Ostroff Proposal SUBMITTED

8/4/2019 Ostroff Proposal SUBMITTED

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M.S. THESIS PROPOSAL DRAFT ZACH OSTROFFNOVA SOUTHEASTERN UNIVERSITY OCEANOGRAPHIC CENTER

EvaluatingAcropora cervicornis Growth and Survivorship in a Line Nursery

I. Introduction

Coral reefs are beautiful and diverse ecosystems of immeasurable importance and intrinsicvalue that provide a great wealth of ecological and economic goods and services (Jameson,1995; Moberg and Folk, 1999; Branderet al., 2007). Regardless of their importance, coralreefs are facing worldwide decline brought about by natural and anthropogenic impacts(Hoegh-Guldberg, 1999; Nystrm et al., 2000; Hughes and Connell, 1999; Hughes et al.,2003; Pandolfi et al., 2003). Near the turn of the century, it was estimated that as much as70% of the worlds coral reefs were directly threatened by human activities (Goreau, 1992;Sebens, 1994; Wilkinson, 1999), and that approximately one in three reef-building coralsfaced elevated extinction risk (Carpenter, et al., 2008). Prominent stresses include climatechange, disease proliferation, coastal eutrophication and sedimentation, ocean acidification,and destructive fishing practices (Sebens, 1994; Hughes and Connell, 1999; Hoegh-Guldberget al., 2007). Impacted by these stressors, coral reefs have been subject to varied, but in manycases dramatic degradation (Bruno and Selig, 2007; Mora, 2007). Caribbean reefs have faredcomparatively worse than those elsewhere, experiencing losses of over 80% of hard coralcover from 1977 to 2001 (Gardner et al., 2003; Figure 1); an average rate of declineapproximately four times greater than in the Indo-Pacific (Mumby and Steneck, 2008).

Figure 1. Observed coral percent cover decline in the greater Caribbean from 1977 to 2001 (Gardneret al.,2003). Triangles and closed circles represent weighted and un-weighted absolute mean coral cover respectively,open circles represent number of studies, and X represent data omissions.

Of the substantial losses of hard coral cover in the Caribbean, those of the Acropora genus havebeen some of the most severe (Aronson and Precht, 2001a; Bruckner, 2003), experiencing losses


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up to 98% in some instances (Milleret al., 2002). Historically, the Acroporidae were the mostprominent corals of Caribbean reef crests, with the elkhorn coralAcropora palmata dominatingthe top five meters of a reef, and the staghorn coral Acropora cervicornis prominently colonizingmuch of the fore reef at depths of about eight to fifteen meters (Goreau, 1959; Goreau and Wells,1967; Woodley and Robinson, 1977; Fig. 2). Regardless of their robust populations of the past,

both A. palmata and A. cervicornis are now listed as Threatened under the United StatesEndangered Species Act (Fed. Reg. 71).

Figure 2. Cross section diagram of DiscoveryBay, Jamaica exhibiting classicAcropora zonation.(adapted from Woodley and Robinson, 1977).

The precipitous population collapse of Caribbean Acropora species is a result of many stresses,and is speculated to also be due to their increased susceptibility to disease and bleachingcompared to other Caribbean coral species (Goreau et al., 1998; Hoegh-Guldberg, 1999;Aronson and Precht, 2001b; Williams and Miller, 2005). With the decline of these andnumerous other reef-building coral species, many Caribbean reefs are shifting from coral-dominated ecosystems to those dominated by other functional groups such as macroalgalcommunities (Done, 1992; Knowlton, 1992; Porter, 1992; Hughes, 1994; McClanahan, 2002;Pandolfi et al., 2005).

If such degradation continues unabated, present day coral reefs may reach a threshold ofirreversible change. As such, a great variety of measures are being undertaken in attempts to haltor reverse this trend (Jaap, 2000; Rinkevich, 2005). Reaction to the decline of reef ecosystemshas brought about many theories on how to best preserve and restore them. Some currentpreservation methods include: establishing marine protected areas, the suppression and reductionof land-sourced nutrient and sediment runoff, and the installation of mooring buoys at dive sitesto lessen anchor damage. Some current restoration (sometimes referred to as rehabilitation)

methods include: the reattachment of coral colonies and stabilization of reef structures following physical impacts such as ship groundings, whole colony and fragment transplantation fromhealthy reefs to denuded reefs, and the cultivation and targeted transplantation of coral colonies.Additionally, the creation of artificial reefs using materials of opportunity (tires, sunken ships,building debris, rock boulders, etc.) or purpose-built structures (reef balls, EcoReefs) providesnew physical habitat for fish, corals, and other reef organisms in attempts to augment existingreef habitat (Abelson, 2006). However, the effectiveness of artificial reefs as a restoration tool isdebated.


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Coral transplantation has many uses (Table 1), and while whole colonies (wild-sourced) havebeen used in the past, more recent methodologies have adopted the use of cultured specimens forrestoration (Edwards and Clark, 1998; Bowden-Kerby, 1997, 1999; Rinkevich, 2000; Lindahl,2003; Herlan and Lirman, 2008; Grablow et al., 2010; Larson, 2010). The life histories andcharacteristics (fast-growing, branching morphologies) of Acropora species such as A.

cervicornis lend them well to such culture.

Table 1 Examples of Coral Transplantation Efforts(Adapted from Edwards and Clark, 1998)

Reason for Transplantation Study

Aid recovery following dynamite fishing Auberson, 1982; Fox et al., 1999Replace corals killed by thermal effluent Birkeland et al., 1979Save corals threatened by pollution, construction, dredging Plucer-Rosario and Randall, 1987;

Newman and Chuan, 1994Reintroduce species into previously polluted areas Maragos et al., 1985

Accelerate reef recovery following ship groundings Gittings et al., 1988;Hudson and Diaz, 1988

Enhance attractiveness of tourism area Bouchon et al. 1981Rehabilitate tourist-damaged reefs; create artificial reefs Rinkevich, 1995; Oren and Benayahu, 1997to relieve diving pressure

Rehabilitate reefs impacted by natural events such Guzman, 1991; Harriot and Fisk, 1988as El Nio and Crown-of-Thorns sea star

II. Species Profile

The staghorn coral, Acropora cervicornis (Fig.3), is a scleractinian coral species with ageographic range limited to shallow waters of the greater Caribbean. Fixing calcium carbonatefrom seawater, colonies contribute to reef growth and act to fortify and stabilize reefs by activelybinding reef rubble as they spread and encrust onto new substrates (Gillmore and Hall, 1976).Generally inhabiting fore reefs from five to twenty five meters of depth, they have also beenobserved as shallow as one meter to depths of fifty meters (Lewis, 1960; Goreau and Wells,1967; Logan, 1969). Their upper extent is limited by wave action and their lower extent by lightavailability. Additionally, small populations are observed in the patch reef habitats of shelteredback reef and lagoonal areas and, more recently, robust thickets have been described in near-shore waters off Fort Lauderdale, Florida, USA (Vargas-ngel et al., 2003).

Colonies of A. cervicornis are branching, and typically light brown in color. Branches areapproximately two centimeters thick, tapered towards their growing tips, with secondary andtertiary branches commonly budding off of parent branches at nearly 45 angles. Branches arecovered in small, distinctly protruding polyps, and terminate in a single axial polyp. Coloniesgrow in ramose bush-like forms, the nature of which (branch density, branch angle, etc.) canbe influenced by varying hydrological conditions (Bottjer, 1980). Growth ofAcropora species isan important contributor to the physical complexity (also termed rugosity or architectural


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complexity) of the reef environment, which positively correlates with fish and reef organismabundance and species richness (Luckhurst and Luckhurst, 1978; Gratwicke and Speight, 2005).

Figure 3: A colony of the study species,Acropora cervicornis. (photo - E. Larson).

Acropora cervicornis, like numerous other branching scleractinian corals, reproduces by twomeans: sexually and asexually. A hermaphroditic species,A. cervicornis colonies produce bothoocytes and spermatocytes (Szmant, 1986). Spawning is synchronous in nature, with all coloniesof a reef spawning together between two and seven nights after the full moon of July or August(Vargas-ngel et al., 2006). The planktonic products of these events are larval coral planulae.An individual planula can drift in the open ocean for extended periods of time before settling

onto a reef, metamorphosing into a polyp, and beginning the process of binary fission andlimestone deposition as it grows into a new colony.

Asexual reproduction is very common in A. cervicornis and is promoted by its branchedstructure. If subjected to a strong force, often storm-driven wave action or collision with animallife, the rigid branches ofA. cervicornis can break off (in whole or part), falling to the substrate.If a branch fragment settles into a stable position of favorable environmental conditions, it canattach and grow into a new colony. It is primarily in this way thatA. cervicornis colonies spreadon a reef, and it is speculated that asexual, not sexual reproduction is the primary means bywhich A. cervicornis grows and expands its population (Highsmith, 1982; Vargas-ngel et al.,2006). This predisposition to asexual reproduction makes cultivatingA. cervicornis colonies anattractive means of supplying restoration efforts.

III. Artificial Coral Culture

The practice of artificial culture of coral varieties such as Acropora takes advantage of suchcorals predisposition to asexual reproduction by fragmentation (Tunnicliffe, 1981; Highsmith,1982; Clark and Edwards, 1995). In essence, fragments of a desired species are secured to aselected substrate, and left to grow as if they had undergone fragmentation in a natural setting. Itis commonly performed in the aquarium trade by hobbyists and entrepreneurs seeking to


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cultivate (for sale and trade) a great variety of coral species. In the aquarium hobby, cement orplastic plugs are commonly used for a convenient and transportable growth substrate, whereas inreef restoration coral cultivation has been adopted on a much larger scale incorporatingnumerous materials such as cement blocks and pedestals, metal and cement frames, and evensuspended networks made of materials from wire and mesh to PVC and fishing line (Fig. 4)

(Bowden-Kerby, 1997; Thorton et al., 2000; Soong and Chen, 2003; Okubo et al., 2005; Quinnand Kojis, 2006; Shafiret al., 2006; Putchim et al., 2008; Shaish et al., 2008; Levy et al., 2010;Nedimyeret al., 2011). Utilizing cultured corals for restoration also avoids many of the negativeattributes associated with whole colony collection and transplantation between reefs (Table 2).

Figure 4. Cultured specimens ofAcropora cervicornis, growing in a suspendednursery constructed of nautical rigging line and wire.

Admittedly, restoration by transplantation has limitations. Current methods favor the productionof fast-growing, branching species when it has been suggested that the introduction of slower-growing, massive species may be more effective in the long term (Edwards and Clark, 1998). Inaddition, restoration efforts using current methodologies can only affect small areas, and can notbe expected to act as a cure-all for any single species or whole reefs in general. It is unrealisticto assume that with transplantation alone one could maintain the health of the worlds reefsindefinitely. However, what can be achieved by targeted transplantation has further-reachingeffects than initial objectives of site-specific regeneration, beautification, etc (Table 3).

Transplantation-based restoration may be able to create and/or maintain hot spots ofreproductive viability for a species, maintaining its presence in an area where it might be subjectto local extinction. For a species such as A. cervicornis, such hot spots of sexual reproductioncould act as consistent sources of new coral recruits, accelerating its natural process of recovery -one that is currently hindered by sharp declines in recruitment and gene flow. This has been aproduct of both population decreases resulting in decreased spawning volume, and the increasingdifficulty of planulae to settle amongst heightening algal abundance on reefs (Kuffner et al.,2007; Vollmer and Palumbi, 2007).


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Table 2 Potential Drawbacks of Whole Colony Transplantation(Adapted from Edwards and Clark, 1998)

Loss of coral colonies from donor reef areas

Higher mortality rates of transplanted corals

Reduced growth rates of transplanted coralsLoss of transplanted colonies due to attachment failure

Reduced fecundity of transplanted colonies due to transplantation stress

Table 3 Potential Benefits of Coral Transplantation(Adapted from Edwards and Clark, 1998)

Immediate increase in coral cover and diversity

Increased recruitment of coral larvae as a result of presence of transplants

Survival of locally rare and/or threatened coral species when primary habitat is destroyed

Reintroduction of corals to areas which are larval supply limited or have high post-settlement mortality

Improved aesthetics of areas frequented by tourists

Increased rugosity and shelter for herbivores in bare areas

IV. Procedure and Methods

Primary objectives of this project include:

1. Investigate effects of suspended nursery culture on Acropora cervicornis growth andsurvivorship.

2. Determine whether significant differences in growth and survivorship betweengenotypes (previously observed Larson, 2010) persist in the alternative farmingtechnique of suspended culture.

3. Investigate whether suspended culture techniques can broaden the nurseryfragmentation season of A. cervicornis, which is currently limited by high summertemperatures.

To accomplish the first two objectives, six modules will be installed at an existing Acropora

cervicornis nursery site off Ft. Lauderdale, FL. To each module, 24 fragments ofA. cervicornis(at approximately 3cm) will be affixed; for a total of 144 fragments. Fragments of threepredetermined genotypes will be harvested from existing colonies in a neighboring nursery inequal number (48 per genotype), and cut to remove existing apical polyps to reduce experimentvariability. Fragments will not be collected from colonies exhibiting disease (such as white bandor rapid tissue necrosis), prominent bleaching, or severe predation.


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On each module, fragments will be attached with shielded copper wire in two configurations:

A. Directly attached in vertical orientation with polyp apertures facing upward, with 10 -15cm of separation between fragments, to module lines (3/8 polyester nautical rigging)(Fig. 5).

B.

Suspended from horizontal module lines, in horizontal orientation with 10 - 15cm ofseparation between fragments and 10cm of separation from the module line (Fig. 6).Wire will first be affixed to individual fragments, followed by the attachment (andcrimping to prevent lateral shifting) of each wire to the module.

Figure 5 (left): Example of anA. cervicornis fragment directly attached to module line in a verticalorientation.

Figure 6 (right): Example of A. cervicornis fragments attached in a suspended, horizontalorientation.

Each module will consist of a single two meter horizontal line supporting twelve horizontalfragments, and two 2.5 meter vertical lines supporting six vertical fragments each (Fig. 7).

Horizontal fragments will be suspended approximately one meter above the substrate (sand), andvertical fragments are to occupy from approximately one to two meters above the substrate.Modules will be secured to the substrate with two 30 long, 4 diameter screw-in groundanchors, and held upright by two support buoys. To lessen possible shading effects of each buoy(6 diameter) on the uppermost vertical fragments, buffer space (~30cm) will be incorporated.


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Figure 7: Suspended nursery module.

For both orientations, relative fragment position on each module will be determined by stratifiedrandom assortment, such that genotypes are distributed evenly between modules andorientations, but placed randomly on each module. This method of placement will mitigate theseverity of possible losses to any one genotype in the event of module failure or loss.

Upon initial fragment transplantation, and at monthly intervals for a period of one year, multiplecharacteristics of fragment growth will be recorded:

1. Mortality partial and/or full mortality of individual fragments.2. Linear Growth* greatest length (horizontal fragments) or height (vertical

fragments). Since horizontal fragments do not have a fixed point of reference(substratum in traditional methods), linear growth will be taken from the single most-linear distance of a fragment that is judged to most closely follow the central axis ofthe fragment (Fig. 8).

3. Tissue Extension* - fragment/colony branches that are not included in linear growthmeasurements will be recorded once they reach

5mm in length (measurement takenfrom apical tip of new branch to outer surface of source branch not to core of sourcebranch; Fig. 8).

4. Attachment fragments will be denoted as attached once tissue grows overattachment wire (and/or module line for vertical fragments) such that at any point,coral tissue completely envelops attachment wire (Fig. 9).

Ground anchors

Su ort buo s

Vertical fragments

Horizontal fra ments

1m

2m

Support buoys


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5. Bleaching, predation, general stressors general fragment condition will be assessedmonthly, with respect to bleaching (including severity), disease, suspected branchbreakage and/or predation, and overgrowth by macroalgae, hydroids, etc. (Fig. 10).

*Linear growth and tissue extension measurement methods selected to replicate those of previous A.cervicornis growth and survivorship study (Larson, 2010) to allow for direct comparison of growth

data. All length measurements taken to the nearest mm using calipers.

Figure 8 (top): Measurement of suspended colony. Linear growth would be taken as measurement1. Tissue extension taken as the sum of all measurements 1, 2, 3, and 4.

Figure 9 (bottom left): Example of fragment attachment, note coral tissue enveloping attachmentwire. (photo - E. Larson)

Figure 10 (bottom right): Example of hydroid overgrowth, a recorded stressor.

To accomplish objective 3, fragmentation trials will be undertaken in months when watertemperatures exceed the favorable maximum of 27C for nursery fragmentation of A.cervicornis. Each monthly trial will include 36 fragments (12 of each genotype), half of whichwill be transplanted via the traditional puck and epoxy method (Larson, 2010 - Fig. 11) at theexisting fixed (hard substrate) nursery occupying the same site, while the remaining half will betransplanted using the aforementioned horizontally suspended attachment method. Growthmeasurements (utilizing the same data collection procedure as in previous component) will only be undertaken upon fragmentation and in the final monitoring effort. General fragmentcondition, with specific attention to mortality, bleaching, and disease will be recorded monthlyfor both suspended and fixed fragments until the conclusion of the study. In situ

1

2 3

4


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temperature loggers at the nursery will allow for fragment condition to be assessed againsttemperature. Since no significant difference in fragment survival was observed betweengenotypes 4, 8, and 10 in the original study by Larson, genotype 6 (which exhibited significantlylower survival) will be used in place of genotype 8 in this component.

Fragments will be installed on additional horizontal tiers that will be outfitted to existingmodules (Fig. 12). Fragment position will be determined by the same stratified random sampletechnique utilized for the initial transplantation.

In previous studies, fragment mortality has been observed at rates greater than 50% whentransplantation was conducted above 27C (56% mortality observed 35 days after fragmentationin ~29C conditions, September 2007 transplantation - Larson, 2010). Increased tolerance tothermal stress has been observed, but not quantified by other facilities conducting suspendednursery culture of A. cervicornis (Ken Nedimyer and Katie Grablow of Coral RestorationFoundation, personal communication, October 2010). If acceptable survivorship is observed insuspended fragments transplanted during summer months, it would demonstrate a degree ofversatility of suspended culture by allowing for the expansion of the fragmentation season.

Figure 11 (left): A. cervicornis fragments attached via traditional puck-epoxy method. (photo - E.Larson)

Figure 12 (right): Suspended module diagram highlighting summer fragment addition.

Statistical Comparison

Much data are available from the previous nursery study (Larson, 2010) of the three genotypes(designated 4, 8, and 10), allowing for direct comparison of the following:

1. Survival - Overall: no significant difference between genotype survival (4, 8, 10 overall)was observed in the previous study (December transplantation). Approximately 90%survival was observed for genotype 4 and 8, with 100% for 10.

2. Survival Orientation: no significant difference was found between horizontal andvertical fragments transplanted in December of the previous study, although verticalfragments exhibited higher survival (83%) than horizontal fragments (62%). Data forindividual genotypes may yet show significant differences.

Summer fragments


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3. Survival Seasonal: 56% and 42% mortality was observed in the September and Octobertransplantations of the previous study, respectively. The planned summer transplantationtrials of this study will incorporate concurrent puck-epoxy transplants for comparison, aswell as encompass the heightened temperature conditions which produced increasedmortality. If observed, significant increases in suspended fragment survival will illustrate

the positive effects of suspended culture techniques on fragment thermal stress tolerance.4. Disease: Genotypes 4, 8, and 10 exhibited zero, three, and one disease event in the

previous study respectively.

5. Growth (Linear Growth and Tissue Extension): per-genotype monthly growth rates areavailable from the previous study, and may be used to compare to measured growth rates(both overall and by month) of current study. Points of comparison include: difference between maximum and minimum growth rates (within genotypes), difference betweenoverall growth rates between genotypes, differences between seasonal maximum andminimum growth rates of genotypes between studies, and the comparison of overallgrowth rates of genotypes by attachment method between studies.

Genotype 4 exhibited the greatest tissue extension in the previous study, significantlyhigher than genotype 10 which showed the lowest tissue extension. No significantdifferences in linear growth were observed between the three genotypes (afterincorporating for impacts to fragments such as disease).

6. Branching Frequency: In the previous study, genotype 4 exhibited the highest branchingfrequency whereas genotype 10 exhibited the lowest frequency.

V. Anticipated Results

A. Suspended, horizontal fragments will exhibit greater overall tissue extension and branchnumber than vertical fragments. Horizontal transplantations have exhibited greater branching than vertically transplanted fragments previously. Additionally, suspendedfragments will have a near-unlimited range of direction in which to grow, versus verticalfragments which will be comparatively encumbered by the presence of the module line.

B. Higher survivorship (per genotype) than previous study for suspended fragments.Preliminary findings of other Acropora cervicornis nurseries employing suspendedculture (Coral Restoration Foundation, Ken Nedimyer, personal communication) haveshown increased survivability of suspended fragments compared to epoxied transplants.

C. Higher temperature-related survivability in suspended fragments (summertransplantations). A significant difference in survival is expected between suspended and puck-mounted fragments during the warm-water transplantation trials; the formerexhibiting greater survivability. High (90%) survivorship is not expected during thewarmest months (August and September averaged 30C and 29C in 2010 respectively), but may occur when temperatures approach the preferred 27C temperature ceiling(October and November 2010 averaged 28C and 25C).

D. Preserved: no significant differences in survivorship between genotypes or orientations.As overall mortality is expected to decrease, relative differences in survivorship willcorrespondingly decrease.


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VI. Merit

Broadening the knowledge of the physiological responses in corals undergoing fragmentation-based reproduction is essential in this age of reef degradation. The relatively young practice ofcoral aquaculture, a method currently utilized in reef restoration, may be greatly furthered by

advances in asexual reproduction of corals. In the case ofAcropora cervicornis, an importantspecies of Caribbean reefs, continuing the research of present and future methods of itsrestoration is paramount. The role thatA. cervicornis and similar coral species play is essentialto the formation, growth, and continued existence of Caribbean coral reefs. Outcomes of thisstudy could effect changes in propagation procedure to better suit the aims of numerous reefrestoration and augmentation efforts around the world, specifically those concerning A.cervicornis in South Florida and the greater Caribbean. Additionally, potential differences insurvival, bleaching stress and disease occurrence between suspended colonies and those beinggrown on substrate may provide insights into the influence water flow and/or interactions withthe benthic community have onA. cervicornis physiology.

VII. Funding and Feasibility

Research costs, including material and monitoring expenses (including boat excursions) will besupported by The Nature Conservancy (TNC), the NOAA Restoration Center, and funding fromthe American Recovery and Reinvestment Act. Laboratory space, equipment, and personnelassistance will be provided by the Benthic Assessment, Monitoring and Restoration Laboratoryof Nova Southeastern University Oceanographic Center (NSUOC) headed by Dr. David Gilliam.Monitoring efforts will be conducted through the NSUOC AAUS scientific diver program.Boating operations will be conducted under Oceanographic Research Vessel (ORV) distinctionthrough NSUOC.

This research will be supported by The Nature Conservancy (TNC) and the NOAA Restoration Center, U.S. Department of Commerce, with funding from the American Recovery and Reinvestment Act (Award

#NA09NFF4630332). The statements, findings, conclusions, and recommendations are those of the author(s) and

do not necessarily reflect the views of The Nature Conservancy, the NOAA Restoration Center or the U.S.

Department of Commerce.


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