Delayed maturation in the colonial coralGoniastrea aspera (Scleractinia): whole-colony mortality, colony growth and polyp egg production

The present study examines (1) the cost of reproduction on colony growth, and (2) relationships among sexual maturity, whole-colony mortality rate and colony growth rate inGoniastrea aspera free from external influences by macrobenthos. Survival of colonies in permanent plots was followed for two years. Egg production by polyps in colonies collected just before the first spawning of a year was estimated by dissecting the polyps. Growth of the colonies (increase in number of polyps) was followed over one annual reproductive cycle. The cost of egg production on colony growth was apparent through colony ontogeny: (1) immature colonies had a greater annual growth rate than mature colonies, but produced almost no eggs; (2) in mature colonies, growth rate was negatively correlated with NE/PV (number of eggs per polyp volume mm^-3). Annual whole-colony mortality was high in colonies with fewer than11 polyps in initial colony size, while mortality was extremely low once a colony grew beyond this size. This critical size for low whole-colony mortality was much smaller than the colony size (40 polyps) which would attain maturity one year later. Age at maturity was estimated as six years. While survival to maturity may be a selective force for the evolution of delayed maturation, the present data suggest that high colony fecundity, achieved after a long growth period as an immature colony, and an abrupt decrease of colony growth rate after maturation are the crucial forces.     

Polyp oriented modelling of coral growth

The morphogenesis of colonial stony corals is the result of the collective behaviour of many coral polyps depositing coral skeleton on top of the old skeleton on which they live. Yet, models of coral growth often consider the polyps as a single continuous surface. In the present work, the polyps are modelled individually. Each polyp takes up resources, deposits skeleton, buds off new polyps and dies. In this polyp oriented model, spontaneous branching occurs. We argue that branching is caused by a so called “polyp fanning effect” by which polyps on a convex surface have a competitive advantage relative to polyps on a flat or concave surface. The fanning effect generates a more potent branching mechanism than the Laplacian growth mechanism that we have studied previously (J. Theor. Biol. 224 (2003) 153). We discuss the application of the polyp oriented model to the study of environmentally driven morphological plasticity in stony corals. In a few examples we show how the properties of the individual polyps influence the whole colony morphology. In our model, the spacing of polyps influences the thickness of coral branches and the overall compactness of the colony. Density variations in the coral skeleton may also be important for the whole colony morphology, which we address by studying two variants of the model. Finally, we discuss the importance of small scale resource translocation in the coral colony and its effects on the morphology of the colony.     

Intraspecific competition in a reef coral: effects on growth and reproduction

Summary Growth rates and reproduction of a branching coral (Stylophora pistillata) were compared in the presence and in the absence of intraspecific competition. Field experiments demonstrated a significant decline in the growth rate of competing colonies compared to noncompeting control colonies; the growth rate slowed in all of the interacting individuals, irrespective of their place in the hierarchy of the intraspecific dominance or of their color morph. In case of immediate killing of the subordinate, the dominant colony grew at a normal rate. In addition to the marked decrease in the growth rate of interacting colonies, the typical symmetry shape of these colonies was changed to an abnormal growth form. The number of female gonads per polyp was significantly reduced in colonies competing intraspecifically, and the typical synchrony in reproduction among different branches of a given colony was changed and desynchronized. Again, these results did not correlate with the hierarchy of dominance. We conclude that intraspecific competition in reef corals involves great investment of energy. The ecological significance and the different pathways of this competition are discussed.     

The effect of gravity on coral morphology

Coral morphological variability reflects either genetic differences or environmentally induced phenotypic plasticity. We present two coral species that sense gravity and accordingly alter their morphology, as characterized by their slenderness (height to diameter) ratio (SR). We experimentally altered the direction (and intensity) of the gravitational resultant force acting along or perpendicular to the main body axis of coral polyps. We also manipulated light direction, in order to uncouple gravity and light effects on coral development. In the experiments, vertically growing polyps had significantly higher SR than their horizontal siblings even when grown in a centrifuge (experiencing different resultant gravitational forces in proximal and distal positions). Lowest SR was in horizontal side-illuminated polyps, and highest in vertical top-illuminated polyps. Adult colonies in situ showed the same pattern. Gravitational intensity also affected polyp growth form. However, polyp volume, dry skeleton weight and density in the various centrifuge positions, and in aquaria experiments, did not differ significantly. This reflects the coral's ability to sense altered gravity direction and intensity, and to react by changing the development pattern of their body morphology, but not the amount of skeleton deposited.     

Within-colony feeding selectivity by a corallivorous reef fish: foraging to maximize reward?

Foraging theory predicts that individuals should choose a prey that maximizes energy rewards relative to the energy expended to access, capture, and consume the prey. However, the relative roles of differences in the nutritive value of foods and costs associated with differences in prey accessibility are not always clear. Coral‐feeding fishes are known to be highly selective feeders on particular coral genera or species and even different parts of individual coral colonies. The absence of strong correlations between the nutritional value of corals and prey preferences suggests other factors such as polyp accessibility may be important. Here, we investigated within‐colony feeding selectivity by the corallivorous filefish, Oxymonacanthus longirostris, and if prey accessibility determines foraging patterns. After confirming that this fish primarily feeds on coral polyps, we examined whether fish show a preference for different parts of a common branching coral, Acropora nobilis, both in the field and in the laboratory experiments with simulated corals. We then experimentally tested whether nonuniform patterns of feeding on preferred coral species reflect structural differences between polyps. We found that O. longirostris exhibits nonuniform patterns of foraging in the field, selectively feeding midway along branches. On simulated corals, fish replicated this pattern when food accessibility was equal along the branch. However, when food access varied, fish consistently modified their foraging behavior, preferring to feed where food was most accessible. When foraging patterns were compared with coral morphology, fish preferred larger polyps and less skeletal protection. Our results highlight that patterns of interspecific and intraspecific selectivity can reflect coral morphology, with fish preferring corals or parts of coral colonies with structural characteristics that increase prey accessibility.     

Fecundity and sexual maturity of the coral Siderastrea siderea at high latitude along the Florida Reef Tract,USA

Siderastrea siderea is one of the most abundant corals at high latitude shallow sites along the Florida Reef Tract (25°–27°N). This species is able to tolerate wide seawater temperature fluctuations and sedimentation stress, but its reproductive status at high latitudes and under marginal environmental conditions is poorly understood. The objectives of this study were to evaluate the reproductive potential of S. siderea along a latitudinal gradient (25°–27°N) and to determine if sexual maturity occurs in small (<12.0 cm) S. siderea colonies. Samples of coral tissue were collected in 2007, 2008, and 2009 at three sites along the latitudinal gradient and were processed for histological analysis. Oocyte size, volume, and abundance were used to calculate fecundity. Results showed that fecundity decreased with increasing latitude and that oocyte volume was the major contributing factor to this variation. Mature oocytes were observed in S. siderea colonies at sizes as small as 1.1 cm in diameter. The ability of S. siderea to reach fertility at high latitude areas suggests this species is able to reproduce under marginal environmental conditions; however, reduction in oocyte size could increase local retention of larvae. The presence of mature oocytes in small colonies suggests that stress can reduce somatic growth and shift sexual maturity to smaller colony sizes.     

Large dwellers of the Silurian Halysites biostrome: rhizosessile life strategies of cystiphyllid rugose corals from the Llandovery of Gotland

An exceptionally well‐preserved, unusual biostrome composed of the framebuilding cateniform tabulate coral Halysites catenularius (Linnaeus, 1767) bears an assemblage of the relatively large solitary cystiphyllid rugosan Cystiphyllum visbyense Wedekind, 1927. The corallites of solitary cystiphyllids are embedded within the ranks of the halysitid colonies, which developed on a soft, muddy substrate and in relatively turbid water. The cystiphyllid larvae successively settled mostly on the ranks of halysitid colonies and on colonies of the tiny phaceloid rugose coral Nanophyllum ramosum Johannessen, 1995, whereas calice‐in‐calice recruitment was not successful for these cystiphyllid corals. Further growth of C. visbyense was supported by rhizoid structures, which were most frequently developed on the cardinal (convex) side of the corallite. The process of formation of the rhizoid structures is here discussed and explained in detail, showing that they were formed by the extension of the basal ectodermal tissue of the polyp. The cystiphyllids, which settled on the walls of living corallites of halysitid colonies, used sweeper tentacles to kill the smaller polyps of the colony to maintain the space around them and expand. Hence, they ultimately used the halysitid colonies only as a hard substrate to stabilize their position on the soft muddy sediment.     

Structure and signaling in polyps of a colonial hydroid

Abstract. After feeding, polyps of colonial hydroids contract regularly, dispersing food throughout the colony via the gastrovascular fluid. Such contractions may trigger signaling pathways that allow colonies to grow in an adaptive manner, i.e., to initiate development of more polyps in food‐rich areas and to suppress polyp development in food‐poor areas. In this context, we investigated the structure and potential signaling of the junction between polyps and stolons in colonies of the hydroid Podocoryna carnea. Using transmission electron microscopy, we found that the density of mitochondrion‐rich epitheliomuscular cells was low in polyp and stolon tissues except at or near the polyp‐stolon junction, where many of these mitochondrion‐rich cells occur in ectodermal tissue. In vivo fluorescence microscopy suggests that these mitochondria are a principal source of the metabolic signals of the colony. Both native fluorescence of NAD(P)H and fluorescence from peroxides (visualized with H₂DCFDA) co‐localize to this region of the polyp. Rhodamine 123 fluorescence suggests that both these metabolic signals emanate from mitochondria. To test whether such metabolic signals may be involved in colony pattern formation, inbred lines of P. carnea were used. Colonies of a runner‐like inbred line grow with widely spaced polyps and long stolonal connections, much like wild‐type colonies in a food‐poor environment. Colonies of a sheet‐like inbred line grow with closely spaced polyps and short stolonal connections, similar to wild‐type colonies in a food‐rich environment. Polyp‐stolon junctions in runner‐like and sheet‐like colonies were imaged for the fluorescence of H₂DCFDA. Densitometric analysis of this signal indicates that the mitochondria in epitheliomuscular cells of runner‐like polyps emit greater amounts of peroxides. Because peroxides and other reactive oxygen species are frequently intermediaries in metabolic signaling pathways, we suspect that such signaling may indeed occur at polyp‐stolon junctions, affecting colony pattern formation in these inbred lines and possibly in hydroid colonies in general.     

Is the remodeling of the polyp prepattern in a hydrozoan planula a function of larval age or size and is it of “adaptive” significance?

Eggs of medusae develop into lecithotrophic planulae that undergo metamorphosis at different ages to form polyps. As planulae age they decrease in size as their yolk stores are utilized. The planulae of most Phialidium medusae develop into polyps where there is a decrease in the size of the holdfast region and a relative increase in the size of the hydranth region as they age. These changes occur independently of the decrease in planula size. In planulae with a decrease in the size of the holdfast region and an increase in the size of the hydranth-forming region there was a 50% decline in polyps that successfully stayed attached to the substrate after metamorphosis. These aged planulae produced an initial hydranth with the same number of tentacles as polyps from full-sized young planulae while young half-sized planulae produced hydranths where the tentacle number was smaller. The first phase of polyp colony growth with a small initial hydranth was slower than growth of a colony with a larger initial hydranth. Predation during this period led to more death in colonies with a small initial hydranth. The decline in successful attachment in aged planulae was not offset by the higher rate of growth and a smaller window of time where predation leads to death, suggesting that this age-related developmental change in planulae was not adaptive.     

The effects of a coral disease on the reproductive output of Montastraea faveolata (Scleractinia: Faviidae)

The direct impacts of coral diseases on coral populations have been assessed by quantifying coral tissue loss and colony mortality, but the determination of the indirect effects of diseases, such as disruptions in life history functions (e.g. reproduction, growth and maintenance), are more difficult to ascertain and have been scant. This study involved a comparison of various measures of reproductive output from histological slides of healthy tissue samples of Montastraea faveolata and tissue samples from colonies with white plague (WP) infections in Dominica (West Indies). Although the variability in the reproductive data was high, WP had significant negative impacts on the percentage of reproductive polyps per cm2, the percentage of reproductive mesenteries within a polyp, oocyte quantity per polyp, mean oocyte volume (mm3), and fecundity (oocyte volume per cm2 of tissue). However, these effects were only observed in the tissue directly impacted by the WP disease "band" and were not observed in tissue samples taken 20 cm away from the lesion. Therefore, the effects of a coral disease (WP) on reproductive output are localized and not expressed colony-wide.     

Sexual Reproduction and Seasonality of the Alaskan Red Tree Coral,Primnoa pacifica

The red tree coral Primnoa pacifica is an important habitat forming octocoral in North Pacific waters. Given the prominence of this species in shelf and upper slope areas of the Gulf of Alaska where fishing disturbance can be high, it may be able to sustain healthy populations through adaptive reproductive processes. This study was designed to test this hypothesis, examining reproductive mode, seasonality and fecundity in both undamaged and simulated damaged colonies over the course of 16 months using a deepwater-emerged population in Tracy Arm Fjord. Females within the population developed asynchronously, though males showed trends of synchronicity, with production of immature spermatocysts heightened in December/January and maturation of gametes in the fall months. Periodicity of individuals varied from a single year reproductive event to some individuals taking more than the 16 months sampled to produce viable gametes. Multiple stages of gametes occurred in polyps of the same colony during most sampling periods. Mean oocyte size ranged from 50 to 200 µm in any season, and maximum oocyte size (802 µm) suggests a lecithotrophic larva. No brooding larvae were found during this study, though unfertilized oocytes were found adhered to the outside of polyps, where they are presumably fertilized. This species demonstrated size-dependent reproduction, with gametes first forming in colonies over 42-cm length, and steady oocyte sizes being achieved after reaching 80-cm in length. The average fecundity was 86 (±12) total oocytes per polyp, and 17 (±12) potential per polyp fecundity. Sub-lethal injury by removing 21–40% of colony tissue had no significant reproductive response in males or females over the course of this study, except for a corresponding loss in overall colony fecundity. The reproductive patterns and long gamete generation times observed in this study indicate that recruitment events are likely to be highly sporadic in this species increasing its vulnerability to anthropogenic disturbances.     

Induction of mixed-function oxygenase system and antioxidant enzymes in the coral Montastraea faveolata on acute exposure to benzo(a)pyrene

Components of the cytochrome P(450) monooxygenase system (MFO) and antioxidant enzymes were investigated in the coral Montastraea faveolata exposed to the organic contaminant benzo(a)pyrene (B(a)P). For bioassays the corals were exposed to increasing concentrations of B(a)P (0.01 and 0.1 ppm) for 24 and 72 h, with water renewal every 24 h. Enzymatic activity of catalase (CAT), superoxide dismutase (SOD) and glutathione S-transferase (GST) were measured in host (polyp) and hosted (zooxanthellae) cells. NADPH cytochrome c reductase activity and contents of cytochrome P(450) and P(420) were only measured in the polyp. Antioxidant enzymes CAT and SOD in polyps and zooxanthellae and GST in polyps increased significantly at the highest concentration and maximum time of exposure. Cytochrome P(420) was found in all colonies, and the cytochrome P(450) content was greatest in the colonies from the highest concentrations of contaminant. NADPH cytochrome c reductase activity and the concentration of pigments did not vary between treatments. This is the first report of the induction of both detoxifying mechanisms, the MFO system and antioxidant enzymes on acute exposure to an organic contaminant in the reef-constructing coral species M. faveolata.     

Experimental fragmentation reduces sexual reproductive output by the reef-building coral Pocillopora damicornis

Natural and anthropogenic disturbances may fragment stony reef corals, but few quantitative data exist on the impacts of skeletal fragmentation on sexual reproduction in corals. We experimentally fragmented colonies of the branching coral Pocillopora damicornis and determined the number and size of planula larvae released during one lunar reproductive cycle. Partially fragmented colonies significantly delayed both the onset and peak period of planula release compared with intact control colonies. Most fragments removed from the corals died within 11–18 days, and released few planulae. The total number of planulae released per coral colony varied exponentially with remaining tissue volume, and was significantly lower in damaged versus undamaged colonies. However, the number of planulae produced per unit tissue volume, and planula size, did not vary with damage treatment. We conclude that even partial fragmentation of P. damicornis colonies (<25% of tissue removed) decreases their larval output by reducing reproductive tissue volume. Repeated breakage of corals, such as caused by intensive diving tourism or frequent storms, may lead to substantially reduced sexual reproduction. Therefore, reef management should limit human activities that fracture stony corals and lead to decreases in colony size and reproductive output. Accepted: 2 February 2000     

Effect of habitat characteristics on the distribution and abundance of damselfish within a Red Sea reef

For coral reef fish with an obligate relationship to their habitat, like Pomacentrid damselfish, choosing a suitable home amongst the reef structure is key to survival. A surprisingly small number of studies have examined patterns in adult damselfish distributions compared to other ontogenetic phases. The aim of this study was to determine which reef and coral colony characteristics explained adult damselfish distribution patterns in a Red Sea reef. The characteristics investigated were reef type (continuous or patchy), coral species (seven species of Acropora), and coral morphology (coral size and branching density). The focal damselfish species were Dascyllus aruanus, D. marginatus, Chromis viridis, and C. flavaxilla. Occupancy (presence or absence of resident damselfish), group size and fish species richness were not significantly different between the seven Acropora species. However, within each coral species, damselfish were more likely to occupy larger coral colonies than smaller coral colonies. Occupancy rates were also higher in patchy reef habitats than in continuous sections of the reef, probably because average coral colony size was greater in patchy reef type. Fish group size increased significantly with coral colony volume and with larger branch spacing. Multi-species groups of fish commonly occurred and were increasingly likely with reduced branching density and increased coral size.     

Analysis of spacing in a periodic pattern

Colonies of hydroids exhibit periodic biological patterns. Polyps form on stolons at fixed distances, obeying distinct rules. The spacing mechanism is based on inhibition emanating from existing polyps, predominantly from the head of the polyp. Removal of polyps from young colonies reduces the distance between initiating polyps and newly formed polyps to 50% of the normal values. Head removal suffices to obtain an almost identical reduction. Polyp enlargement which increases the distance between the inhibition-emitting head and the stolon tissue reduces the intrastolonal range of inhibition. In the stolon tissue, decrease of inhibitory activity occurs. An increase in the stolon/polyp ratio of a colony reduces bud distances. The decay is, in part, due to loss of inhibitory activity into the external medium: if colonies are incubated in conditioned culture medium derived from crowded colonies having normally large interpolyp distances, bud distances increase in test colonies. The effectiveness of transfer of inhibitory activity from tissue into the medium depends on culture conditions. If convection is increased by agitation of the culture medium, the distances between polyp and bud decreases; viscosity enhancement of the culture medium reduces convection and bud distances become larger. This effect is compensated for by additional agitation of the viscosity enhanced culture medium. Our results support the idea that a lateral inhibition mechanism controls polyp spacing in the stolon and that inhibition is based on diffusible inhibitory compounds.     

Scleractinian coral diseases in south Florida: incidence, species susceptibility, and mortality

There are limited quantitative data available documenting the natural, or non-epizootic, occurrence of scleractinian coral diseases over multiple years. Individual coral colonies exhibiting black band disease (BBD), white plague (WP), dark spots syndrome (DSS), and white band disease (WBD) were monitored 3 times per year on 5 south Florida reefs over a 2 yr period. Surveys included measurements of coral population composition, coral diversity, disease type, coral species affected, colony size, percent of colony affected, and the number of lesions or active infections per colony. Data on re-infections of the same colonies, multiple infections per colony, disease duration, disease-associated tissue mortality, and coral recruitment are also presented. A total of 674 coral colonies exhibiting coral diseases were tagged and monitored. DSS was the most common syndrome (n = 620 infected colonies), but BBD and WP infections caused the largest amount of coral tissue death. The only disease that exhibited a linear increase in incidence with elevated temperature was BBD. DSS and BBD were the most persistent conditions, and WP infections were comparatively short-lived, with obvious signs of disease typically disappearing after 2 to 3 mo. The only disease that caused total colony death as oppposed to partial mortality during the survey period was WBD. WP and DSS incidence was significantly positively correlated with the relative frequency of the species most commonly affected by each disease at each study site. Of the 61 colonies examined in the recruitment survey, only 5 scleractinian coral recruits were identified. The most commonly recorded colonizer of exposed coral skeleton was filamentous/turf algae, thus indicating the potential for a shift towards algal-dominated reef communities.     

Mechanisms of interaction between macroalgae and scleractinians on a coral reef in Jamaica

After several decades of disturbance, many coral reefs in the Caribbean are dominated by macroalgae. One process affecting this transition is coral-macroalgal competition, yet few studies have addressed the mechanisms involved. In this study, we investigated competition between the tall and bushy macroalga Sargassum hystrix (J. Agardh) and the branching coral Porites porites (Pallas) on a shallow reef in Jamaica. Experiments were designed to expose coral branches to different treatments to test the role of shading and abrasion by Sargassum on coral growth and polyp expansion. Corals exposed to Sargassum grew significantly more slowly (80% reduction) than controls, but this effect was absent when corals were caged to prevent physical contact with macroalgae. Light levels were reduced in both the algal and cage treatments, but shading apparently had little effect on the growth of corals in cages. Short-term measurements of integrated net water flow did not detect variation among treatments. In algal-mimic treatments, where clear plastic strips could touch but not shade the corals, growth rates were 25% lower than controls, but this effect was not statistically significant. Thus, the growth of corals in contact with Sargassum was reduced by abrasion and, to a lesser extent, by factors unique to living macroalgae. Analysis of polyp expansion showed that polyps were more frequently retracted when corals were in contact with macroalgae or algal-mimics compared to controls or cage treatment; the frequency of polyp contraction was correlated positively with growth. Together, these results suggest that abrasion-mediated polyp retraction is one of the primary mechanisms of competition utilized by tall (ca. 17 cm) macroalgae against scleractinian corals.     

Asexual reproduction in scyphistomae of Aurelia sp.: Effects of temperature and salinity in an experimental study

The growth and survival of colonies and individuals within sedentary polyp colonies of moon jellyfish (Aurelia sp.) was investigated at three temperatures and three salinities in laboratory experiments. Growth rates of colonies (number of polyps and number of buds in the colony) and individuals (number of buds per active scyphistomae) significantly increased with temperature, but were not affected by salinity. Survival was high in all treatment combinations indicating a wide tolerance to environmental conditions. However, scyphistomae at the lowest temperature had a greater percentage of larger individuals and slower population growth rate than those at warmer temperatures. These results suggest that the reproductive strategy to maximise production of Aurelia sp. is to increase the size of scyphistomae colonies by asexual budding when conditions are good (warmer temperatures and abundant food generally during spring and summer). Budding activity slows, but the size of scyphistomae increases, during the colder winter period leading up to strobilation, resulting in the production of a greater number of ephyrae. The trigger for strobilation is possibly stressful conditions. However, if trigger conditions do not occur, the colony of scyphistomae can continue to grow and survive through a broad range of conditions spanning many seasons, thus ensuring survival of the population.     

Food selection of a corallivorous butterflyfish Chaetodon austriacus in the Red Sea, with particular emphasis on size and fish symbionts of corals

The exquisite butterflyfish Chaetodon austriacus feeds mainly on Acropora , Pocillopora , Montipora and Stylophora in the northern Red Sea. Large colonies of Acropora are preferred over smaller colonies and other coral genera, whereas avoidance is indicated for corals that are occupied by certain coral-associated gobiid fishes of the genus Gobiodon . It is suggested that, apart from coral identity, colony size and potential repellent effects of particular coral symbionts are both important sources of variation in the food selectivity of corallivorous chaetodotids.     

Tumor formation on the coral Montipora informis

A high incidence of tumors (i.e., abnormal skeletal growth) was observed on Montipora informis Bernard, 1897 (Acroporidae) coral colonies on the shallow reef flat of Sesoko Island, Okinawa, Japan. Tumors were recognized as slightly hemispherical protuberances and were characterized by fewer numbers of polyps per surface area, fewer zooxanthellae per polyp, and finer skeletal structures than normal. In addition, fecundity was significantly reduced in tumored areas. Tumor formation appeared localized, stable and non-epidemic. During the summer of 1998, coincident with the global coral bleaching event, tumored tissue suffered higher mortality than normal tissue. The mechanism triggering tumor formation remains unknown.