For these taxa (Montipora and Pocillopora), there was also significant temporal and spatial variation in prevalence of partial mortality. The severity of injuries ranged from 3% to more than 80% and varied among coral taxa, but was fairly constant spatially and temporally. This shows that some injuries have considerable longevity and that corals may invest relatively little in regenerating tissue over sites of previous injuries. Inter-colony variation in the severity of injury also had no apparent effect on the realized growth of individual colonies, suggesting that energy diverted to regeneration has a limited bearing on overall energetic allocation, or impacts on other life-history processes (e.g., reproduction) rather than growth. Establishing background levels of injury and regeneration is important for understanding energy investment and life-history consequences for reef-building corals as well as for predicting susceptibility to, and capacity to recover from, acute disturbances.
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Life history processes of scleractinian corals play a crucial role in the development and maintenance of coral reefs. Factors such as recruitment, survival, and longevity of the corals determine the reef characteristics. These factors depend on environmental parameters as well as on physiological intrinsic traits of the primary reef builders, the scleractinian corals. Despite intensive studies, we still lack a good understanding of the mechanisms of stress response and recovery of corals at the organism and community levels (Buddemeier and Smith 1999). Buddemeier and Smith (1999) proposed that “in any environment there exists a “hyperspace” where interacting variables create a multidimensional space of optimal or acceptable conditions in terms of organism response”. As long as all or most of the variables are within the central core of this hyperspace, the organisms may display resilience. An organism or community may remain viable or seem to thrive even in response to environmental change that reduces its volume of optimal hyperspace (Buddemeier and Smith 1999). Further change, however slight, may then push them over the threshold causing a disintegration of adaptive responses. If we perceive a coral as a unit where resources are limited and partitioned to a multitude of life functions (see Fig 10.1), we can envision the mechanism in which the “hyperspace” may act upon it. Corals that suffer injury to tissue, suffer damage to reproduction, growth and the ability to withstand stress, and by definition exist in a shrunken hyperspace. These corals will also ultimately be more susceptible to disease.
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Corals vary in their ability to resist breakage from mechanical force. Biomechanical theory predicts that resistance to mechanical injury should be determined by the morphological properties of coral colonies. However, a general model to predict resistance of corals to mechanical impact from structural characteristics has been lacking. In order to determine whether resistance can be predicted from colony structure, I use a suite of 6 variables-skeletal density, branch thickness, branch spacing, branch length, colony height and colony (projected) area-to quantitatively describe the structure of colonies from 12 species of scleractinian coral. Experimental analysis of the resistance of colonies from each species, which range from massive and tabular growth forms to various branching models, showed that damage susceptibility varied widely. Resistance ranged from very high for species such as Leptoria phrygia, Porites cylindrica and Porites spp. in which there was no reduction in colony area as a result of mechanical stress, to low in species such as Pocillopora damicornis, Seriatopora hystrix and Montipora sp, which suffered over 60% damage. Regression tree analysis revealed a strong relationship between resistance and colony morphology, producing a model which accounted for 76% of the variation in resistance using only 4 of the structural variables: colony height, branch thickness, branch spacing and colony area. Skeletal density and branch length were relatively unimportant in determining susceptibility to breakage. The regression tree model showed potential for development as a predictive tool in reef management, as it was able to predict susceptibility to breakage of corals using 4 easily measured morphological variables with 57% reliability (estimated by cross-validation).
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In their recent paper, Bongiorni et al. (2003a) claim that nutrients released from fish farms (Fig. 1A) at Eilat (Red Sea) 'may not necessarily lead to the demise of coral reefs as is commonly presumed' (p. 137). Their study, aimed at assessing the 'impacts of a commercial fish farm near Eilat, Israel, on some life-history para-meters of selected coral species' (p. 138), showed that 7 mo after experimentally suspending branches of Acropora eurystoma corals at the fish farm site, there was an accelerated growth rate and weight increase in these branches compared with corals at a reference site across from the Interuniversity Institute (IUI). They further showed that in colonies of the coral Stylophora pistillata growing on artificial substrata at depths of 10 to 17 m close to the fish farm, there were significantly more polyps with developing testes and oocytes at the fish farm site, compared with the reference site. It is important to critically examine this work since the authors conclude that their results 'challenge the pre-vailing notion … that nutrient effluents released from intensive net-pen fish-farms in the Gulf of Eilat is the major detrimental factor contributing to the degrada-tion of the coral reefs along the Israeli coast' (Bongiorni et al. 2003a, p. 143). We contend that the methodology and experimental design used in that study are unsuit-able for obtaining a correct assessment of the impact of commercial fish farms. Moreover, we claim that their results actually attest to the opposite, i.e. to the detri-mental effect that eutrophication has on the coral species examined. Experimental design and methodology. In their study, Bongiorni et al. (2003a) transplanted coral frag-ments of the branching coral Acropora eurystoma onto PVC plates suspended at a depth of 6 m in the vicinity of the fish farm. The seafloor at that site is approxi-mately 22 m deep. The reasoning behind mooring the plates at such a shallow depth in the water column was to exclude 'possible impacts from resuspended bottom sediments' (p. 139). But by doing so they also pre-vented the coral fragments from 'experiencing' the true concentrations of both particulate and dissolved matter that naturally growing colonies on the sea floor experience. Much of the nutrients released from the fish farms is in the form of particulate materials that accumulate on the sediment. Unfortunately, Bongiorni et al. (2003a) failed to provide an accurate description of the benthic environment under the fish cages; fish feces and sinking food particles have an adverse effect on natural communities, making the benthic habitat anaerobic and supporting the growth of bacterial mats (Fig. 1A,B), protozoans and of a sulfide-resistant com-munity. Sinking particulate matter can be resuspended by water motion, where it can be colonized by muco-polysaccharide-producing diatoms and microbes (Fabricius & Wolanski 2000, Wolanski et al. 2003). By settling on the coral colonies this particulate matter may cause polyp suffocation and disease (Fabricius & Wolanski 2000). In addition, resuspension of this par-ticulate matter increases the nutrient levels that nat-ural colonies encounter at both sites, compared to the nutrient levels experienced by the colonies grown by Bongiorni et al. at 6 m depth. The experimental design presented in their paper is thus inadequate for assess-ing nutrification effects on natural coral communities. The authors state that 'During the experiment, algae and encrusting invertebrates were removed from the PVC plates on a monthly basis' (Bongiorni et al. 2003a, p. 139; emphasis added). In doing so, they artificially eliminated one of the most important environmental effects of fish farms on corals: stimulating growth of benthic algae (Fig. 1C) that may smother the corals (Fig. 1D) (Laws 1992, Genin et al. 1995, McCook 1999).
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Three types of injuries commonly generated by biotic and abiotic agents on the periphery of coral colonies are tissue mortality, scraping injuries and breakage. To determine if recovery (amount of regeneration) was affected by injury type, colonies of Acropora hyacinthus and Montipora tuberculosa were inflicted with tissue, scraping and breakage injuries and their recovery monitored over 24 days. The influence of the tissue available for regeneration (i.e. the zone of tissue of bordering or partially injured polyps from which regeneration can occur) and the amount of settlement of algae on recovery was also measured for the different injury types and species. Overall, the amount of regeneration was up to four times greater for a scraping injury than tissue mortality and breakage. Furthermore, the amount of regeneration of the breakage injuries was approximately two-times greater for M. tuberculosa than for A. hyacinthus colonies. The regeneration rate of all injury types declined over time for both species. Recovery of injuries was influenced by the amount of algae that colonised the lesion and the zone of tissue available for regeneration. In general, the amount of regeneration of injuries was lower for injuries with a small zone of basal tissue available for regeneration, and the amount of regeneration was negatively correlated with the amount of algal settlement. The results of this study are important for understanding the role of partial mortality in the demography of scleractinian corals. Injuries impact on the population dynamics of colonies because regenerative processes require resources usually available for other processes (e.g. growth and reproduction), and slow recovery rates potentially increase the chance of whole colony mortality.
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This article aims to review 1) the major and most frequent human-induced physical disturbances and their consequences on coral reef habitats using a multi-scale approach, and 2) the scale-related indicators and conceptual aspects used to detect and measure the effects of these physical impacts. By physical disturbances, we mean direct perturbations that lead to the destruction/erosion of the carbonate framework. Human-induced direct physical disturbances are numerous from coastal development, tourism, harvesting, accidents and nuclear/weapon testing. Since methods for monitoring and measuring indicators are generally scale-implicit, coral reefs are first presented according to different ecological-spatial scales of organization, from colony to region (colony, reefscape, reef zone. whole reef, island and region). In this way, it is easier to link a couple (habitat, disturbance) to their potential indicators and to the descriptors they target. Three classes of descriptors, related to the response of the living, component of coral reef ecosystem, are considered here: stony coral, reef fishes and the human uses. A synthesis of the different options for coral habitat assessments is proposed. We sort them according to their objectives (monitor, initial status or improvement of knowledge), their specificities (identification or not of a specific disturbances) and their scale of investigation (small, meso- or large scales). Usually, the majority of the indicators of human-induced disturbances are non-specific. They reveal that something is happening but not the actual causality and can only detect differences across time or space. A major weakness lies in the difficulty in deconvoluting the signals from a conjunction of stressors occurring at different scales. As such, a hierarchical concept of disturbances in coral reefs would be the next logical step to enhance our capabilities in monitoring and forecasting coral reefs status.
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The reproductive biology, development, and planula behavior of the gorgonian Pseudopterogorgia elisabethae were studied at 2 sites in the Bahamas between 1996 and 2001. Colonies were gonochoric, and females brooded planulae on the colony surface. Gonads were observed only in colonies 18 cm high or larger. Spawning was asynchronous within and between sites but was concentrated 2–10 days after the new moons from late November through early January. Fertilized eggs developed into planulae over 1–2 days and the planulae remained attached to the surface of the female colony for an additional 2–4 days. Planulae were negatively buoyant and field observations suggest that larvae may settle within tens of meters of the maternal colony. P. elisabethae is harvested for natural products, and information on the reproduction of this commercially important species is crucial to the understanding of its population biology and to the development of management plans for the conservation of the species.
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Six coral species of the genus Acropora and two species of the genus Porites were studied during experiments on cultivation of reef-building scleractinian corals. The research has established species-specific factors and others affecting regeneration of fragments and growth of new colonies in these coral species. The accretion of donor fragments and new branches averaged from 40 to 160 mm per year, depending on the coral species, colony size, and season of transplantation. An average monthly accretion of medium and larger transplants and growth of new branches were 1.2–1.3 times higher at spring cultivation than at autumn transplanting. When transplanted, coral fragments of medium and larger sizes survived well and showed higher growth rates in all species studied. These transplants developed the highest number of new branches, and their buds and formed the largest colonies. Prolongation of the cultivation time from 1 to 1.5 years caused a 1.2–1.4 fold accretion of transplants.
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