Testing the first phase of the ‘gardening concept’ as an applicable tool in restoring denuded reefs in Tanzania

Studies on coral reef restoration through a two-step coral gardening protocol have lately proved it to be a viable solution for future reef restoration. This involves a first step of gardening small colonies in mid-water nurseries and a second step, their transplantation, upon reaching suitable size, onto the pre-surveyed damaged areas. We established in September 2007 two mid-water nurseries, each holding 10,000 fragments measuring 2 cm average initial size, at 4 m depths (high tide) in Zanzibar and Mafia Islands, Tanzania. Each nursery comprised six species, each of which was represented by three genotypes. During 9 months, we followed developments by analyzing and comparing survivorship and growth rates of fragments between the different nurseries, species and genotypes. A significant difference between species survival and growth rates was observed in acroporid species, in Pocillopora verrucosa and Millepora sp., which showed better success than Porites cylindrica. In both sites, Millepora suffered no mortality and other species exhibited low mortality, ranging (per coral genotype) between 3% and 24% in Zanzibar (most cases below 10%) and between 13% and 44% (mostly below 25%) in Mafia Island. Most of fragments’ mortality occurred during the first two nursery months. Coral species in Zanzibar nursery also performed better in growth rates than those in Mafia, but in both sites, farmed corals were ready for transplantation just 9 months after the nursery was set up. Economic evaluations involved in the overall nursery set-up and the results indicated that the coral gardening approach could be used in Tanzania to generate large quantities of coral colonies for the restoration of damaged reefs at relatively low cost.     

Genotype and attachment technique influence the growth and survival of line nursery corals

The Caribbean staghorn coral, Acropora cervicornis, was once a dominant habitat creating coral, but its populations have declined dramatically in recent decades. Numerous restoration efforts now utilize coral gardening techniques to cultivate this species, growing colonies on fixed structures or from line/suspended nurseries. Line nurseries have become increasingly popular because of their small footprint and ease of use, replacing fixed structures in many nurseries. To evaluate the efficacy of the line technique, this study evaluated growth, condition, and survivorship of A. cervicornis nursery colonies of three distinct genotypes grown via two line nursery techniques (suspended and direct line attachment [vertical]). Direct line attachment of nursery colonies resulted in poor survival (43%) and growth (9.5 ± 1.33 cm/year), whereas suspended culture had 100% survival and increased growth (61.1 ± 4.19 cm/year). Suspended culture had significantly reduced disease prevalence and prevented colony predation. Suspended coral growth was also comparable to a neighboring fixed structure nursery (55.2 ± 7.86 cm/year), and found to be as effective in propagating corals as fixed structures.     

Growth Dynamics of the Threatened Caribbean Staghorn Coral Acropora cervicornis: Influence of Host Genotype,Symbiont Identity,Colony Size,and Environmental Setting

Background

The drastic decline in the abundance of Caribbean acroporid corals (Acropora cervicornis, A. palmata) has prompted the listing of this genus as threatened as well as the development of a regional propagation and restoration program. Using in situ underwater nurseries, we documented the influence of coral genotype and symbiont identity, colony size, and propagation method on the growth and branching patterns of staghorn corals in Florida and the Dominican Republic.

Methodology/Principal Findings

Individual tracking of> 1700 nursery-grown staghorn fragments and colonies from 37 distinct genotypes (identified using microsatellites) in Florida and the Dominican Republic revealed a significant positive relationship between size and growth, but a decreasing rate of productivity with increasing size. Pruning vigor (enhanced growth after fragmentation) was documented even in colonies that lost 95% of their coral tissue/skeleton, indicating that high productivity can be maintained within nurseries by sequentially fragmenting corals. A significant effect of coral genotype was documented for corals grown in a common-garden setting, with fast-growing genotypes growing up to an order of magnitude faster than slow-growing genotypes. Algal-symbiont identity established using qPCR techniques showed that clade A (likely Symbiodinium A3) was the dominant symbiont type for all coral genotypes, except for one coral genotype in the DR and two in Florida that were dominated by clade C, with A- and C-dominated genotypes having similar growth rates.

Conclusion/Significance

The threatened Caribbean staghorn coral is capable of extremely fast growth, with annual productivity rates exceeding 5 cm of new coral produced for every cm of existing coral. This species benefits from high fragment survivorship coupled by the pruning vigor experienced by the parent colonies after fragmentation. These life-history characteristics make A. cervicornis a successful candidate nursery species and provide optimism for the potential role that active propagation can play in the recovery of this keystone species.     

Mid-water rope nursery—Testing design and performance of a novel reef restoration instrument

Fast degradation of coral reefs worldwide has promoted the exploitation of active restoration instruments, one of which is the ‘gardening concept’. This concept comprises two phases: (1) establishing in situ coral nurseries for rearing large numbers of coral fragments; (2) their transplantation onto denuded reefs. This study tested the design and performance of a novel mid-water floating nursery instrument, a ‘rope nursery’. This nursery accommodated small coral fragments attached to a rope, creating an easily constructed nursery bed that is rapid and inexpensive. Two sets of experiments were conducted: the first tested two mid-water rope nursery prototypes in small-scale trials that tested depth, coral genotypes and construction stability, whereas the second set incorporated lessons learned from the first set, and was designed to carry larger numbers of colonies. These highly economical nurseries (US$ 0.11/fragment) revealed high survivorship low detachment and fast growth rates compared to previous coral-nursery types. Moreover, the coiling force of the ropes adequately held fragments without adhesives, and the minimal surface area of rope nursery beds provided not only improved water flux around farmed corals, but also reduced proliferation of fouling organisms. The rope nursery prototypes studied here attest to the diversity of their potential uses under various conditions and demands, making the construction of large scale nurseries a very feasible target. This restoration instrument was proven to be an effective coral reef rehabilitation tool.     

Coral reef restoration in the Eastern Tropical Pacific: feasibility of the coral nursery approach

Due to the worldwide degradation of coral reefs, the active restoration of these ecosystems has received considerable attention in recent decades. This study investigated (1) the feasibility of using coral nurseries for restoration projects, (2) the minimum size required for a Pocillopora damicornis (Pocilloporidae) coral fragment to survive and grow in a nursery, and (3) the optimal transplant size of a fragment when transplanted to a degraded reef at Gorgona Island (Colombian Pacific). For this investigation, 230 fragments were transplanted directly to El Remanso reef, and another 150 fragments were maintained in in situ nurseries. Every 2 months, the length, weight, and survival of the fragments were recorded. After growing for 134 days in the nurseries, the 52 surviving fragments were transplanted to El Remanso reef, and after 5 months, the same variables were measured. Among the nursery‐reared fragments, the largest (4 to <8 cm) had the highest survival and growth rates, whereas among the directly transplanted fragments, the smallest fragments (<2 cm) had the highest survival and growth rates. However, the nursery‐reared fragments acquired greater structural complexity (arborescent morphology), and they were all alive 156 days after transplantation and presented a maximum linear growth rate of over 2 cm, which was higher than that of the directly transplanted fragments. Apparently, the arborescent morphology acquired during the nursery period provides advantages to the colonies that favor greater success when transplanted. Therefore, nursery‐reared fragments of P. damicornis between 2 and 4 cm are the most appropriate for use in restoration projects.     

The role of in situ coral nurseries in supporting mobile invertebrate epifauna

Coral nurseries are commonly employed to generate coral material for reef restoration projects, but observations of epifaunal organisms utilising the nurseries for food and shelter indicate that they can also provide important functions beyond that of coral propagation. To examine the level of biodiversity that can be supported by coral nurseries, and investigate if epifaunal communities were influenced by the presence of live coral tissue, we compared the abundance, diversity and community composition of mobile invertebrate epifauna associated with live and dead fragments of three coral species (Pocillopora acuta, Echinopora lamellosa, Platygyra sinensis) that were reared in an in situ nursery. A total of 418 mobile invertebrates spanning 63 taxa were recorded from 22 coral colonies. The three coral species hosted significantly different epifaunal communities, most likely a consequence of the difference in growth forms of the coral hosts. Significant differences in epifaunal communities were only observed between live and dead colonies of P. acuta, indicating that resource provisioning in this species is particularly influenced by the presence of live tissue. Our findings showed that coral nurseries can support a range of mobile invertebrates and function as tools to conserve threatened mobile invertebrates. This ecological function is under-studied and should be assessed in restoration programs for the conservation of corals and associated fauna.     

Detachment of Porites cylindrica nubbins by herbivorous fishes

A form of active restoration for coral assemblages involves culturing coral nubbins at nursery sites before transplantation to recipient reefs. Incidental grazing and/or directed predation by local fish assemblages are major sources of dislodgement and mortality for coral nubbins in nurseries. However, the rate of coral nubbin detachment, how this varies across fish taxa, and whether nubbin size affects rates of detachment warrant further investigation. We used field and aquaria experiments to examine the effect of incidental grazing and predation on the detachment of Porites cylindrica nubbins of different sizes (0.5, 1, 2, 3, 4, and 5 cm height). Short‐term (6 hours) exposure of nubbins to local fish assemblages at Lucero Reef, northwestern Philippines, caused higher detachment (1.93% ± 0.53 SE) compared to caged controls (0.16% ± 0.16 SE), with no detectable effect of nubbin size. To identify the impact of individual fish species, nubbins were exposed to one of four locally abundant herbivorous and corallivorous fish species in aquaria for 8 hours. Nubbin detachment was greater when exposed to Chlorurus spilurus (1.20–36.2%) and Siganus fuscescens (0.00–15.0%) than Chaetodon lunulatus (0.00–4.00%) and Chaetodon kleinii (0.00–1.20%), with the smallest nubbins (0.5 cm) being the most vulnerable. Our results suggest that incidental grazing by herbivorous fishes, especially parrotfishes, may potentially be an important source of detachment and likely mortality of nubbins. Optimizing coral nursery protocols should consider potential trade‐offs between excluding grazing fishes and the accumulation of algal material on caging structures to minimize nubbin mortality and improve coral restoration success.     

In Situ Coral Nurseries Serve as Genetic Repositories for Coral Reef Restoration after an Extreme Cold‐Water Event

During an unusual cold‐water event in January 2010, reefs along the Florida Reef Tract suffered extensive coral mortality, especially in shallow reef habitats in close proximity to shore and with connections to coastal bays. The threatened staghorn coral, Acropora cervicornis, is the focus of propagation and restoration activities in Florida and one of the species that exhibited high susceptibility to low temperatures. Complete mortality of wild staghorn colonies was documented at 42.9% of donor sites surveyed after the cold event. Remarkably, 72.7% of sites with complete A. cervicornis mortality had fragments surviving within in situ coral nurseries. Thus, coral nurseries served as repositories for genetic material that would have otherwise been completely lost from donor sites. The location of the coral nurseries at deeper habitats and distanced from shallow nearshore habitats that experienced extreme temperature conditions buffered the impacts of the cold‐water event and preserved essential local genotypes for future Acropora restoration activities.     

Propagation of the threatened staghorn coral Acropora cervicornis: methods to minimize the impacts of fragment collection and maximize production

Coral reef restoration methods such as coral gardening are becoming increasingly considered as viable options to mitigate reef degradation and enhance recovery of depleted coral populations. In this study, we describe several aspects of the coral gardening approach that demonstrate this methodology is an effective way of propagating the threatened Caribbean staghorn coral Acropora cervicornis: (1) the growth of colonies within the nursery exceeded the growth rates of wild staghorn colonies in the same region; (2) the collection of branch tips did not result in any further mortality to the donor colonies beyond the coral removed for transplantation; (3) decreases in linear extension of the donor branches were only temporary and donor branches grew faster than control branches after an initial recovery period of approximately 3–6 weeks; (4) fragmentation did not affect the growth rates of non-donor branches within the same colony; (5) small branch tips experienced initial mortality due to handling and transportation but surviving tips grew well over time; and (6) when the growth of the branch tips is added to the regrowth of the fragmented donor branches, the new coral produced was 1.4–1.8 times more than new growth in undisturbed colonies. Based on these results, the collection of small (2.5–3.5 cm) branch tips was an effective propagation method for this branching coral species resulting in increased biomass accumulation and limited damage to parental stocks.     

Improved sustainable maintenance for mid-water coral nursery by the application of an anti-fouling agent

The ‘gardening concept’ for reef restoration focuses on coral colonies farming in mid-water nurseries before their transplantation onto denuded reef areas. Nurseries situated in a nutrient-enriched environment significantly curtail nursery time, but extend labor, as nursery construction and farmed corals must be frequently cleaned from competing fouling organisms. Mass farming of corals calls, therefore, for efficient and cheap maintenance methodologies, which we here tested by employing Aqua-guard M250, an anti-fouling agent used in the fish farming industry. We found that this anti-fouling paint, while reducing fouling organisms on nursery components during the crucial phase of coral ramets' development from nubbins and small fragments sizes to colony sizes suitable for transplantation, is not toxic to maricultured coral fragments that staged more than 2 cm away from the paint. Applying small quantities of such antifouling paint to coral nurseries, while restricting its use to nursery components that are not in direct contact with farmed coral material, reduces fouling coverage and cleaning procedures by 90%.     

Macroalgae reduces survival of nursery‐reared Acropora corals in the Florida reef tract

Recent declines in coral populations along the Florida reef tract have prompted the establishment of coral restoration programs which raise coral species, such as the threatened Acropora cervicornis, in nurseries ready for outplanting. Large numbers of nursery‐reared coral colonies have been outplanted along the Florida reef tract in recent years, yet few studies have characterized benthic habitats that are considered optimal for colony survival. In 2016, we surveyed 23 A. cervicornis restoration sites, located at six different reefs in the upper Florida Keys. We examined the condition of the outplanted corals and quantified the benthic assemblages adjacent to the outplanted coral colonies. We found that where A. cervicornis survived for more than 1 year, the substrate significantly supported less brown macroalgae of the genus Dictyota than at sites where A. cervicornis had died. Coral survival was highest at sites with less than 15% Dictyota cover. These results suggest that the habitat conditions that supported Dictyota spp. were not conducive to A. cervicornis growth and survival. Restoration practitioners should avoid attaching nursery‐raised corals to substrate with Dictyota spp. cover greater than 15%.     

In situ nurseries enhance coral transplant growth in sedimented waters

In situ nurseries have been a crucial part of coral reef restoration initiatives for the past two decades. However, the advantages over direct transplantation in sedimented waters has yet to be examined. In the present study, we showed that Pachyseris speciosa and Pocillopora damicornis fragments reared in in situ nurseries (NR) in Singapore’s sedimented waters grew significantly faster (by three to five times) than those which were directly transplanted (DT) onto the substrates. The increased growth rate during the nursery phase augmented the size of NR transplants, and had a flow-on effect on their performance during the post-transplantation phase. Overall, the maximum diameter of the NR transplants was 1.8–2.7 times larger than DT transplants after 11 months. The growth enhancement of the nursery-reared transplants improved the cost-effectiveness of our restoration effort: the estimated cost per centimetre growth of NR transplants was one-fifth of the DT corals despite the additional costs incurred to construct the nurseries. These results highlight that coral nurseries are beneficial to reef restoration in chronically sedimented waters.     

Employing a highly fragmented,weedy coral species in reef restoration

A major determinant in reef restoration programs is the choice of species employed. In this paper, we concentrate on the potential use of Montipora digitata (Dana, 1846), a highly fragmented and weedy coral species, for reef restoration and for recreation of reef physiognomy in Bolinao, The Philippines, under the ‘gardening concept’. Coral ramets (n = 1960) were reared in a shallow lagoon nursery for 1 year and then, the resulting nursery-grown colonies were transplanted onto denuded reef knolls at two different sites and according to three transplantation designs (grid design with high or low density and patch design). The nursery grown ramets, which had initially exhibited a 99% survivorship, were reduced to 87% by a super typhoon. Low detachment rates, low mortality (<1% both) and low bleaching (<3% over most months), have been indicating good nursery conditions for corals. Monitoring transplanted colonies for over 15 months showed no significant differences between the sites or among the three different transplantation designs. The major events of mortality and detachment were documented during the first 3 months post-transplantation and in the wake of a bleaching event. M. digitata transplants exhibited rapid growth (a 384% increase in ecological volume) and frequent fragmentation. Some fragments remained by their colonies of origin entangled in the attached branches, while other fragments fell onto the surrounding sandy substrate, forming M. digitata ‘beds’ around the knolls. In both, nursery and transplanted corals, creation of complex 3D structures was followed by recruitment of juvenile fishes into aggregations. Above results reveal that employing fast growing, weedy and easily fragmenting branching species, which are also considered as ‘modifier species’, in reef restoration may result in a fast regeneration of reef physiognomy. Instead of ‘copying’ natural processes, reef managers should therefore concentrate on recovering the reef's ability for self-maintenance.     

Assessment of the nursery species pool for restoring landscapes in southeastern Brazil

Brazil has committed to fulfilling international restoration goals and to enforcing environmental legislation that will require private landowners to undertake ecological restoration of 21 million hectares of degraded and deforested landscapes. To support a broad range of restoration practices, a well‐established supply chain capable of representing regional plant diversity is essential. This study investigated the restoration species pool in native plant nurseries in São Paulo state, located in southeastern Brazil, and evaluated their geographic distribution, similarity of their plant stocks, and the proportion of species represented from regional floras. Despite a lack of technical assistance and a significant presence of nonnative species (126 species, average 7.5 species/nursery), we found an impressive native species richness in plant nurseries (561 species, average 86.4 species/nursery) from both the Atlantic Forest and Cerrado domains, representing 38–44% of regional floras. There was a huge bias toward tree and shrub species (96.6%) and an absence or underrepresentation of other growth forms, as well as of savanna specialists, animal‐dispersed, and threatened species. The dissimilarity of species observed among the different nurseries surveyed highlights their role in representing regional diversity, which reflects their regional seed collection practices. Effective assistance and training are essential to address issues related to misidentification of species, underrepresentation of most functional plant groups, and the presence of nonnative species, as well as to support the supply chain, which is currently undergoing a market downturn.     

Coral Reef Restoration (Bolinao,Philippines) in the Face of Frequent Natural Catastrophes

Restoration of coral reefs is generally studied under the most favorable of environmental conditions, a stipulation that does not always reflect situations in the field. A 2‐year study (2005–2007), employing the “reef gardening” restoration concept (that includes nursery and transplantation phases), was conducted in Bolinao, Philippines, in an area suffering from intense human stressors. This site also experienced severe weather conditions, including a forceful southwesterly monsoon season and three stochastic environmental events: (1) a category 4 typhoon hit the Bolinao's lagoon (May 2006) impacted farmed corals; (2) heavy rains (August 2006) caused seepages of freshwater, followed by reduced salinity that impacted transplanted colonies; and (3) a bleaching event (June 2007) caused by warming of seawater, severely impacted both nursery and transplanted corals. This study analyzes the effects of these natural catastrophes on restoration efforts, and presents the successes and failures of recently used restoration instruments. Our results show that (1) in the nursery phase, consideration should be paid to depth‐flexible constructions and tenable species/genotypes prioritization and (2) for transplantation acts, site/species deliberation, timing, and specific site selections should be taken into account. Only the establishment of large‐scale nurseries and large transplantation measures and the adapting of restoration management to the frequently changing environment may forestall extensive reef degradation due to the combination of continuous anthropogenic and worsening global changes.     

Simple ecological trade-offs give rise to emergent cross-ecosystem distributions of a coral reef fish

Ecosystems are intricately linked by the flow of organisms across their boundaries, and such connectivity can be essential to the structure and function of the linked ecosystems. For example, many coral reef fish populations are maintained by the movement of individuals from spatially segregated juvenile habitats (i.e., nurseries, such as mangroves and seagrass beds) to areas preferred by adults. It is presumed that nursery habitats provide for faster growth (higher food availability) and/or low predation risk for juveniles, but empirical data supporting this hypothesis is surprisingly lacking for coral reef fishes. Here, we investigate potential mechanisms (growth, predation risk, and reproductive investment) that give rise to the distribution patterns of a common Caribbean reef fish species, Haemulon flavolineatum (French grunt). Adults were primarily found on coral reefs, whereas juvenile fish only occurred in non-reef habitats. Contrary to our initial expectations, analysis of length-at-age revealed that growth rates were highest on coral reefs and not within nursery habitats. Survival rates in tethering trials were 0% for small juvenile fish transplanted to coral reefs and 24-47% in the nurseries. As fish grew, survival rates on coral reefs approached those in non-reef habitats (56 vs. 77-100%, respectively). As such, predation seems to be the primary factor driving across-ecosystem distributions of this fish, and thus the primary reason why mangrove and seagrass habitats function as nursery habitat. Identifying the mechanisms that lead to such distributions is critical to develop appropriate conservation initiatives, identify essential fish habitat, and predict impacts associated with environmental change.     

Ecological impacts of coral gardening outplanting in the Maldives

As coral reefs continue to degrade at an alarming rate, coral restoration efforts are increasing worldwide in an attempt to keep up with the global challenge of preserving these iconic ecosystems and the many services they provide. Coral gardening, the farming and outplanting of coral fragments, is a commonly applied practice; however, regional validation is required before upscaling can be considered. This study follows up from the successful farming of fragments in mid-water rope nurseries, by reporting on the successive outplanting of these corals. Specifically, 60 Pocillopora verrucosa colonies were outplanted to a degraded reef at different depths (1–12 m), applying three arrangement patterns (equal, clustered, random). After 1 year, 72% were considered successfully outplanted (alive and still attached), with detachment being the main challenge at wave-impacted shallow depths, while loose coral rubble caused more partial mortality at depth. Outplanting stress was observed at 1–6 m depth, but had no impact on survival or growth. Drupella sp. predation was most common at 3 m and 79% of colonies hosted mutualistic fauna after 1 year. Outplanting significantly benefitted the reef environment with a higher fish abundance and diversity along with a higher increase in natural coral cover (H = 2.7; 6.2% increase) in comparison with the control sites. These are promising results, considering that the restoration site has shown little natural recovery in the last few years (coral cover <4%). We hope that our findings provide useful initial insights and help to guide effective restoration practices in the Maldives.     

Feasibility of early outplanting of sexually propagated Acropora verweyi for coral reef restoration demonstrated in the Philippines

Over the last 20 years, coral sexual propagation techniques for reef restoration have been steadily developed and improved. However, these techniques involve considerable time and costs to grow coral propagules. There is a need to examine the optimal size of juvenile corals for outplantation. Here, we outplanted sexually propagated small (3–5 mm diameter) and large (10–15 mm diameter) Acropora verweyi corals at 4 months after fertilization at two sites in northwestern Philippines, and compared their survival and radial growth rate after a year. A. verweyi coral juveniles (n = 240) exhibited an overall mean survival of 29.5% and growth rate of 11.12 ± 6.2 mm/year (mean ± SD). Large colonies had a significantly higher growth rate than smaller colonies. Although survivorship of large juveniles was significantly better than that of the smaller ones at one site, it did not differ significantly at the other. Each 4‐month‐old coral cost US$1.52 to produce, while the cost of each of the outplanted juveniles (n = 240) was about US$2.67, whereas the cost of each survivor about a year after outplantation was US$11.47. Results suggest that A. verweyi reared in ex situ nurseries for only 4 months can survive reasonably well when outplanted onto coral reefs.     

Strategies for Gardening Denuded Coral Reef Areas: The Applicability of Using Different Types of Coral Material for Reef Restoration

Recreational and other human activities degrade coral reefs worldwide to a point where efficient restoration techniques are needed. Here we tested several strategies for gardening denuded reefs. The gardening concept consists of in situ or ex situ mariculture of coral recruits, followed by their transplantation into degraded reef sites. In situ nurseries were established in Eilat's (Northern Red Sea) shallow waters, sheltering three types of coral materials taken from the branching species Stylophora pistillata (small colonies, branch fragments, and spat) that were monitored for up to two years. Pruning more than 10% of donor colonies' branches increased mortality, and surviving colonies displayed reduced reproductive activity. Maricultured isolated branches, however, exceeded donor colony life span and reproductive activity and added 0.5–45% skeletal mass per year. Forty‐four percent of the small colonies survived after 1.5‐year mariculture, revealing average yearly growth of 75 ± 32%. Three months ex situ maintenance of coral spat (sexual recruits) prior to the in situ nursery phase increased survivorship. Within the next 1.5 years, they developed into colonies of 3–4 cm diameter. Nursery periods of 2 years, 4–5 years, and more than> 5 years have been estimated for small colonies, spat, and isolated branches, respectively. These and other results, including the possible use of nubbins (minute fragments the size of a single or few polyps), are discussed, revealing benefits and drawbacks for each material. In situ coral mariculture is an improved practice to the common but potentially harmful protocol of direct coral transplantation. It is suggested that reef gardening may be used as a key management tool in conservation and restoration of denuded reef areas. The gardening concept may be applicable for coral reefs worldwide through site‐specific considerations and the use of different local coral species.     

Ontogenetic changes in the capacity of the coral Pocillopora damicornis to originate branches

Most colonial corals vary intraspecifically in growth forms, and the diversity in branching morphology is especially striking. While the effects of environmental factors on growth forms have been studied, the genetic control of coral branching patterns has received little attention. The discovery of ontogenetic changes in the capacity to originate branching would set the stage for studies of how branch formation is genetically controlled. During experiments investigating contact reactions in the coral Pocillopora damicornis, we observed that young colonies derived from settled planulae and colonies regenerated from adult branch tips assumed different growth forms. Young colonies formed at least one branch from the central region of the colony, while colonies regenerated from adult branch tips (3-5 mm long) did not form branches during the 9-month observation period. This pattern was invariable, regardless of the types and outcomes of the contact experiments or the orientation of the branch tips. However, some fragments taken from 1- or 2-year-old colonies formed branches. This suggests that the rate of branch formation in P. damicornis colonies decreases with age. These findings will facilitate investigations of the mechanism of coral branch formation at the molecular level.