Growth and population dynamic model of the reef coral Fungia granulosa Klunzinger, 1879 at Eilat, northern Red Sea

The lack of population dynamic information for most species of stony corals is due in part to their complicated life histories that may include fission, fusion and partial mortality of colonies, leading to an uncoupling of coral age and size. However, some reef-building corals may produce compact upright or free-living individuals in which the above processes rarely occur, or are clearly detectable. In some of these corals, individual age may be determined from size, and standard growth and population dynamic models may be applied to gain an accurate picture of their life history. We measured long-term growth rates (up to 2.5 years) of individuals of the free-living mushroom coral Fungia granulosa Klunzinger, 1879 at Eilat, northern Red Sea, and determined the size structure of a population on the shallow reef slope. We then applied growth and population models to the data to obtain estimates of coral age, mortality rate, and life expectancy in members of this species. In the field, few F. granulosa polyps suffered partial mortality of >10% of their tissues. Thus, the majority of polyps grew isometrically and determinately, virtually ceasing growth by about 30-40 years of age. Coral ages as revealed by skeletal growth rings were similar to those estimated from a growth curve based on field data. The frequency of individuals in each age class on the reef slope decreased exponentially with coral age, indicating high mortality rates when corals were young. The maximum coral age observed in the field population (31 years) was similar to that estimated by application of a population dynamic model (30 years). Calculated rates of growth, mortality and life expectancy for F. granulosa were within the range of those known for other stony corals. Our results reveal a young, dynamic population of this species on Eilat reefs, with high turnover rates and short lifespans. Such information is important for understanding recovery of coral reefs from disturbances, and for application to the management of commercially exploited coral populations.     

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     

Sexual reproduction in the tropical corallimorpharian Rhodactis rhodostoma

Abstract. Polyps of the tropical corallimorpharian Rhodactis rhodostoma segregate sexes between center and edge positions within aggregations produced by clonal replication. On a reef flat at Eilat, northern Red Sea, infertile polyps and males occur mainly along the edges of clonal aggregations, while females mostly occupy central positions within each aggregation. In addition, on the inner to middle reef flat where polyps of this species are abundant, aggregations consist mostly of females. On the outer reef flat, where polyps are rare, a sampled aggregation consisted mostly of males and infertile polyps. Male polyps are significantly smaller than females, and the smallest polyps are infertile. Fecundity increases significantly with polyp size in females, but testis size and number do not vary with body size in males. Oocytes are present in polyps during most of the year and gradually increase in size until annual spawning in June-July during the period of maximum day length. Testes do not vary significantly in size during the year and remain a small proportion of body mass (>8%). In contrast, females invest up to 30% of their body mass into gonads during the months immediately before spawning. The annual spawning of gametes coincides with a temporary drop in the frequency of clonal replication by polyps. We estimate that each female polyp of R. rhodostoma may release up to 3000 large eggs (500 μm in maximum diameter) each summer. The high investment of this corallimorpharian in sexual production of planktonic propagules may allow rapid dispersal to reef habitats distant from parent populations.     

Effects of anemonefish on giant sea anemones: expansion behavior,growth, and survival

The symbiosis between giant sea anemones and anemonefish on coral reefs is well known, but little information exists on impacts of this interaction on the sea anemone host. On a coral reef at Eilat, northern Red Sea, individuals of the sea anemone Entacmaea quadricolor that possessed endemic anemonefish Amphiprion bicinctus expanded their tentacles significantly more frequently than did those lacking anemonefish. When anemonefish were experimentally removed, sea anemone hosts contracted partially. Within 1–4 h in most cases, individuals of the butterflyfish Chaetodon fasciatus arrived and attacked the sea anemones, causing them to contract completely into reef holes. Upon the experimental return of anemonefish, the anemone hosts re-expanded. The long-term growth rate and survival of the sea anemones depended on the size and number of their anemonefish. Over several years, sea anemones possessing small or no fish exhibited negative growth (shrinkage) and eventually disappeared, while those with at least one large fish survived and grew. We conclude that host sea anemones sense the presence of symbiotic anemonefish via chemical and/or mechanical cues, and react by altering their expansion behavior. Host sea anemones that lack anemonefish large enough to defend them against predation may remain contracted in reef holes, unable to feed or expose their tentacles for photosynthesis, resulting in their shrinkage and eventual death.     

Effects of allogeneic contact on life-history traits of the colonial ascidian Botryllus schlosseri in Monterey Bay

The formation of chimeric colonies following allogeneic contact between benthic invertebrates may strongly affect colony fitness. Here we show that, in a field population of the colonial ascidian Botryllus schlosseri in Monterey Bay, California, more than 20% of all colonies occur in allogeneic contact with conspecifics. We experimentally assessed the effects of allogeneic contact on the following life-history traits under natural field conditions: growth, age and size at first reproduction, and egg production (fecundity). When compared with isolated colonies, and in some cohorts also with colonies that rejected allogeneic neighbors, colonies that fused with neighbors incurred reduced fitness in terms of most life-history traits measured. We propose that one of the benefits of precise allorecognition is that, in fused colonies, it limits the unit of selection to chimeric individuals composed of closely related kin.     

A complex allorecognition system in a reef-building coral: delayed responses,reversals and nontransitive hierarchies

A highly polymorphic and complex allorecognition system in the coral Stylophora pistillata was revealed in the field by assaying branch pair combinations among 11 colonies (181 assays) for 24 months. Replicates of between-colony combinations exhibited consistent outcomes, in both time scale and type of response. Different allogeneic combinations exhibited one of two main outcomes, either unilateral rejection, or an array of other incompatible reactions following a state of non-fusion. These responses were partially linked with color morphs (purple dominated yellow). An additional 22 isogeneic grafts resulted in complete fusion. Unilateral rejection occurred 1–7 months following initial contact. Nonfusion usually developed into skeletal suture barriers after 3–9 months, and then into unilateral colony-specific overgrowths at 6–23 months with some reversals in direction at 18–22 months. During this process, small lesions usually developed on the tissue of the subordinate partner, which were either overgrown by the dominant partner or healed. After two years, a network of overgrowths among colonies was established with essentially hierarchial properties, but some nontransitive interactions also occurred. The colonies segregated into three distinct histocompatibility groups; within each group, colonies engaged in nonfusion. Between groups, colonies exhibited nonfusion or rejected each other in a group-specific manner. Based on the results, we discuss the terminology used for fusion versus rejection phenomena in scleractinian corals, the possible genetic background for self-nonself recognition in Stylophora, and the methodological artifacts associated with the use of short-term allorecognition assays. Correspondence to: B. Rinkevich     

Abundance and clonal replication in the tropical corallimorpharian Rhodactis rhodostoma

Abstract. The corallimorpharian Rhodactis rhodostoma appears to be an opportunistic species capable of rapidly monopolizing patches of unoccupied shallow substrate on tropical reefs. On a fringing coral reef at Eilat, Israel, northern Red Sea, we examined patterns of abundance and clonal replication in R. rhodostoma in order to understand the modes and rates of spread of polyps across the reef flat. Polyps were abundant on the inner reef flat (maximum 1510 polyps m⁻² and 69% cover), rare on the outer reef flat, and completely absent on the outer reef slope at >3 m depth. Individuals cloned throughout the year via 3 distinct modes: longitudinal fission, inverse budding, and marginal budding. Marginal budding is a replicative mode not previously described. Cloning mode varied significantly with polyp size. Approximately 9% of polyps cloned each month, leading to a clonal doubling time of about 1 year. The rate of cloning varied seasonally and depended on day length and seawater temperature, except for a brief reduction in cloning during midsummer when polyps spawned gametes. Polyps of R. rhodostoma appear to have replicated extensively following a catastrophic low-tide disturbance in 1970, and have become an alternate dominant to stony corals on parts of the reef flat.     

Factors controlling the expansion behavior of favia favus (Cnidaria: Scleractinia): effects of light, flow, and planktonic prey

Colonies of the massive stony coral Faviafavus were exposed to different flow speeds and levels of light, and to the addition of zooplankton prey. The relative importance of each factor in controlling polyp expansion behavior was tested. The coral polyps fully expanded when they were exposed to low light intensity (0-40 micromol m(-2) s(-1)) and high flow speed (15 cm s(-1)), regardless of prey presence. They also partially expanded under low and medium light (40-80 micromol m(-2) s(-1)) at medium flow speed (10 cm s(-1)). The corals expanded their polyps only when they were exposed to light levels below compensation irradiance (Icom: light level at which photosynthesis = respiration), which was determined to be about 107 +/- 24 micromol m(-2) s. The results presented here indicate that high flow, low light, and the presence of planktonic prey induce coral expansion. There is a hierarchy of response to these stimuli, in which light level and flow speed are dominant over prey presence. Coral response to these three factors is probably due to the relative importance of gas exchange and zooplankton prey.