The Octocoral Fishery in the Southeastern U.S. and Gulf of Mexico

MARK CHIAPPONE, PAOLA ESPITIA, LEANNE M. RUTTEN, and STEVEN L. MILLER

Introduction with ~1.8 million households owning Conservation Commission (FWC) in The marine aquarium hobby has a saltwater aquarium (American Pet Florida. Regulation of the collection witnessed a resurgence in popular- Products Association, 2014). Within of live tropical, ornamental, and ma- ity within the United States since the the United States, Florida represents rine in Florida is governed by 2007–09 recession (Bricker et al.1), the largest component of the ornamen- Rule Chapter 68B-42 of the Florida tal fishery, supplying live marine life Administrative Code (FAC). The Flori- 1Bricker J. B., Bucks, A. Kennickell, T. Mach, to the aquarium industry with over 9 da regulations went into effect in 1991 and K. Moore. 2011. Surveying the aftermath of million individual per year, and have undergone several revisions, the storm: changes in family finances from 2007 encompassing over 600 fish, inver- affecting bag limits, proper collection to 2009. Working paper 2011–17, Mar. 2011, Fed. Reserve Bd., Finance and Econ. Discussion tebrate, and plant species (Rhyne et techniques, reporting requirements, Ser., 38 p. (Online at https://www.federalreserve. al., 2009). Management of this multi- and area restrictions (and more) for gov/pubs/feds/2011/201117/201117pap.pdf). species fishery is under the authority recreational and commercial marine of the Federal Gulf of Mexico Fish- life collectors (Larkin et al., 2001). The authors are with the Halmos College of Natural Science and Oceanography, Nova ery Management Council, the South With a Saltwater Products License, Southeastern University, 8000 N. Ocean Drive, Atlantic Fishery Management Coun- commercial collectors can exceed the Dania Beach, FL 33004. Corresponding author cil in EEZ waters off North Carolina, daily recreational bag limits that differ is Mark Chiappone (email: [email protected]). South Carolina, and Georgia, and the by species group. To collect and sell doi: https://doi.org/10.7755/MFR.80.3.2 State of Florida’s Fish and Wildlife tropical fish and invertebrate species

ABSTRACT−Octocorals, also known This study assessed some of the character- collected in the Florida Keys in two of the as gorgonians and soft , were previ- istics of the octocoral fishery by evaluating collector categories represented < 0.004% ously managed by the U.S. South Atlantic FWC Trip Ticket data, conducting interviews of the estimated population sizes. The col- (SAFMC) and Gulf of Mexico (GMFMC) with octocoral collectors, and analyzing oc- lectors and aquarium hobbyists interviewed Fishery Management Councils through tocoral life history information and avail- stated that they would welcome and use a a joint Fishery Management Plan able fishery-independent data on population field guide to help with octocoral identifica- (FMP). Because octocorals are mostly col- densities and sizes. Based on interviews with tion, which would ultimately provide better lected from Florida waters, the Florida collectors, a synthesis of trip ticket results, fishery-dependent taxonomic resolution for Fish and Wildlife Conservation Commission and population abundance estimates, the collected species. The collection of octo- (FWC) is currently tasked with managing long-term stability of the octocoral fishery is corals below the State of Florida quota of octocoral collection, including the moni- not likely to change significantly. The social 70,000 colonies per year threshold, which toring of colony landings, in the Economic dynamics of the aquarium industry to seek has yet to be reached according to landings Exclusive Zone (EEZ) adjacent to Florida. colorful, rare, and exotic marine species for data, likely does not adversely affect the Collection of 70,000 colonies per year total, home aquaria places octocorals at the lower octocoral populations targeted. This con- which applies to both state and EEZ waters end of the list of desired species. Octocoral clusion is based upon the large population off Florida, is permitted under Rule 68B- distribution and abundance information ob- estimates determined for octocorals relative 42.006 of the Florida Administrative Code, tained from an extensive search of the lit- to the small number of colonies collected. but has never been exceeded according to erature, along with available age, growth, This conclusion assumes that the distribu- available landings data. Before octocoral and habitat data, suggests that current in- tion and population sizes of targeted spe- management responsibility was transferred formation is comprehensive and definitive, cies will continue to be relatively stable or to the State of Florida, the SAFMC was con- such that stock assessment or population dy- increase. Information collected by the State cerned that octocoral landings data, as op- namic modeling could be considered but are of Florida through trip tickets is probably posed to population data, were being used probably not required to assess the status adequate to understand and manage the to set Acceptable Biological Catch limits of collected octocoral species. For multiple octocoral fishery. Minor reporting clarifi- and Overfishing limits under the new Fed- sampling periods, over a decadal period cations and better taxonomic resolution in eral fishery management standards. An ad- (1999–09) in the Florida Keys, where most reporting would help improve the accuracy ditional concern with the octocoral fishery octocoral collection occurs, abundance es- of collecting data, but improved accuracy and other organisms captured for the marine timates presented for 15 species illustrate is not required to assess the current state of aquarium and ornamental fisheries is that that population sizes are large (tens of mil- the fishery—the octocoral fishery is sustain- large taxonomic groups are lumped togeth- lions to hundreds of millions of colonies, per able and would likely remain sustainable at er, including species with potentially differ- species) and abundance is stable or increas- colony collection levels orders of magnitude ent life histories and ecological functions. ing. For example, the numbers of colonies larger.

18 Marine Fisheries Review such as mollusks, plants, live rock, ferable Dive (MLN), and Marine Life Deep Water Habitat Area of Particu- and live sand, a Restricted Species en- Bycatch (MLB). To qualify for these lar Concern adjacent to Florida waters dorsement and Marine Life endorse- endorsements, collectors had to show (FAC 68B-42.0036). ment are both required. proof of commercial marine life land- During collection, no power tools The State of Florida has collected ings and sales above a $5,000 thresh- may be used and only one inch of commercial landings and effort data old between the licensing years of substrate around the perimeter of the since November 1984, maintained 1999 and 2003 (FAC 68B-42.0065). base of the octocoral holdfast may be in their Marine Fisheries Informa- This process eliminated the part-time removed (FAC 68B-42.007 and 68B- tion System, commonly referred to collectors and brought the number of 42.008). Octocorals must be collected as “Trip Ticket” data (Larkin et al., active ML endorsement holders in the alive and stored in a re-circulating live 2001). State of Florida landings data State of Florida down from 743 to 168 well or oxygenated system aboard the on marine life collection date back to (Adams et al., 2001). About 10% of all collection vessel (FAC 68B-42.0035). 1994. The Florida Marine Life Asso- ML endorsements are currently MLN There are additional area closures for ciation (FMLA), the official industry (M. Bademan, FWC, personal com- marine life collection in southeast- organization for marine life collectors, mun.). Collectors wishing to obtain an ern Florida, including National Parks petitioned the Florida Marine Fish- ML endorsement may now only do so (Everglades, Biscayne, Dry Tortugas), eries Commission in 1988 to adopt through the transfer of a pre-existing John Pennekamp Coral Reef State standards for the collection of tropi- endorsement, since no new endorse- Park, and portions of the Florida Keys cal marine life species. Catch regula- ments are currently being issued. National Marine Sanctuary, includ- tions were adopted as a result and the ing the Key Largo Management Area collection of marine life and tropical Management and Regulation (formerly Key Largo National Marine ornamental data were included in the of the Fishery Sanctuary), the Looe Key Manage- Trip Ticket reporting system begin- Octocorals were managed in the ment Area (formerly Looe Key Na- ning in 1994. Originally designed to U.S. South Atlantic (North Carolina tional Marine Sanctuary), and various track the sales of seafood products to Florida) and U.S. Gulf of Mexico smaller no-take zones including Sanc- from Florida waters, the Trip Ticket (Florida to Texas) from 1990–2011 tuary Preservation Areas, Special-Use/ reporting program requires wholesale under the joint Gulf of Mexico and Research-Only Areas, and Ecological dealers to report information about South Atlantic Coral Fishery Manage- Reserves (15 CFR 922 Subpart P). saltwater products purchased (Section ment Plan (FMP). Restructuring of 379.362, Florida Statutes, and 68E- the Coral FMP in 2011 (Amendment Collector and Fishery 5.002, FAC). 7) subsequently excluded the manage- Characteristics Subsequent concerns over the num- ment of octocorals in Federal waters Collector Demographics ber of new Marine Life (ML) en- off Florida, thereby transferring man- dorsement holders in the industry agement responsibility to the State of Of the 168 ML tiered endorsement holders in Florida in 2013, only 64 prompted the FMLA to petition the Florida (FAC 68B-42.009). Octocorals 2 FWC to limit the entry of new ma- are defined by the State of Florida as reported octocorals landings. After rine life collectors (Division of Ma- “any erect, non-encrusting species of consulting two marine life collectors rine Fisheries, 2002, cited in Larkin the Subclass , except the familiar with the fishery, 53 octo- et al., 2001), although an assessment species flabellum and G. coral collectors were identified as ac- of past and current trends about pos- ventalina” (FAC 68B-42.002). Rec- tive collectors based in the Florida sible exploitation of individual species reational collectors may collect up to Keys (n=30 collectors), the Florida had not been conducted (Larkin et al., six octocoral colonies/day with a Flor- east coast (n=14), and the Florida west 2001). As a result, the State of Flori- ida Recreational Saltwater Fishing Li- coast (n=9). A letter that described the da passed Senate Bill 1506 in 1998, cense (FAC 68B-42.005). There are no goals of this study with a request for which imposed a 4-yr moratorium on trip or daily collection limits on com- additional information about the octo- the issue of new ML endorsements mercial landings; however, the annual coral fishery was sent to the 53 active (Florida Statute 370.06(2)(d)(2)). Fur- quota for octocorals collected in State collectors. A second (and in some cas- ther restrictions were introduced in of Florida and adjacent Federal wa- es a third) request for information was 2005 for those wanting to collect or ters is currently 70,000 colonies (FAC sent via email to collectors (n=30). sell marine life species. In addition 68B-42.006). Area restrictions on oc- Telephone and in-person interviews to a Saltwater Products License and a tocoral collection exist in Atlantic Fed- were conducted with 15 collectors, Restricted Species endorsement, col- eral waters north of Cape Canaveral, including 8 from the Florida Keys, 5 lectors needed to apply and qualify Biscayne National Park, and in the from the Florida east coast, and 2 from for a ML endorsement. The endorse- Stetson-Miami Terrace Deep Water 2Florida Fish and Wildlife Conservation Com- ments are Marine Life Transferable Coral Habitat Area of Particular Con- mission, Tallahasse, Florida. Unpubl. data, Nov. Dive (MLD), Marine Life Non-trans- cern, as well as the Pourtales-Terrace 2014.

80(3) 19 the Florida west coast. Interview ques- Table 1.—Octocorals typically collected in the Atlantic Ocean off Florida (H. Feddern, personal communication). tions addressed the fishery (i.e., spe- Species group Scientific name Collector common name cies collected, colony size, collection Other Diodogorgia nodulifera yellow finger/colorful sea rod gorgonian sites, and collection frequency), col- sp. brown tree gorgonian Muricea sp. silver gorgonian lection methods (i.e., collection tech- nutans large gorgonian niques, holding facility set-up, and Pterogorgia citrina yellow ribbon gorgonian shipping logistics), and social issues Purple Antillogorgia sp. purple plume gorgonian Antillogorgia sp. purple willow gorgonian within the fishery (i.e., purpose for Antillogorgia elisabethae purple frilly gorgonian starting a business, amount of time flavida purple brush gorgonian Pterogorgia anceps purple ribbon gorgonian spent collecting, and collector eti- quette). Information gathered from in- Red Diodogorgia nodulifera red finger gorgonian terviews and field visits to the Florida exserta orange tree gorgonian Keys (5 collectors visited) and Miami (2 collectors visited) was used to sum- marize octocoral fishermen and fishery Octocorals are mainly collected for octocoral collection. As a result, ma- characteristics. the aquarium trade and not for the rine life collectors spread their efforts In the State of Florida, octocoral curio trade. A few opportunistic ML across collecting and selling marine collectors are required to hold a Salt- endorsement holders hold a MLB en- fishes and a variety of invertebrates water Products License, Restricted dorsement, which allows them to keep such as crabs, corallimorpharians, Species endorsement, and a ML tiered whole octocorals that are incidentally anemones, and snails. Table 1 lists the endorsement (MLD, MLN, or MLB). caught by nets. Since the qual- octocorals commonly collected for the These endorsements were either pur- ity of specimens obtained from by- aquarium trade in Florida (Feddern4) chased before the moratorium on the catch is typically not suitable for the and Table 2 is an expanded list of spe- issue of new ML endorsements in marine life trade due to tissue damage cies that was provided by marine life 2005, or they were purchased from an and broken branches, such colonies collectors interviewed and by FWC. existing license holder, typically as a are dried and sold for the curio trade.3 package including the entire business Because shallow-water octocorals are Collection Methods (e.g., boat, warehouse, holding tanks, found in great abundance through- Aquarium owners generally prefer and collection sites). This is an ex- out hard-bottom and coral reef habi- clean, brightly colored purple, red, or pensive purchase considering the cost tats of south Florida, these fauna are yellow octocoral specimens that are (~$35,000 US) of the ML endorse- typically easy to find and can be col- heavily branched with large polyps in ment.3 Many (n=11) of the collectors lected year-round. In contrast, because good condition (e.g., no visible signs interviewed are lifelong ocean and deeper water (> 30 m depth) species of tissue loss or bleaching). Size mat- aquarium enthusiasts who have been such as Diodogorgia nodulifera, Swif- ters, with specimens usually small diving since the 1960s and 1970s; tia exserta, and spp. are enough to fit comfortably in home thus, they have extensive experience more challenging to collect because of aquariums (< 20 cm in height). Oc- to know where to efficiently find their strong currents and the constraints of tocorals are collected using snorkel- target organisms. Other collectors deeper diving, only a few of the col- ing, SCUBA, or hookah diving rigs. (n=4) learned the trade through an ap- lectors interviewed target deeper water A hammer and chisel or dive knife is prenticeship of sorts, helping a current (> 30 m) species. typically used to remove whole colo- ML holder while learning proper col- Despite the general ease in finding nies (Fig. 1). As per FAC 68B-42.007, lection techniques. Over half (58%) of octocorals, no ML endorsement hold- the holdfast and substrate with a maxi- the octocoral collectors contacted live ers collect and sell octocorals exclu- mum diameter 1-in or less can be re- in the Florida Keys (Monroe County), sively. In a few cases (n=3), octocorals moved along with the colony. Some of with the remaining residing on either represent nearly 50% of total marine the collectors interviewed believe that the Florida east coast (22%) or west life sales for collectors; however, more preserving the substrate with the hold- coast (20%). While octocorals are col- commonly (n=12) octocorals comprise fast increases the survivorship of a col- lected throughout Florida waters, 80% 20% or less of total marine life sales. ony, but no formal studies have been of reported landings come from Atlan- The fishery is demand-driven, which conducted to test this observation. tic waters2, with the majority of the means that octocorals are primarily Octocorals collected off the west collection effort focused on Florida collected upon request from whole- coast of Florida, such as Leptogorgia Keys shallow-water species. salers, retail stores, public aquariums, spp., are often removed by hand from and aquarium hobbyists. The demand nearshore artificial reefs and bridge for octocorals within the aquarium in- 3Anonymous marine life collectors interviewed in southeastern Florida, Personal commun., dustry has never been sufficient to sup- 4Feddern, Henry. Marine life collector, Florida 2013–14. port a business focused exclusively on Keys. Personal commun., Oct., 2013.

20 Marine Fisheries Review pilings. This is similar in the Florida Keys; however, both locations also involve substantial collection direct- ly from the natural seabed. Collected colonies are placed in mesh bags or plastic collection containers during the dive, and then stored in covered species-specific live wells or 5-gallon buckets with air stones during trans- port to a holding facility. Water chang- es are done throughout the day (P. Espitia, personal observ.). Collectors usually fulfill octocoral orders within an hour, so the remainder of the day is spent collecting other marine spe- cies. Collectors with wholesale orders noted that they collect as many as 600 octocorals per month, and then distrib- ute several large orders to wholesalers. Collection sites are species-specific and several types of marine life (e.g., emerald crabs, Mithraculus sculptus) may be targeted along with octocorals Figure 1.—Octocoral collector, Dr. H. Feddern, using a hammer and chisel to remove at a single site. All of the collectors octocoral colonies to be sold in the aquarium trade. interviewed specified that they have multiple sites from which colonies are removed and that they are conscious to

Table 2.—Octocorals collected in the Atlantic Ocean and the Gulf of Mexico off Florida based upon personal communi- rotate sites to reduce collecting stress cations with Florida collectors during 2013–14. Asterisk (*) indicate the most highly demanded octocoral taxa. on individual site locations. Although Species Collection collectors do not officially share their group Scientific name Collector common name(s) location work site locations with each oth- Other Antillogorgia acerosa golden plume gorgonian Atlantic er, they generally avoid one another. Diodogorgia nodulifera* yellow finger/colorful sea rod gorgonian Atlantic Muricea sp.* delicate spiny sea rod Atlantic Some Florida Keys marine life col- Muricea elongata orange spiny sea rod/rusty gorgonian Atlantic Muricea muricata spiny sea fan Atlantic lectors noted that in the past they Muricea pinnata long spiny sea fan/silver gorgonian Atlantic were followed by other collectors who homomalla black sea rod Atlantic Plexaurella fusifera brown tree gorgonian Atlantic would try and learn where they were * giant slit pore/large polyp gorgonian Atlantic collecting, especially where cryptic or Pterogorgia citrina* yellow ribbon/yellow sea whip gorgonian Atlantic prized specimens that are worth more Purple Antillogorgia spp. purple frilly/purple whip gorgonian Atlantic, Gulf money were located. Antillogorgia acerosa* Purple or golden willow gorgonian Atlantic Antillogorgia elisabethae* purple frilly gorgonian Atlantic The octocoral collectors interviewed Antillogorgia kallos* purple plume/bi-pinnate gorgonian Atlantic Briareum asbestinum* corky sea finger/sea stalk gorgonian Atlantic stated they are cautious about storage Eunicea spp. purple knobby/knobby sea rod/warty sea rod gorgonian Atlantic methods, because negative interac- Eunicea flexuosa* purple candelabra/swollen knob candelabrum gorgonian Atlantic Muriceopsis flavida* purple brush/bottle brush gorgonian Atlantic tions between species can occur. For sp. porous sea rod Atlantic example, the slimy sea plume, Antil- purple tree gorgonian Atlantic Pterogorgia anceps* purple ribbon/angular sea whip gorgonian Atlantic, Gulf logorgia Americana, is not sold in the Pterogorgia guadalupensis purple blade/flat-blade/grooved-blade sea whip gorgonian Gulf aquarium trade because the excess

Red Diodogorgia nodulifera* red finger/colorful sea rod gorgonian Atlantic slime it produces fouls the water dur- barbadensis Devil’s sea whip Atlantic Icilliogorgia schrammi orange deep-water fan Atlantic ing storage and shipment, killing other Leptogorgia punicea sea whip/carmine sea spray gorgonian Atlantic, Gulf octocorals within the same aquarium Leptogorgia virgulata sea whip gorgonian Gulf Nicella schmitti red/bushy sea whip Atlantic system. Other “sea plumes” (Antil- Swiftia exserta* orange tree/red polyp gorgonian Atlantic logorgia spp.) reportedly are caustic to other octocoral genera. The high concentration of prostaglandins in (Bundy, 1985) causes caustic reactions when stored with other octocorals, even individu-

80(3) 21 Figure 3.—Eco-Gorgs® are kept upright in flat plastic egg crates within a greenhouse holding facility in the Florida Keys.

Figure 2.—(Greenhouse style octocoral holding facility in the Florida Keys.

als from the same species. One collec- store octocorals to maximize profit by treated water. Octocoral colonies with tor reported that some morphologies minimizing the number of specimens slender branches, such as sea plumes, of Eunicea flexuosa will “burn” the collected and maximizing their sur- are stored upright with their holdfasts tissue of other octocorals, causing it vival after collection. This contrasts, buried in a bed of sand or fine gravel to slough off. Plexaurella species and for example, with the way that some substrate, while those with more ro- some deepwater species such as El- marine ornamental species, particular- bust branches (e.g., Eunicea and Pseu- lisella barbadensis, Nicella schmitti, ly fishes, are collected internationally doplexaura species) are suspended and Swiftia exserta are reportedly sen- such as in the Philippines, where large upside-down with their holdfasts se- sitive to physical damage and to al- numbers are collected from reefs and cured within a piece of foam (similar lelochemical interactions with other only a few survive to be sold to con- to a pool noodle). Fragmented colo- species, so collectors keep these fauna sumers (Wood, 2001). nies can be stored upright with their in individual bags during collection Collected octocorals are usually plugs secured within sheets of plastic and storage. Collectors who sell the housed in long raceway aquaculture egg crates (Fig. 3). deepwater octocoral Diodogorgia nod- tanks (~4.6 m x 0.6 m x 0.6 m) located The length of time octocorals are ulifera, on the other hand, described in greenhouses or warehouses (Fig. 2). held within a warehouse facility be- this species as hardy and indicated that Two of the facilities visited have di- fore being shipped ranges from several colonies can be housed in close prox- rect access to bay or inlet waters and hours to several weeks. Some collec- imity with conspecifics and S. exser- had a flow-through water filtration sys- tors wrap wet paper towels or news- ta. While none of this information is tem. Other operations mix their own paper around individual colonies, and proprietary, collectors have learned by saltwater (n=12) or have it trucked then place six wrapped colonies in a trial and error how to best collect and in (n=1), and then ship octocorals in plastic bag with a small amount of wa-

22 Marine Fisheries Review Figure 5.—Eunicea flexuosafrag - ment is super-glued into a concrete plug to be sold as an Eco-Gorg®. Figure 4.—Branch fragments being clipped from a Eunicea flexuosa colony to be sold as Eco-Gorgs®. ter. Wrapping the colonies keeps them A growing number of collectors are um asbestinum, Eunicea calyculata, E. moist and self-contained, which helps adopting the Eco-Gorg®5 technique, flexuosa, E. succinea, Muricea elon- to minimize contact with other colo- in existence since at least the early gata, Plexaura homomalla, and Ptero- nies if they start to deteriorate during 1990s, to reduce potential collecting gorgia guadalupensis. One deepwater shipping. For high-volume shipments impacts on colonies and populations. collector interviewed uses this tech- of 100 or more colonies, this technique The Eco-Gorg® technique involves nique to collect Swiftia exserta more is too labor intensive. Some collectors clipping branch tips ~10-cm in length quickly and efficiently in challenging have had inconsistent results with wet from larger established colonies (Fig. high current, deep-diving conditions. paper towels, so they instead pack in- 4), instead of removing entire colonies S. exserta is collected in Palm Beach dividual colonies in small plastic bags from the seabed. From 1 to 20 branch County along the edges of the south- and inject hospital-grade oxygen along tips per colony are removed and after east Florida reef tract from 18–46 m with a small amount of water to keep the colony base of each clipping is depth, where high densities of large the colonies wet. Another method used stripped of its tissue, they are inserted colonies are intermixed with Diodo- is to place only the holdfast in water into concrete plugs and super-glued gorgia nodulifera and M. elongata. and then hand-vacuum-seal the bag; into place (Fig. 5). The clippings are Colonies tend to be short and rigid in however, many collectors noted that kept in holding tanks for a few weeks high-current environments. One col- air exposure kills the colonies. De- to allow tissue to grow over the con- lector interviewed also finds these pending on the species, a dozen or so crete plug. The collectors interviewed deeper water taxa in Miami-Dade colonies can be placed in a single bag stated that if they keep octocorals lon- County at 27 m depth; however, den- (e.g., D. nodulifera and Leptogorgia ger in their holding facility, then the sities are lower and colonies are taller spp.), while other octocorals must be colonies acclimate better to an aquar- and more flexible at these sites. Be- packed individually (e.g., P. homomal- ium environment, thus making them tween five and ten branch clippings, la). To minimize costs associated with hardier for customers. Species frag- 20-cm long, are removed from colo- colony holding and shipping, one col- mented and marketed as Eco-Gorgs® nies over one meter in height. As many lector transfers his collection from the include Antillogorgia acerosa, Briare- as 70 clippings can be collected in ten vessel to live wells in the back of a minutes. Not all octocorals can be col- truck, and then delivers the octocorals lected using the Eco-Gorg® technique. 5Mention of trade names or commercial firms directly to wholesalers, which keeps does not imply endorsement by the National Ma- D. nodulifera colonies, for example, the process simple and lowers costs. rine Fisheries Service, NOAA. lack a proteinaceous central axis and

80(3) 23 thus cannot be adhered to a plug or and Coral Magazine (coralmagazine. warm temperatures in south Florida in other substrate for growing out in a com). Such networks have contributed 2014 resulted in massive bleaching of holding facility. greatly to the expanding knowledge a number of octocoral species includ- Aquarium hobbyists generally de- base of octocoral care and sales. ing Antillogorgia spp., B. asbestinum, sire fragmented colonies for several The international market for Carib- Muricea sp., and Plexaurella spp. In reasons. First, they are a sustainable bean octocorals exists because inter- addition, elevated concentrations of alternative to removing whole colonies national hobbyists seek out exotic, the marine Karenia bre- from the reef. In addition, they offer seemingly rare species to keep in their vis created an algal bloom off the west the potential to introduce more vari- home aquariums. The collectors inter- coast of Florida6, killing a substan- ety to aquariums because specimens viewed stated that they ship octocoral tial number of octocorals as stated by that may otherwise be too large can colonies to Canada, Israel, Ukraine, one collector. In such cases, collectors be made available as smaller colonies. and Asia, with several also shipping must travel further distances to find The fragments also tend to be har- to the European Union despite need- suitable specimens to collect, result- dier, since they have already survived ing a health certificate from the U.S. ing in price increases to offset extra storage in the artificial systems main- Department of Agriculture, along with fuel costs. Deepwater octocoral collec- tained by the collectors or wholesal- veterinarian inspections of collection tors have an apparent competitive ad- ers. Aquarium hobbyists also state that stocks and holding facilities. - vantage over shallow-water collectors Eco-Gorg® fragments are easy to es- ny in particular has become a leading because demand for their specimens tablish in their aquariums because they manufacturer of aquariums and aquar- is relatively high, due to the vibrant are already accustomed to receiving ium products that serve a popular reef- coloration (red, orange, and yellow) fragments of stony corals and coralli- keeping hobby. When the Cold War of the deepwater colonies. Collectors morpharians attached to plugs. Marine political divisions blocked access to also work to emphasize the “special” life vendors have begun to adopt the aquarium resources, aquarists in East nature of deepwater octocorals by par- fragmentation process and on occa- Germany were forced to dedicate their tially filling orders, or only collecting sion request large whole colonies that husbandry efforts to a limited number these species every couple of months. they clip themselves and then sell as of species, mainly soft corals from the Intentional or not, the perception re- fragments. Mediterranean Sea, and to construct mains that deepwater octocorals are their own filtration systems. The re- “special,” so this places a higher val- Economics of the Fishery unification of Germany gave rise to ue on colonies (Anonymous collector, Octocoral collectors sell to whole- greater access to tropical marine - personal commun.). However, deepwa- salers, retail stores, hobbyists, and mals; however, the popularity of reef- ter octocorals are also more difficult to public aquariums both domestically centric aquariums remained and now maintain in good condition in aquaria, and internationally. One interviewed many home aquariums in Germany are reducing demand. collector, for example, supplies octo- dominated by invertebrates, especially Internal costs associated with col- corals primarily to public aquariums, octocorals (Luty, 2014). lecting marine life organisms are high but also sells to wholesalers in Chi- The mark-up rate from collector to and include operational costs, mainte- na and Japan and a few domestic pet retailer is approximately 10-fold, with nance costs, and other associated costs stores. Wholesale dealers and retail retailers purchasing colonies from col- (Table 3). Per week, the collectors in- stores work closely with one another, lectors for $2 to $5 per colony and then terviewed stated that they could spend so marine life collectors are careful selling those colonies for $20 to $50 $400 on collection trips. For this rea- not to undercut wholesalers by dealing each. Working with wholesalers tends son, octocoral collectors sometimes directly with retail stores. to bring reduced prices per colony for work with other marine life collectors Collectors who sell directly to hob- the collector; however, collectors can to fill gaps in their own supplies. Oth- byists are typically working with a sell 100 colonies at a time rather than er costs include annual renewal fees “purist” market—where buyers are just 5–15 colonies. In addition, once such as the $550 fee for a Wholesale trying to recreate a reef ecosystem a relationship is established between Dealer License. Unforeseen costs in- in their home aquaria. Buyers look- a collector and a wholesaler, whole- clude fines from FWC; for example, ing for high-quality Caribbean spe- salers show loyalty to their collectors for not using an air stone or recircu- cies that are sold at reasonable prices and will subsequently purchase from lating live well to keep octocorals in a hear about these direct-sale-collectors them despite competitors offering low- healthy condition, or the revocation of by word-of-mouth. This results in a er prices. Collectors do have influence an ML license for failure to report Trip steady demand for octocorals that is over their distribution and prices, es- Ticket data (e.g., loss of the purchase additionally driven by social media pecially when healthy specimens are outlets and marine aquarium forums difficult to find or if collecting condi- 6FWC. 2014. Red tide current status, accessed such as ReefBuilders (reefbuilders. tions such as inclement weather make 14 Dec. 2014 at https://myfwc.com/research/ com), Reef Central (reefcentral.com), collecting challenging. The extensive redtide/statewide/.

24 Marine Fisheries Review Table 3.—Costs associated with owning a marine life Table 4.—Annual landings, number of collection trips, average price per colony, and estimated value of octo- collection business according to interviewed octo- corals in State and Federal waters in Florida, 1994–2012, based upon commercial marine life trip ticket data coral collectors in Florida. (FWC, 2014).

Costs Year No. collected Total trips Average price/colony ($) Estimated value ($) Operational 1994 32,106 764 3.80 121,983 Fuel (personal vehicles) 1995 35,976 889 2.42 87,186 Fuel (boat) 1996 37,057 702 2.80 103,739 Warehouse facility rental/mortgage 1997 44,867 725 2.34 104,866 Aquarium system setup 1998 40,743 832 2.23 91,029 Electricity 1999 36,621 790 2.23 81,662 Staff/crew, food, ice 2000 43,100 997 2.50 107,914 2001 45,894 956 2.60 119,304 Maintenance 2002 38,138 880 2.51 95,731 SCUBA gear 2003 44,484 926 2.63 117,140 SCUBA tank fills 2004 43,708 1,018 2.74 119,802 Boat 2005 41,452 905 2.88 119,515 Aquarium products (water, salt, etc.) 2006 48,567 1,004 2.96 143,538 Packing and shipping supplies 2007 44,273 864 3.29 145,759 2008 42,642 826 3.57 152,029 Associated 2009 42,481 784 2.93 124,533 License/endorsement renewal 2010 32,237 695 4.08 131,529 Insurance (personal vehicle) 2011 28,942 701 3.52 101,780 Insurance (boat) 2012 33,228 625 3.85 127,839 Insurance (facility) International shipping fees

such as temperature-related bleaching price increases, so they often turn to price and sales associated with the ML events and algal blooms, can weaken less expensive sources or alternative license). and cause octocoral mortality, making species from international markets. it difficult to find viable specimens to For instance, a collector interviewed Challenges in the sell. Even healthy colonies can quick- from the Florida Keys who was mining Octocoral Fishery ly deteriorate and become unviable if natural live rock in the 1980s stated The biggest challenge faced by oc- they are not handled or stored prop- that once the State of Florida prohib- tocoral collectors is the relatively low erly. In addition, maintaining octocoral ited live rock collecting in State waters demand for colonies by aquarium hob- colony health in an aquarium system is in 1992 (FAC 68B-42.008), marine byists, as well as finding and retaining challenging because octocorals require aquarium hobbyists shifted their at- customers. Maintaining live octocorals adequate spacing among colonies, wa- tention to Haiti to acquire inexpensive in aquaria is also a difficult and rela- ter quality, appropriate lighting (e.g., Caribbean live rock. During economic tively costly hobby, compared to main- lower light output for deeper water downturns, such as the recession of taining only fishes. In addition, there octocorals), and relatively strong wa- 2008, the price per octocoral colony are substantial physical challenges ter flow. Deepwater species also re- declined (Table 4) because of less dis- inherent when diving and collecting quire a supplemental food source since posable income to spend on luxury marine life. Long days spent on the they lack symbiotic microalgae (zoo­ aquarium items, which, in turn, placed water mean exposure to the elements, xanthellae). Aquarium system con- pressure on collectors’ daily mainte- venomous marine life such as lionfish, struction and maintenance is also ex- nance and operation costs (Table 3). Pterois spp., and jellies, and physi- pensive. Some collectors who do not Coupled with the relatively low de- cal fatigue. Marine life collection can live near a clean source of saltwater mand for octocorals (historically and also require diving expertise, espe- have to pay for saltwater transported to presently), collectors typically rely on cially when diving in strong currents their warehouses. other marine life species to increase and deeper water. Palm Beach Coun- Economically, the octocoral indus­ profits. During the past 20 years in ty is notorious for its high-current re- try supplies the larger aquarium trade, Florida, snails, crabs, starfish, , gime due to the proximity of the Gulf which is itself a luxury hobby, espe- and anemones constituted the most Stream to the coast, so collectors pre- cially as related to keeping live corals popular aquarium invertebrates sold to fer to drift dive with their boat. Many and octocorals. Price fluctuations and the aquarium trade (Table 5). collectors operate alone while diving octocoral supply problems are largely Future of the Fishery because they feel safer not having to uncontrollable by collectors. For ex- keep track of a dive buddy. Limited ample, when healthy octocoral colo- The state of the octocoral fishery market potential and difficulty col- nies become scarce due to bleaching, has remained relatively stable since lecting combine to make octocoral algal blooms, or other factors related 1994 (FWC, 2014). In the last six collecting a niche business, usually a to environmental stress, collectors years, however, there has been a de- small part of a larger enterprise that must travel further distances to find vi- cline in demand of octocorals by involves collecting a diversity of ma- able colonies, which, in turn, raises the aquarium hobbyists (Table 4). The ma- rine organisms. price of octocoral colonies. Hobbyists rine aquarium hobby is susceptible to Adverse environmental conditions, are usually unaware of the causes of trends, much like the fashion industry

80(3) 25 Table 5.—Annual landings and economic value of marine invertebrate species groups collected in Florida, 1994– ity. Demand then increased when they 2012, based on commercial trip ticket data (FWC, 2014). were re-introduced due to demand for Annual landings Value ($) Change in landings, Change in value apparently sustainable and ecologically Invertebrate group 1994 2012 1994–2012 (%) 1994 2012 1994–2012 (%) friendly products. Working against the Snails 290,211 3,308,614 1,040 173,812 449,102 158 trade is the fact that the most colorful Crabs 117,889 2,667,574 2,163 100,617 543,191 440 Sand Dollars 643,105 1,484,091 131 64,631 143,110 121 species are typically the most difficult Shrimp 33,719 1,754,445 5,103 32,374 353,760 993 Starfish 314,071 25,414 -92 298,922 40,705 -86 to collect, as they are usually the deep- Anemones 307,891 65,626 -79 164,671 136,621 -17 er water species found in high-current Octocorals 32,106 33,228 3 121,983 127,839 5 regimes. Such fauna are also more dif- ficult to maintain in good condition in hobbyist aquariums. In the mid-1990s, changing styles from year-to-year. One (Ho, 2011). Since octocorals are not for example, “red” octocorals were of the most popular current trends is considered rare or exotic (at least in marketed at a higher price point be- related to aquarium lighting. Advances the U.S.), they are not currently part of cause they were clearly attractive as in lighting technology have replaced this trend. ornamentals, but demand could not be large, heat-producing, energy-draining What would need to occur for the sustained. lights with more compact, energy-effi- octocoral fishery to boom in Florida The octocoral collectors interviewed cient LED lights that provide focused or the greater southeastern U.S. and have a positive outlook on the future of light into reef tanks (Joshi, 2010). Ac- Gulf of Mexico? The only real boom the octocoral fishery, yet are realistic tinic lights have also become popular in octocoral collection—and it was when discussing the future. Collectors in reef aquariums because they high- small—occurred in the mid-1990s af- have experienced trends that fluctuate light the fluorescence within coral pol- ter culture techniques were developed with the vagaries of the aquarium trade, yps. As a result, saltwater aquariums that were relatively easy for home yet they have adhered to collecting and are no longer dominated by just fish- aquarium hobbyists to adopt. During selling octocorals typically because es, but by other colorful species such this period, octocorals became associ- there are reliable buyers who provide as corallimorpharians and zoanthids. ated with keeping seahorses in aquari- a relatively small but steady income. A Since most octocorals apparently do ums, which elevated octocoral demand few of the collectors interviewed spoke not fluoresce under such lighting con- slightly. On a limited scale, the encrust- optimistically about the potential for ditions, they are not currently part of ing octocoral Erythropodium caribaeo- growth as hobbyists begin to learn more this trend. rum was once collected for a unique about octocoral husbandry require- Aquarium hobbyists also seek short- secondary metabolite that had pharma- ments through aquarium hobby online cuts in reducing the time it takes to ceutical properties. The biocompound, forums. Technological developments in perform maintenance duties on home however, was synthesized quickly by a power heads that simulate wave-driven aquaria, such as siphoning out excess Canadian company, subsequently re- water movement may also facilitate oc- food and detritus or removing . sulting in reduced demand for this spe- tocoral husbandry. Collectors are ad- As a result, “cleaner” invertebrates cies (Taglialatela-Scafati et al., 2002). ept at the use of social media outlets to such as snails and crabs are now com- The only other actively collected octo- market their marine organisms. Formal, mon elements in aquariums. Because coral for the pharmaceutical industry potentially costly, marketing approach- marine hobbyists often spend a lot of presently, at least that we are aware es have not apparently been attempted time, effort, and money, they enjoy of, is Antillogorgia elisabethae, which based upon the responses from the col- sharing and showing off their aquaria. is collected in the Bahamas and per- lectors interviewed. Some collectors Several marine aquarium forums have haps elsewhere (Castanaro and Lask- lamented the difficulty in finding reli- “tank of the month” pages (e.g., www. er, 2003). Typically, when a bioactive able customers who keep octocorals in reefcentral.com, Reefkeeping Maga- compound is isolated from a marine their aquaria, noting flat sales over the zine, and Ultimate Reef) that encour- species with potential for commercial years, compared with sales of other in- age hobbyists to send in photographs application, considerable effort is ex- vertebrate organisms such as crabs and of their home aquariums for others pended to synthesize the compound in corallimorpharians. Many collectors to learn from or to reserve “bragging the laboratory, since collection from interviewed mentioned that they would rights” for a job well done. Part of this natural populations rarely provides a welcome a concerted effort on the part social dynamic is seen in the variety of reliable or sustainable source of the of the scientific community to validate species within these tanks. Exotic and bioactive material. It is possible that the sustainability of octocoral collec- rare species such as the peppermint an octocoral species could suddenly tion, which they think might help pro- angelfish, Centropyge boylei, a deep- become fashionable and result in in- mote interest and boost sales. Indeed, water angelfish known to only inhabit creased demand. For example, Eco- that is one reason why so many collec- the Cook Islands, are highly sought Gorgs® were first introduced in the late tors were willing to be interviewed for after, despite the $20,000 retail value 1990s, and then declined in popular- this study.

26 Marine Fisheries Review Figure 6.—(A) Annual landings and (B) dockside value of invertebrates for the marine aquarium trade in Florida, 1994–2012 (FWC, 2014).

Commercial Marine al., 2001). During this same period, Sand dollars are collected for the cu- Life Landings and Values octocorals represented only 4.4% of rio trade and experienced a 57% in- all invertebrates collected (Larkin et crease in collection levels since 1994 Invertebrates al., 2001). Instead, gastropod snails, (FWC, 2014). As demand increases A principal driver of the marine life crabs, sand dollars, shrimps, starfish, for such organisms, so too does their fishery in the U.S. South Atlantic and and anemones represented the top in- dockside value. The dockside value of Gulf of Mexico is that it is a demand- vertebrate groups collected (Fig. 6A). ornamental cleaner shrimp, for exam- based fishery for aquarium hobbyists Since the State of Florida FWC began ple, increased by 52% on an average and for the curio trade. Historically, keeping public records of commercial annualized basis from $32,374 in 1994 the marine aquarium trade has been landings in 1994, declines in landings to $353,760 in 2012. Crab, snail, and fish-centric; however, developing tech-­ of starfish and anemones have given sand dollar values exhibited similar in- nologies in lighting and filtration have rise to dramatic increases in the col- creases in value (Fig. 6B, Table 5). advanced the husbandry of inverte- lection of shrimps, crabs, and snails brate species, thereby opening the on an annualized basis (Fig. 6A, Table Octocorals hobby to greater demand for other reef 5). The increased demand for organ- Octocorals are valued because at organisms (Rhyne and Tlusty, 2012). isms that graze algae and scavenge least some specimens provide aesthet- For example, during 1990-98, inver- detritus reflects demand by hobbyists ic beauty to home aquaria. However, tebrate landings accounted for an av- for organisms that provide an aquari- because most octocorals lack the color erage of 63% of the total value of um service (i.e., control and removal vibrancy of other invertebrates, they annual dockside landings within the of algal growth and detritus) for their have not experienced the same col- Florida marine life fishery (Adams et home aquaria (Rhyne et al., 2009). lection intensity as other marine life

80(3) 27 Figure 7.—Average annual landings (bars) and dockside value (line) of octocorals collected in Florida, 1994–2012 (FWC, 2014).

organisms. Since 1990, the FWC has the life history traits and husbandry ter the 2008 recession was likely the collected annual landings data that in- requirements for several octocoral spe- most important factor that drove pric- cludes the number of octocoral colo- cies, providing aquarium hobbyists es upwards, but low demand by hob- nies landed; however, landings are with a user’s manual for maintaining byists reduced the number of colonies not partitioned by taxonomic classifi- octocorals. By 2006, an article de- collected. The most recent data show a cation but instead by color categories scribing octocoral propagation tech- 13% increase in colony landings and (FAC 68B-42.001). According to the niques was published in Reefkeeping a $0.33 increase in price per colony FAC (Rule 68B-42), the term “octo- (Hiller, 2006), another popular online from 2011 to 2012 (Fig. 7, Table 4). coral,” which also includes “soft cor- marine aquarium magazine, making As reported in the FWC Trip Tick- al,” means “any erect, non-encrusting these techniques more widely acces- et reports (the following regional and species of the subclass Octocorallia, sible to the home hobbyist. Also dur- species group specific data are un- except the species ing this time, the use of octocorals in published), octocoral landings and and .” FWC also seahorse aquariums became popu- dockside value data are available for collects commercial octocoral fishery larized as images and discussions of Florida on a statewide basis (Fig. 8) data pertaining to the number of col- “seahorse-friendly” organisms became and further partitioned for the Florida lection trips made, average price per trending topics in marine aquarium east (Fig. 9) and west coasts (Fig. 10). colony, and average dockside value of forums (www.reefcentral.com/forums, Approximately 80% of octocorals are octocorals using trip ticket reports. As access­ed in 2006). The collectors in- collected from the Atlantic Ocean off reflected in nearly 20 years of avail- terviewed noted that requests for yel- the Florida Keys and the east coast of able commercial trip ticket data, oc- low octocorals increased during this Florida from Monroe County to Palm tocoral annual landings increased by same period, perhaps due to increased Beach County. In Florida Keys north- only 3%, while the price per colony online forum traffic. ward to Palm Beach County, com- has nearly doubled, from $2.23 in the Since the mid-2000s, the demand for pared to other areas of Florida and late 1990s to $4.08 in 2010 (Fig. 7, Ta- octocorals has declined annually, but the rest of the southeastern U.S., oc- ble 4). Noteworthy is that the price per the high price-per-unit has produced tocoral populations are diverse—spe- specimen is considerably higher than some of the highest grossing years cies from all of the color groups (e.g., for many other invertebrate groups, since 1994 for octocoral collection, as other, purple, and red) are collected which generally sell for less than reported in FWC Trip Ticket data. The (Fig. 9A). The smaller octocoral fish- $1.00 per specimen. In 2006, the oc- estimated value of the octocoral fish- ery on the west coast of Florida in tocoral fishery experienced a surge in ery peaked at $145,759 in 2007 and Gulf of Mexico waters, concentrated hobbyist demand, resulting in 48,567 $152,029 in 2008 (Fig. 7, Table 4). At mainly in Hillsborough and Pinellas colonies collected during 1,004 com- the height of the recession in 2009, the Counties, is comprised mainly of oc- mercial trips (Fig. 7, Table 4). Julian price per colony declined to $0.67 and tocorals in the “other” and “purple” Sprung (2004), a marine aquarium then increased from $1.15 to $4.08 categories. The taxa collected within expert and author of several books in 2010. Demand dropped by 24% in the “red” species group are Leptogor- on the topic, wrote an article in Ad- 2010 (Fig. 7, Table 4). The need for gia species and Muricea pendula (Fig. vanced Aquarist Magazine detailing collectors to generate lost income af- 10A and Table 2). The “red” species

28 Marine Fisheries Review Figure 8.—(A) Annual octocoral landings and (B) dockside value of octocoral species groups (other, purple, and red) in State and Federal waters off Florida, 1994–2012 (FWC, 2014). code elsewhere usually refers to Swif- data show the greatest dockside value diversify and find new income sourc- tia exserta, which has red polyps, but of octocorals ($48,128) compared to es, “Eco-Gorgs®” was developed. To it is possible that other octocorals with the value of “other” octocorals from create Eco-Gorgs®, several small (< a red rind or colony axis are report- the Gulf of Mexico ($25,955) (Fig. 9B 20 cm) clippings were removed from ed under this code. Contrary to what and 10B). larger adult colonies (Fig. 4), glued would be expected from an industry Demand for “purple” octocorals be- to concrete plugs (Fig. 5), and mar- that values colorful, seemingly rare gan to increase in the early 2000s keted as a sustainable alternative to species, the demand in the aquarium (Fig. 8), a period in which small collecting whole colonies. Additional trade is primarily for octocorals in the (< 20 cm) whole colonies of Antil- desirable species (color and/or large “other” species group (Table 6). On logorgia species, Briareum asbesti- polyps) such as Muriceopsis flavida average, nearly half (47%) of all of the num, and Pterogorgia anceps were and Plexaurella nutans were then in- octocorals collected annually are spe- being sold to hobbyists as an alterna- troduced to the market. Octocoral clip- cies within the “other” species group tive to the mostly brown octocorals pings had previously been introduced (18,340 octocorals), primarily (74%) sold within the “other” species group. to the market in 1998, and while they on the Atlantic Ocean of Florida However, the diversity of “purple” were valued as having a relatively low (Table 6). However, on average they octocorals available was limited be- impact on the ecosystem, they did not are sold at a lower price per unit than cause of the difficulty in finding small, introduce any additional selection to those collected on the Florida west whole colonies suitable for home the market. Increased interest in eco- coast ($2.67/colony vs. $3.75/colony) aquaria. After the 2008 economic re- friendly approaches and the introduc- (Table 6). The difference in unit col- cession, collectors began exploring tion of new species to the market were ony price has not hampered the value alternative options for increasing the what made Eco-Gorgs® more suc- of “other” octocoral species collected diversity of octocorals that could be cessful the second time around. The in Atlantic waters; current (as of 2012) brought to market. From this need to dockside market value of “purple”

80(3) 29 Figure 9.—(A) Annual octocoral landings and (B) dockside value of octocoral species groups (other, purple, and red) in State and Federal waters in the Atlantic Ocean off Florida, 1994–2012 (FWC, 2014).

Table 6.—Average annual landings (no. colonies), numbers of trips, price per colony, and economic dockside Initial attempts in 1994 to make value of octocorals collected in Florida in the Atlantic Ocean and Gulf of Mexico during 1994–2012, based on un- published commercial trip ticket data (FWC, 2014). a large profit margin off the color- ful “red” octocoral varieties (Table 2) Area Species group Landings No. of trips Price/colony ($) Value ($) by selling them at over $6.50/colony Atlantic Ocean Other 13,569 351 2.67 36,018 Purple 11,638 329 2.66 30,046 failed. The following year, “red” oc- Red 6,734 104 3.42 23,083 tocorals were sold at $2.48/colony

Gulf of Mexico Other 4,771 190 3.75 17,897 and have since sold for an average Purple 2,317 92 2.74 6,322 Red 454 14 2.63 763 of $3.14/colony. Though these octo- corals fit the desired color profile of

an ornamental organism, they are typi- octocorals peaked in 2008 at $57,171 “purple” octocoral taxa in the Atlantic cally found at depths ranging from 25 (Fig. 8B) and in 2009, demand for supports 84% of the overall collection to 60 m and may lack zooxanthellate. “purple” octocorals reached a high of of these species (Table 6). Purple oc- Without zooxanthellae, deepwater oc- 19,228 colonies (or fragments) collect- tocoral dockside values in the Atlantic tocorals rely solely upon tentacle cap- ed (Fig. 8A). surpassed that of the “other” octocoral ture of for their nutrition. Within the “purple” species group, species group from 2002–09 (Fig. 9B), Keeping deepwater octocorals in home eight species or taxa of octocorals are while landings were greatest in 2009 aquaria therefore means hand-feeding collected in Atlantic Ocean waters, three (Fig. 9A). As of 2012, the average an- the colonies live or frozen food mix- of which are also collected in the Gulf nual price per colony of “purple” octo- tures. Presumably because they are of Mexico: Antillogorgia species (sea corals was higher than any other year more difficult to care for, demand for plumes) and the sea whips Pterogorgia ($4.30/colony) for this and species “red” octocorals declined in Florida by anceps and P. guadalupensis (Table 2). collected within the “other” octocoral 75% since 2006 (Fig. 8A). Collection The greater diversity and abundance of group complex. efforts have also declined from 272

30 Marine Fisheries Review Figure 10.—(A) Annual octocoral landings and (B) dockside value of octocoral species groups (other, purple, and red) in State and Federal waters in the Gulf of Mexico off Florida, 1994–2012 (FWC, 2014)

trips annually in 2001 to 111 trips in Biology and Ecology are presently designated into one of 2012 (Sheridan7), with only a handful three color categories (purple, red, and of collectors willing to work at deeper Overview other), although octocoral collectors diving depths and with strong currents. A concern raised with the fisheries often distinguish genera and species. Those that do collect deeper water col- for marine ornamental, aquarium, and FWC biologists have proposed a new onies focus their efforts on the east reporting system based upon mor- curio species in the U.S. South Atlan- 8 coast of Florida off of Palm Beach, tic and elsewhere is the lumping of phology (Brinkhuis ). At issue is that Broward, and Miami-Dade Counties. species with potentially different bio- species with life history patterns that On average, 104 collecting trips are geographic and habitat distributions, render them perhaps more resilient to made annually, yielding 6,733 “red” as well as life history parameters (e.g., collection (e.g., rapid growth, high re- octocorals with a dockside value of growth and reproduction) in broad cat- cruitment, and broad distribution) are $23,082 (Table 6). In comparison, on egories (Rhyne et al., 2009). The octo- lumped with species that may be less average, 454 “red” octocorals (primar- coral fishery in the U.S. South Atlantic resilient to over-collection because of ily Leptogorgia species, Table 2) were and Gulf of Mexico, where the ma- slower growth, delayed reproduction, collected in the Gulf of Mexico during jority of the octocoral collection oc- lower recruitment, slower regenera- 10 out of the last 19 years, yielding an curs in Florida waters, is no exception. tion from injury, and/or restricted geo- average annual dockside value of $763 Octocorals collected in Florida waters graphic distribution or habitat/depth (Table 6). of the U.S. South Atlantic and Gulf distribution. One goal of this study of Mexico span genera and families was to address the poor taxonomic that are categorized into color variet- 7Sheridan, N. 2014. Octocoral harvest from state vs. federal waters by Gulf of Mexico vs. Atlantic ies for reporting under the FWC Trip 8Brinkhuis, Vanessa. Florida Fish and Wildlife waters, 1994-2013. FWC unpubl. data, personal Ticket program. Quotas and landings Conservation Commission, Fish and Wildlife commun. Res. Inst., Personal commun., Oct., 2013.

80(3) 31 resolution in the octocoral fishery to of mortality) for the main suite of oc- skeletal elements consisting of calcar- evaluate if sub-components of the fish- tocorals collected for the marine life eous spicules or sclerites (Lewis and ery might be more or less sustainable, aquarium and ornamental trades in the Von Wallis, 1991). In most octocorals, and thus perhaps require different U.S. South Atlantic and Gulf of Mex- the sclerites are arranged in three dis- management strategies. By address- ico. Much of the available literature tinct layers in the coenochyme, gener- ing these data gaps and providing the stems from octocoral studies in Flori- ally small (0.05–1 mm), and produced best scientific information to the South da, as well as the wider Caribbean or by specialized cells (Lewis and Von Atlantic Fishery Management Coun- tropical western Atlantic. Primary and Wallis, 1991). The outer cortex or sur- cil and the State of Florida, improved gray literature sources were reviewed, face layer of the octocoral colony can management of the octocoral fishery along with field survey information, to be variable in sclerite composition, but is a hopeful outcome, with ideally the compile species profiles (Appendix). generally, only one type of sclerite oc- least economic impact to the collectors Cairns et al. (2002) list 102 octocoral curs (Bayer, 1961). Octocoral sclerites, that depend financially upon octocoral taxa for U.S. Atlantic waters out to along with field characters such as resources. the edge of the continental shelf, not branching patterns, are often needed The narrative below provides an including Puerto Rico and the U.S. to confirm field identification (Bayer, overview of octocoral biology and Virgin Islands. Octocoral collectors 1961). ecology, with a focus on those species in Florida target at least 23 species, Octocorals are modular organisms, targeted in the octocoral fishery in the not including species groups such as similar to reef-building corals (Scler- U.S. South Atlantic and Gulf of Mexi- Antillogorgia spp., Eunicea spp., and actinia), where the basic module is the co. The number of species investigated Muricea spp. Information for 51 oc- polyp; growth is indeterminate, with was determined by consultation with tocoral species among 16 genera was no indication of senescence (Jackson, collectors, as they are only required to part of this study, including all of the 1991). Thus, portions of a colony may note color categories on Trip Tickets, targeted species in the octocoral fish- die due to injury, overgrowth by oth- and available landings data from the ery, as well as many sister taxa, espe- er organisms, or (Jackson, State of Florida do not show any fur- cially where collectors noted genera 1991; Castanaro and Lasker, 2003). ther taxonomic resolution. Dr. H. Fed- instead of species. Octocorals are benthic, sessile, passive dern, an octocoral collector, indicated suspension feeders, but many species, that he personally collects at least ten General Aspects of Biology especially in shallower water, derive genera, with seven taxa identified to Octocorals include the soft corals, nutrients from intracellular algae (zoo- the species level from samples con- sea fans, blue corals, and sea pens and xanthellae) and thus can be classified firmed by State of Florida biologists belong to the Phylum (Coel- as zooxanthellate or azooxanthellate. using sclerite samples (Table 1). Fur- enterata), an ancient (pre-Cambrian) Azooxanthellate corals rely solely on ther consultation and interviews with and diverse assemblage of mostly ma- particle capture by the tentacles to ful- other collectors indicated a rather rine organisms also including jellies, fill their nutritional requirements. broad range of families, genera, and sea anemones, corals, and hydroids Classification of octocorals, like for species that are targeted, represent- (Fabricius and Alderslade, 2001; many groups of organisms, has under- ed by 15 genera and at least 23 spe- Cairns et al., 2002; Brusca and Brus- gone significant changes in the past cies (Table 2). As most of these taxa ca, 2003). Diversity within the phylum decades, with various reclassifications are collected primarily in the Florida stems from two fundamental aspects and renaming (Bayer, 1961, 1981; Fa- Keys, the focus of the overview of oc- of many cnidarian life cycles: 1) the bricius and Alderslade, 2001; Cairns tocoral biology and ecology concerns tendency to form colonies by asexual et al., 2002). Besides scientific names, those genera and species identified reproduction and 2) dimorphic alterna- various common names are used by by collectors, along with sister taxa tion of generations as polyps and me- octocoral collectors, recreational div- within genera in cases where only a dusa exhibited by some groups. Like ers, and scientists (Voss et al., 1969; (e.g., Eunicea spp. or Muricea other cnidarians, octocorals are dip- Cairns, 1977; Cairns et al., 2002). spp.) was identified as being targeted. loblastic organisms (epidermis, gas- Within the phylum, octocorals are A substantial compilation of species- trodermis, and mesoglea) at a tissue grouped under the class that specific information is presented in grade of construction, characterized contains nine orders and ~6,000 extant the Appendix9, which synthesizes by radial symmetry, tentacles, stinging species. Within the Class Anthozoa all much of the available information on structures (cnidae with nematocysts), of the octocorals are in the Subclass geographic distribution, depth range, an incomplete gastrovascular cavity Octocorallia (=Alyconaria). They are habitat, and life history information with only one opening (the mouth), characterized as colonial organisms (e.g., growth, recruitment, and sources and a middle layer of mesenchyme or containing polyps with eight hollow, mesoglea (Brusca and Brusca, 2003). marginal, and usually pinnate (feath- 9The appendix is available in the online ver- sion of this article (doi: https://doi.org/10.7755/ The mesoglea in octocorals, called the ery) tentacles, along with eight com- MFR.80.3.2). coenochyme, can contain fibers and plete mesenteries, with free or fused

32 Marine Fisheries Review sclerites embedded in mesenchyme and water flow regimes (Bayer 1961; cur below 50 m depth and/or are found (Brusca and Brusca, 2003). The pin- Lewis and Von Wallis 1991). Octo- in temperate waters (Bayer, 1961; nate or feathery tentacles are covered corals are one of the more difficult Cairns et al., 2002). Taxa with deep- with sensory cells and stinging cap- groups of organisms to identify, and er depth distributions, beyond where sules (cnidae), and may contain sym- may explain why the Federal and State most octocoral surveys are conducted, biotic algae (zooxanthellae). Stolons of Florida reporting requirements cur- often have sparse or no biological in- or coenochyme connect the polyps, rently consist of color categories. formation (see Appendix). with new polyps usually formed by budding from existing stolons. Octo- Biogeography Habitat and Depth Distribution corals comprise a diverse assemblage Octocorals are broadly distributed In subtidal habitats, specifically in of organisms commonly referred to as in the marine environment, present areas with available hard substratum, soft corals, sea fans, gorgonians, blue in every ocean basin from the tropics octocorals can reach densities ap- corals, and sea pens. In the literature to polar regions, from the intertidal proaching 100 colonies/m2 (Yoshio- and in the vernacular, the term “soft zone to abyssal depths (Bayer, 1961; ka, 1998). Environmental parameters coral” sometimes refers to Octocoral- Fabricius and Alderslade, 2001). The such as light, water depth, water mo- lia, which have no massive skeleton or western central Pacific harbors the tion, sedimentation, and availability internal axis, but sometimes this term greatest number of octocoral species of appropriate substrata are the most also includes sea fans (Fabricius and (90 genera and 23 families), but the important determinants of octocoral Alderslade, 2001). The term “sea fan” western Atlantic also include a rela- distribution. For zooxanthellate spe- is sometimes used as an alternative tively high number of species (Bayer, cies, the light regime, influenced by for gorgonian, which are Octocorallia 1961; Cairns et al., 2002). The Amer- depth and water clarity, but also sub- other than sea pens that grow upwards ican Fisheries Society (AFS) Com- stratum steepness and stability, are im- from the substratum with the support mittee on Common Names of Aquatic portant factors in spatial distribution. of an internal axis (Fabricius and Al- Invertebrates (CNAI) lists 102 oc- In azooxanthellate octocorals, larvae derslade, 2001). tocoral taxa for U.S. Atlantic waters preferentially settle in low light micro- Soft corals and gorgonians were (exclusive of Puerto Rico and the environments and depend on particle formerly classified separately in the U.S. Virgin Islands) that inhabit con- capture to meet nutritional require- Orders and Gorgonacea, tiguous shore waters on or above the ments. Octocorals can flourish in areas respectively; however, they are now continental shelf to a depth of 200 with strong currents and light to mod- grouped together under the Order Al- m. The focus herein is on geographic erate wave action. Many species re- cyonacea (Bayer, 1981; Fabricius and distributions pertaining to those gen- quire consistent and moderately strong Alderslade, 2001; Cairns et al., 2002). era and species targeted by the octo- currents to maximize food encounter Most of the octocorals collected for coral fishery from North Carolina to rates (Fabricius and Alderslade, 2001). the marine aquarium and ornamental Texas, with a particular emphasis on This is most evident, for example, in (curio) trades in the U.S. South Atlan- south Florida. sea fans, where colony orientation is tic and Gulf of Mexico consist of sea Broad-scale (i.e., wider Caribbean) perpendicular to the predominant cur- whips, sea rods, and sea plumes, dis- geographic patterns of 51 octocoral rent direction to maximize food parti- tributed among 4 suborders and 12 species are listed in Table 7. The ma- cle interception. Stronger currents not families (Cairns et al., 2002). Colo- jority of shallow-water (< 20 m depth) only transport food towards and waste nies are usually brightly colored and species have a relatively broad distri- away from colonies, but also stimulate arborescent with a firm internal axial bution across the wider Caribbean, photosynthesis. Tolerance to sediment skeleton composed of horny protein- largely mirroring the distribution pat- loads varies widely among octocoral aceous material (gorgonin) that con- terns of stony corals, concentrated in taxa, with some adapted to clearer wa- tains modified collagen and may be warmwater (18–31ºC) regions, pos- ters, while others flourish in moderate- mineralized. Occasionally the skeleton sibly due to the relatively narrow ly turbid waters. Some azooxanthellate is calcareous, but colonies are always temperature tolerance of microalgae species are quite abundant in turbid covered with a thin layer of sclerite- (zooxanthellae) (Fabricius and Alder- environments (Opresko, 1973; Fabri- filled mesoglea, which provides colony slade, 2001). Approximately 170 spe- and Alderslade, 2001). support and protection (Bayer, 1961; cies of gorgonian octocorals occur in Lewis and Von Wallis, 1991). Surface the West Indian Province (wider Ca- Life History sclerite (spicule) shape is used to iden- ribbean), which includes Bermuda, Life history refers to an organism’s tify octocorals (Bayer, 1961), along the Bahamas, the Greater and Lesser lifetime pattern of growth, develop- with patterns in the colony growth axis Antilles, south Florida, the Gulf of ment, and reproduction, reflected in (Cairns, 1977). Octocoral growth mor- Mexico, and the Caribbean coasts of an array of adaptations relating to phology and even the structure of the Central and South America (Cairns, physiology, morphology, and behav- spicules can vary over different depth 1977). Many of these taxa (~100) oc- ior. Life history includes traits that

80(3) 33 Table 7.—Biogeography of octocorals (Cnidaria, Anthozoa, Alcyonaria = Octocorallia) in the Wider Caribbean, with a focus on taxa occurring on the Atlantic and Gulf coasts of Florida, especially those collected for the marine aquarium/curio trade (shaded). Asterisks (*) indicate the most demanded taxa in the U.S. South Atlantic octocoral fish- ery. Species are grouped by suborders and families according to Cairns et al. (2002).

Wider Caribbean region1 Octocoral species 1 2 3 4 5 6 7 8 Suborder Family Briareidae Briareum asbestinum X X X X X X

Family Diodogorgia nodulifera* X X X X X X Erythropodium caribaeorum X X X X X X X Iciliogorgia schrammi X X X X X

Suborder Family Eunicea asperula X X X X E. calyculata X X X X X X X Eunicea (=Plexaura) flexuosa* X X X X X X X E. fusca X X X X X X E. knighti X X X X X X E. laciniata X X X X X X E. mammosa X X X X E. palmeri X X X E. succinea X X X X X E. tourneforti X X X X X X X Muricea atlantica X X X X X X X M. elongata X X X X X X X M. laxa X X X X X X X M. muricata X X X X X M. pendula M. pinnata Muriceopsis flavida* X X X X X M. petila X X X Plexaura kuna (=Plexaura A) X P. homomalla X X X X X X Plexaurella dichotoma X X X X X X P. fusifera X X X X X X P. grisea X X X X X P. nutans* X X X X X X X P. pumila X X X X X Pseudoplexaura crucis X X X X P. flagellosa X X X X X X P. porosa X X X X X X P. wagenaari X X X X X X Swiftia exserta* X X

Family Antillogorgia (=Pseudopterogorgia) acerosa* X X X X X X X X A. americana X X X X X X X A. bipinnata X X X X X X A. elisabethae* X X X X A. kallos* X X A. rigida X X X X X X X Gorgonia flabellum X X X X X G. ventalina X X X X X X Leptogorgia (=Lophogorgia) cardinalis X X X X L. punicea L. virgulata X X X Pterogorgia anceps* X X X X X P. citrina* X X X X X X P. guadalupensis X X X X X

Suborder Family Ellisellidae Ellisella barbadensis X X X X X E. elongata X X X X Ellisella (=Nicella) schmitti X X 1(1) Gulf of Mexico, including the west coast of Florida, (2) southeastern Florida including the Florida Keys, (3) Bahamian archipelago (Bahamas and Turks and Caicos), (4) Hispaniola, Puerto Rico, U.S. Virgin Islands, (5) Antillean (Lesser Antilles from the British Virgin Islands to Barbados and Grenada), (6) NW Caribbean (Cuba, Jamaica, Cayman Islands, Caribbean coast of Mexico, and Belize), (7) Continental Caribbean (Honduras to Suriname, including Trinidad, Tobago, Aruba, Bonaire, and Curacao), and (8) Bermuda.

affect an organism’s “schedule” of reproduction (semalparity vs. iter- adaptations relating to an organism’s reproduction and survival; how an oparity); type of larval output (inter- interactions with other organisms. organism reproduces (including re- nal brooding vs. external spawning Most marine populations, including productive mode); timing of repro- and development); and number of octocorals, are dynamic in both space duction in relation to environmental offspring produced per reproductive and time. Survival of adults is one of factors, age, or size; frequency of episode. Life history also includes the principal factors affecting popula-

34 Marine Fisheries Review tion changes. For example, if survival than injured colonies, thus sublethal maturity when colonies are over 20 of adults is high, then the current pop- injuries were concluded to be partly cm in height. Pterogorgia citrina, ulation size plays a significant role in responsible for intra-specific variation a small bushy octocoral that rarely explaining the population size in the in growth (Yoshioka and Yoshioka, grows above 30 cm in height, may future. As many octocorals brood lar- 1991). Thinly branched species (A. reach sexual maturity sooner, but little vae, often throughout the year, once an acerosa and A. americana) are rela- is known about this species. To pro- assemblage of reproductive colonies tively fast growing, while species with vide adequately sized octocorals to is established, it tends to persist. At thicker branches (e.g., P. dichotoma) home aquarists, collectors typically the same time, the generation of off- grow slower, presumably represent- take colonies that are less than 20 cm spring and survival of young are also ing a trade-off between colony branch in height. In doing so, collectors leave important. elongation and branch thickening (Yo- larger colonies behind that are the shioka and Yoshioka, 1991). major contributors of gametes to the Age and Growth population. For example, colonies of Growth rate data are limited for Reproduction and Recruitment Eunicea flexuosa over 70 cm in height most octocoral species (Cary, 1914; Like many marine invertebrates, oc- produce 93–98% of the eggs released Yoshioka and Yoshioka, 1991). Colony tocorals exhibit a two-stage (bi-partite) during spawning (Beiring and Lasker, growth is usually estimated by chang- life cycle, consisting of a benthic, ses- 2000). The octocoral collectors inter- es in colony height. Like other modu- sile stage (post-settlement juvenile viewed indicated that they rotate col- lar organisms, such as stony corals, to adulthood) and a planktonic lar- lection sites and are selective in which there is high variability in growth and val stage (planula). The oval-shaped, colonies they collect. size is often a poor indicator of colony featureless planula larvae settle out is common in age (Yoshioka and Yoshioka, 1991). In from the water column, with the spa- octocorals (Fabricius and Alderslade, general, faster-growing octocoral spe- tial pattern of settlement influenced 2001) and often the predominant mode cies tend to be found at higher densi- by substratum type, light intensity, of reproduction, but not in all spe- ties (Yoshioka, 1997). Previous studies and integrity (Wahle, 1985). Crustose cies (e.g., Leptogorgia virgulata, Go- suggest a general pattern of relatively coralline algae are preferred substrate telli, 1988). In asexual reproduction, low mortality and high variability in for octocoral recruitment (Benayahu et colonies may form runners or stolons octocoral growth rates (Yoshioka and al., 1989; Lasker and Kim, 1996). In from the base of the colony, or extend Yoshioka, 1991), with variability in contrast, the presence of algal films or branches out from the colony. The par- growth roughly equivalent to average sediment can impede recruitment and/ ent colony then translocates parts of growth increments. Thinly branched or lead to greater post-settlement mor- its own body mass through the stolon, taxa, such as Antillogorgia and Muri- tality (Gotelli, 1988). Upon settlement, and a new daughter colony forms that cea species, can grow up to 4 cm/yr, the planula larvae metamorphose into separates from the parent. The stolon while thicker branched species, such a founder polyp, initially forming a is then resorbed, resulting in two col- as Plexaurella, barely grow 1 cm/yr short stalk at one end that is attached onies. Octocorals also can bud, frag- (Yoshioka and Yoshioka, 1991). Birke- to the substratum. The other polyp end ment, and undergo colony fission, land (1974) recorded a mean growth flattens and eight tentacle buds devel- where constrictions are formed in the of 2.6 ± 1.9 cm (mean ± 95% CI) for op around the mouth opening (Fabri- colony and eventually divide into two the common sea fan, Gorgonia venta- cius and Alderslade, 2001). Within a independent smaller colonies. In bud- lina, over a 230-day period off coastal few days, a complete octocoral polyp ding, small buds are produced on the Panama. Birkeland (1974) recorded an develops; at this stage, some zooxan- edge or base of a colony, which even- annual growth rate of 2 cm/yr for the thellae species begin to take in plank- tually drop off and attach to the sub- same species in the Cayman Islands. tonic zooxanthellae through the mouth stratum as separate colonies. Some Yoshioka and Yoshioka (1991) stud- (the zooxanthellae are not digested, species are known to drop polyp bun- ied a multi-species octocoral assem- but are incorporated into the octo- dles consisting of 5–10 polyps that blage on two shallow-water reefs (< coral tissue), while in other species, sink to the substratum, or by “auton- 10 m) off the southwest coast of Puer- the are supplied with zooxanthel- omy of colony ends” (i.e., fragmenta- to Rico over a 4.5-yr period. Growth lae prior to their release into the water tion sensu Walker and Bull, 1983). rates, as determined by changes in col- column (Benayahu et al., 1992). Al- Octocorals sexually reproduce us- ony heights, ranged from 0.8–4.5 cm/ though a few deeper water octocoral ing a diversity of strategies parallel- yr among 12 taxa. The mean growth taxa remain permanently solitary as ing scleractinian reef-building corals rate was 1.4–2.6 cm/yr for most spe- single polyps, all other octocorals de- (Kahng et al., 2011). Most octocorals cies, ranging from a low 0.8 cm/yr in velop from a solitary founder polyp have male and female reproductive Plexaurella dichotoma to a high of 4.5 and form colonies derived from bud- structures in separate colonies, which cm/yr in Antillogorgia americana. Un- ding (Fabricius and Alderslade, 2001). is referred to as the gonochoristic injured colonies generally grew faster Octocorals typically reach sexual mode of reproduction (Fabricius and

80(3) 35 Alderslade, 2001). Three types of sex- ready or nearly so for metamorphosis. faster than injured colonies (Yoshioka ual reproduction have been identified A final reproductive strategy within and Yoshioka, 1991). in octocorals: broadcasting (broad- the octocorals is the external brooding Sources of octocoral mortality in- cast spawning) of eggs and sperm, of larvae. The fertilized eggs develop clude detachment from the substra- internal brooding of larvae, and ex- into larvae within mucus pouches on tum (Wahle, 1985), fracture of the ternal brooding of larvae (Benayahu the surfaces of the mother colonies skeleton, overgrowth (Wahle, 1980), and Loya, 1984; Benayahu, 1991; (Brazeau and Lasker, 1990). Brooded disease outbreaks (Naglelkerken et Lasker and Stewart, 1992). Broad- larvae, which are negatively buoyant, al., 1997), and thermal stress (Lasker, cast spawning has been observed in may settle within a few meters of the 2005). Octocorals are susceptible to alcyoniid soft corals and some gorgo- mother colony. abrasion and dislodgement by storm nians, which release large numbers of Existing information on octocoral waves and associated movement of eggs and sperm into the water where recruitment and post-settlement mor- sand and rubble (Wahle, 1985; Fabri- external fertilization occurs. Spawn- tality suggests that, in contrast to cius and Alderslade, 2001). Some Ca- ing is often synchronized according established adult colonies, post-set- ribbean octocorals (e.g., encrusting to lunar phase and/or water tempera- tlement mortality can be quite high. Briareum and Erythropodium), along ture. Subsequently, planula larvae de- For example, Gotelli (1988) observed with some branching species, especial- velop from fertilized eggs and remain that ~64% of Leptogorgia virgulata re- ly sea fans (Gorgonia spp.), are quite in the plankton for days to weeks un- cruits did not survive the first year after abundant in wave-exposed environ- til settlement and metamorphosis into settlement. With established popula- ments. However, many octocorals are founder polyps. Larvae may be car- tions, survival of additional post-set- more abundant in less wave-exposed ried a considerable distance—tens to tlement octocoral recruits is generally environments, including lagoon patch hundreds of km away from the pa- higher, while survival of established, reefs and deeper fore-reef habitats (Fa- rental colonies based upon oceano- larger colonies (10+ cm in height) may bricius and Alderslade, 2001). In shal- graphic conditions (Gotelli, 1988). be more or less constant (~94% per yr) lower water, sea fan colony orientation Some species (i.e., Plexaura kuna and (Yoshioka, 1998). While established is perpendicular to current direction Eunicea fusca) exhibit multiple strat- colonies may or may not be a key fac- (Wainwright and Dillon, 1969), pre- egies, reproducing sexually (brooding tor underlying octocoral population sumably an adaptation to high wave- or broadcast spawning) and asexually fluctuations, they are essential to the energy conditions and to maximize through fragmentation and vegeta- persistence of populations (Yoshioka, food particle interception. tive propagation (Kahng et al., 2011), 1998). Sediment burial and abrasion stress which can support high population are important sources of mortal- growth rates (Lasker, 1988). Mortality Rates, Sources ity for both established colonies and Like scleractinian corals, the de- of Mortality, and newly settled recruits (Gotelli, 1988), mography of octocorals, including Regeneration from Injury along with weakening of the substra- sexual maturity and reproductive out- tum (Yoshioka and Yoshioka, 1991). put, may be more based upon size Lacking major disturbances, such Colony detachment generally results (height) rather than age (Wahle, 1983; as storms and disease outbreaks, octo- from the failure of the underlying sub- Kapela and Lasker, 1999). Octocorals corals appear to exhibit low mortality strate rather than the colony holdfast have size-dependent reproduction abil- rates from the few studies available. (Cary, 1914; Kinzie, 1973; Birkeland, ity, although some (e.g., Leptogorgia Over a 230-day period off the coast 1974). The major source of mortality virgulata) may be reproductively com- of Panama, for example, Birkeland (40–71%) in one study (Yoshioka and petent within two years of settlement (1974) documented > 95% survivor- Yoshioka, 1991) was due to basal frac- (Gotelli, 1988), while other taxa may ship of sea fans (Gorgonia ventalina). ture (breakage of the axial skeleton not reach reproductive competence un- Probably the most comprehensive near the colony base (22–46%) where til much later post-settlement (Beiring study of survivorship rates in a multi- greater rates were attributed to higher and Lasker, 2000). species octocoral assemblage is the wave energy. Injury to the colony bas- Internal brooding of larvae is com- work of Yoshioka and Yoshioka (1991). al tissue caused by physical abrasion mon in many gonochoristic octocorals At two sites on the southwestern coast increases the probability that a colony (e.g., family Xeniidae), including some of Puerto Rico, they recorded an an- will eventually experience basal frac- taxa in the wider Caribbean (Brazeau nualized survivorship rate of 91.6– ture (Yoshioka and Yoshioka, 1991). and Lasker, 1990). Sperm are released 92.4%, with no significant differences Detachment and basal fracture are into the water (not the eggs), usually among the 12 octocoral taxa sampled. largely non-selective among octocoral a few hours after sunset, and the eggs Sublethal (i.e., not whole colony) inju- species and are not likely due to pre- that are fertilized develop into larvae ries were responsible for some of the dation (Harvell and Suchaneck, 1987; within the female colonies. Larvae are high intra-specific variation in growth; Lasker and Coffroth, 1983; Yoshioka released days to weeks later and are uninjured colonies generally grow and Yoshioka, 1991).

36 Marine Fisheries Review Ecological and Economic some species have a reverse pattern, and distribution lead to a better under- Roles of Octocorals possibly due to strong or slack current standing of the octocoral fishery and flow, high irradiance, and/or distur- potential management improvements Most octocorals are suspension bance by predators (Fabricius and Al- or alternatives? Next, is the current feeders, obtaining small food particles derslade, 2001). Preston and Preston octocoral fishery reporting structure from the water column. The octocoral (1975) found that the degree of polyp sufficient, or are there changes to diet consists of small (< 20 μm) partic- extension in an assemblage of mostly implement that would improve man- ulate organic matter such as bacteria, zooxanthellate octocorals was related agement or understanding of the fish- phytoplankton cells, and to plankton availability, but light inten- ery? Relatedly, is there a need for an (Fabricius and Klumpp, 1995; Fabri- sity was most important in daily activ- octocoral species identification guide cius et al., 1995; Ribes et al., 1998). ity patterns. to help standardize reporting require- Most of the food captured is assimi- Because branching octocorals grow ments in the U.S. South Atlantic? lated near the point of ingestion (Mur- upright from the substratum, they offer Finally, what fishery-independent as- doch, 1978). Larger particles that a canopy structure that provides habi- sessment and monitoring data are encounter tentacles are captured and tat for a diversity of invertebrates and available for octocorals in the region swallowed if suitable, typically con- fishes (Cairns, 1977; Wahle, 1980). that provide octocoral population- sisting of early crustacean and bivalve Patton (1972) documented level assessments? Are these efforts larval stages (Fabricius et al., 1995; symbionts on Leptogorgia virgulata adequate to assess the status of popu- Ribes et al., 1998). As octocoral nema- from the northern Gulf of Mexico, lations using current data from the oc- tocysts are relatively small compared while Spotte and Bubucis (1996) de- tocoral fishery? to jellies, hydroids, and sea anemones scribed the caridean shrimps associ- The marine aquarium trade has both (Fabricius and Alderslade, 2001), par- ated with Antillogorgia americana. commercial and recreational compo- ticle capture is restricted to smaller From the information compiled on nents, and similar to other fisheries, is food items. The octocoral epidermis is octocorals as represented in the Ap- complex in terms of the diversity and covered with microvilli, so absorption pendix, it is clear that many informa- number of invertebrate and fish spe- of dissolved organic matter is likely tion gaps exist, particularly growth, cies targeted, the different types of an additional food source (Schlichter, age, longevity, mode of reproduction, gear and approaches used to capture 1982). Octocoral feeding rates are af- and recruitment rates for deeper-water organisms, geographic areas, and ac- fected by food concentrations and cur- taxa. There are relatively few long- cess points (Bohnsack et al., 1994). rents (Fabricius et al., 1995). term studies of growth and survivor- There is a long history of marine life Azooxanthellate octocorals, because ship (Yoshioka and Yoshioka, 1991), collection that dates back to at least they do not harbor zooxanthellae, ob- and most studies of octocoral species the 1960s in the south Florida area, tain all of their nutrition by suspen- or assemblages have been short in du- where a diversity of invertebrate and sion feeding. Many shallow-water ration (< 1 yr) and usually limited to fishes were collected, primarily for the octocorals host symbiotic algae (zoo- one or a few species (Yoshioka and Yo- marine aquarium industry but also for xanthellae, Dinoflagellata) to supple- shioka, 1991). the curio trade (Voss et al., 1969; Jaap ment their dietary needs and are thus and Wheaton, 1975; Robins, 1976; light dependent (Kinzie, 1973). Zoo- Analysis of Current Derr, 1992; Bohnsack et al., 1994). xanthellae are embedded in gastro- Data Collection Programs Some of these activities are now man- dermal cells or in membrane-bound Current data collection programs aged, including the moratorium on sea vacuoles within the gastrovascular for octocoral management in the U.S. fan collection, restrictions on live rock cavity. Zooxanthellate octocorals uti- South Atlantic and Gulf of Mexico collection, and prohibitions on the use lize photosynthetic products from their were reviewed as part of this study, of chemicals for collecting (FAC Chap. microalgae to contribute to their ener- particularly fishery independent stud- 68B-42). In Florida waters, the collec- gy needs, but still must rely upon par- ies in southeast Florida and the Florida tion of live tropical, ornamental, and ticle capture by the tentacles (Fabricius Keys, since most octocorals are landed marine species such as fish, inverte- and Klumpp, 1995). Octocorals appar- from these areas. The narrative below, brates, and plants is regulated by FAC ently have a lower photosynthetic effi- along with the management recom- Chap. 68B-42. Implementation of the ciency that may be related to the low mendations section, addresses the “Marine Life Rule” went into effect in surface area of octocoral colonies in following questions raised about the 1991 and has undergone subsequent relation to biomass (i.e., photosynthet- U.S. South Atlantic octocoral fishery. amendments. Bag limits, collection ic yields are low and respiration rates First, is the level of data collected at techniques, and area restrictions have are relatively high) (Fabricius and the State of Florida and Federal levels been set for recreational and commer- Alderslade, 2001). Most zooxanthel- sufficient to effectively manage the oc- cial marine life collectors in both Fed- late octocorals generally expand their tocoral fishery? Second, does existing eral and Florida waters, and included tentacles and feed during the day, yet information on octocoral abundance restricted collection of certain species

80(3) 37 and licensing and endorsement re- depth distributions (e.g., Diodogorgia, colony density, colony size structure, quirements. Octocorals are currently Ellisella, Iciliogorgia, and Swiftia). and population abundance estimates. considered to be ornamental species. These taxa are under-sampled or are For the southeastern Florida and Historical (Bohnsack et al., 1994) and not targeted in historical and current Florida Keys areas, the available in- current collection levels (FWC, 2014), assessment and monitoring programs. formation on octocoral population in terms of numbers of individuals or Despite these constraints, there is evi- parameters (distribution, density, size colonies removed, are lower than for dence that even some of the deeper structure, and abundance estimates) most other marine life fishery species water octocorals are broadly distrib- can be summarized as follows. First, and groups (Table 4). uted and occur in high densities along most surveys of octocorals were con- The historical and current reporting much of the southeastern coast of the ducted in relatively shallow-water structure for the Florida marine life U.S. (David10). There are obvious gaps (< 15 m) hard-bottom and coral reef fishery dating back to 1990 consists in knowledge related to the distribu- habitats, although a few studies sur- of landings by numbers of individu- tion and abundance of some octocoral veyed deeper hard-bottom and reef als or colonies, as opposed to the sizes genera and species. However, the rel- environments (e.g., Goldberg, 1973; and/or weights of organisms captured atively broad distribution of most oc- Wheaton, 1981, 1987). Second, most (Bohnsack et al., 1994; FWC, 2014). tocorals in southeastern Florida and historical studies conducted species in- Exceptions to this pattern are for ma- estimates of current population sizes ventories of octocorals in one or more rine plants (weight) and live rock presented below, relative to the small habitats, with information presented as (weight or number of boxes) when numbers of octocorals collected per species presence or absence (e.g., Voss this was permitted. Since 1990, octo- year for most taxa, suggests that 1) ad- and Voss, 1955; Voss et al., 1969; Da- coral landings, which are reported as ditional reporting requirements, while vis, 1982; Wheaton, 1987; reviewed in the number of colonies collected, are desirable, may not be necessary, and Levy et al., 1996). Third, relatively few grouped into broad categories based 2) the octocoral fishery in Florida is historical and more recent studies have on color (e.g., yellow, purple, or red), sustainable. The number of octocorals quantified octocoral colony density growth form (e.g., sea blades), or other collected in Florida appears to be a (numbers of colonies per unit area) by (Bohnsack et al., 1994; FWC, 2014). minor percentage (<0.1%) of the pop- habitat and depth (e.g., Goldberg, 1973; In Florida, landings were initially re- ulation abundance estimates, where Opresko, 1973; Kissling, 1977; Jaap, ported from the county where an or- they exist, for aggregate and individu- 1984; Wheaton and Jaap, 1988; Chi- ganism was first sold, not necessarily al taxa. appone and Sullivan, 1994), and these where an organism was collected. The studies have typically been of limited FWC currently collects octocoral data Fishery Independent duration (< 2 yr). A few sampling ef- in terms of numbers of colonies col- Data Sources forts provide estimates of octocoral lected, number of collection trips, av- Fishery-independent assessment and holdfast or canopy cover, but, except erage price per colony, and average monitoring available for octocorals in for encrusting species, they do not pro- dockside value of colonies. On both the U.S. South Atlantic and Gulf of vide information on octocoral densities the Florida Atlantic Ocean and Gulf of Mexico regions are described below, (Chiappone and Sullivan, 1997; Ruzic- Mexico coasts, information gathered with a focus on results that provide ka et al., 2009; Gilliam, 2012). Even from octocoral collectors indicates that population-level assessments for octo- fewer studies recorded octocoral size among these color groupings, there are corals. Results address whether or not (height) structure for branching octo- at least 29 octocoral species among 19 these data are adequate to assess the corals species (e.g., Opresko, 1973). A genera, with at least 12 nominal taxa status of octocoral populations in the small number of projects such as the (species and/or genera) characterized absence of finer-resolution fishery-de- Coral Reef Evaluation and Monitoring as in high demand (Tables 1 and 2). pendent data from the octocoral fish- Program (CREMP), Broward County’s There are aspects of octocoral bi- ery. As most octocorals are collected in Marine Biological Monitoring Pro- ology and ecology that are relatively southeastern Florida from Palm Beach gram, and local academic institutions, similar across taxa, such as the need County to Monroe County (Table 5), through the use of permanently marked for stable hard substrata for settlement the focus of this section provides an sampling locations, have monitored and growth. On the other hand, some overview of historical and current changes in octocoral canopy cover and/ species exhibit differences in habitat studies of octocorals in the area. Sam- or colony densities either annually or distribution and life history parameters pling efforts are briefly reviewed that intermittently, dating back in some cas- such as growth rate and reproduction. provide data on geographic distribu- es to the mid-1990s (Blair and Flynn,­ For many of the targeted taxa, there is tion, habitat and depth distribution, 1989; Moyer et al., 2003; Beaver et al., a paucity of available information on 2006; FWC, 2006; Thanner et al., 2006; habitat distribution, population abun- Gilliam et al., 2007). Finally, only one dance, and life history parameters, es- 10David, Andrew. NOAA Natl. Mar. Fish. Serv., program to our knowledge has used pecially for species that have deeper Panama City, FL. Personal commun., Nov. 2014. a survey sampling design, structured

38 Marine Fisheries Review by habitat, depth, geographic region, Dade to Palm Beach Counties, Banks 30 colonies/m2 (Gilliam et al., 2003; and management zone, to quantify et al. (2008) summarized several stud- Sathe et al., 2008). octocoral densities that can be scaled ies describing hard-bottom and reef Further south in the Florida Keys, to abundance estimates (Miller et al., community structure (Goldberg et al., several historical surveys provide oc- 2002; Smith et al., 2011). 1973; Blair and Flynn, 1989; Moyer et tocoral distribution and density data The majority of available informa- al., 2003; Beaver et al., 2006; FWC, from nearshore hard-bottom and tion on octocorals in Florida consists of 2006; Thanner et al., 2006; Gilliam et patch reef environments (Voss and species lists or inventories from various al., 2007). Historical surveys of octo- Voss, 1955; Opresko, 1973; Chiap- studies, dating back to the 1950s (e.g., corals north of the Florida Keys are pone and Sullivan, 1994). Chiappone Bayer, 1961), combined with informa- somewhat limited. Goldberg (1973) and Sullivan (1994) conducted pres- tion on habitat and depth distribution conducted benthic surveys of stony ence-absence and density surveys of (reviewed in Levy et al., 1996). These corals and octocorals on nearshore stony corals and octocorals at several surveys were usually conducted within patch reefs (9 m depth), the outer nearshore hard-bottom and patch reef a proposed or existing protected area reef from 16–30 m depth, and the sites in the middle Florida Keys. Av- such as: Biscayne National Monument fore reef (32–44 m depth) off Boca erage octocoral densities ranged from (now Biscayne National Park) (Voss Raton, Florida. Octocoral diversity 0.02–4.3 colonies/m2, with the lowest et al., 1969; Jaap, 1984), Ft. Jefferson (39 species) and densities were high species richness and colony densities National Monument (now Dry Tortu- (25 colonies/m2). The greatest octo- on either nearshore hard-bottom hab- gas National Park) (Davis, 1982), Key coral diversity occurred on the outer itat in Florida Bay, or on patch reefs Largo National Marine Sanctuary (now reef where the convergence of shal- dominated by massive corals. On the the Key Largo Management Area), low and deepwater octocorals occurs Atlantic Oceanside, hard bottom habi- Looe Key National Marine Sanctuary from 15–30 m depth. Faunal cover for tat exposed to moderately strong tidal (now the Looe Key Management Area) some habitats is similar to that fur- flow was dominated by the generaEu - (Antonius et al., 1978; Wheaton and ther south in the Florida Keys, with nicea, Pseudoplexaura, and Briareum. Jaap, 1988), and Florida state parks hard-bottom areas often dominated On Florida Keys patch reefs, his- (John Pennekamp Coral Reef State by octocorals and sponges. Octocorals torical surveys, mostly from Biscayne Park). Most historical octocoral stud- in the genera Eunicea and Muricea, National Park (BNP), as reported by ies report species occurrences related along with Briareum asbestinum, are Opresko (1973), Jaap and Wheaton11, to benthic habitat types; these data the dominant taxa from the 48 species and Jaap (1984), show variable octo- are largely captured in the life history reported. Based on averages of data coral densities ranging from 6.9–46.8 summaries in the Appendix. In other collected by Blair and Flynn (1989), colonies/m2. Antillogorgia americana, instances, taxonomic or field guides Thanner et al. (2006), and Gilliam et Briareum asbestinum, Eunicea flex- provide more general geographic dis- al. (2007), octocoral density through- uosa, E. succinea, Gorgonia ventali- tribution patterns (e.g., Bayer, 1961; out the southeastern Florida reef tract na, and Plexaura homomalla were the Cairns, 1977; Colin, 1978). With few is approximately 7.7 colonies/m2. most dominant among 29 species pres- exceptions, species inventories were Some areas in Palm Beach County ent (Opresko, 1973). Wheaton (un- usually in shallow-water (< 15 m) have densities of more than 30 octo- publ. data as reported in Wheaton and hard-bottom and reef habitats. In the coral colonies per m² (Gilliam et al., Jaap (1988)) observed 31 species in early 1970s, however, Goldberg (1973) 2003), while hard-bottom habitats off similar habitats in the Biscayne Na- conducted stony coral and octocoral Miami-Dade County have octocoral tional Park area during surveys con- surveys to 44 m depth off Palm Beach densities between 2.9–21.2 colonies/ ducted over five years. In hard-bottom County (described below). In 1975, m² (Blair and Flynn, 1989). areas of Biscayne National Park, in- Wheaton (1981) conducted qualita- Broward County’s Marine Biologi- cluding nearshore tidal passes, beneath tive surveys (species identification and cal Monitoring Program monitors the bridges, and seaward of the intertidal photographs) at Carysfort Reef in the status and trends of coral and hard- zone, octocoral densities were 38.3 upper Florida Keys using replicate belt bottom communities. Since 1997, the colonies/m2 and on transitional bank transects sampled to 21 m depth and Broward County Department of Plan- reefs, densities were 25.2 colonies/m2 recorded 27 taxa. Additional visual ning and Environmental Protection (Jaap, 1984). surveys of octocorals using a submers- and the National Coral Reef Institute Surveys conducted at six sites with- ible were conducted in 1979 at French at Nova Southeastern University have in Looe Key National Marine Sanctu- Reef, Elbow Reef, and Carysfort Reef quantified octocoral densities along within the Key Largo Coral Reef Ma- belt photo-transects at 25 permanent 11Jaap, W. C., and J. L. Wheaton. 1977. Unpub- lished report to the National Park Service on rine Sanctuary at depths of 27–56 m sites on the nearshore and offshore coral research in Biscayne National Monument. (Wheaton, 1987). reef terraces (Gilliam et al., 2004a). Cited In J. Dupont, W. Jaap, and P. Hallock. 2008. Further north along the southeast- At these sites, octocoral density was A retrospective analysis and comparative study 2 of stony coral assemblages in Biscayne National ern Florida reef tract from Miami- 8.9 ± 13.9 colonies/m and as high as Park, FL (1977–2000). Carib. J. Sci. 44:334–344.

80(3) 39 ary reported mean octocoral densities ment Monitoring (SCREAM) program taining targeted reef and hard-bottom ranging from 1.1–20.6 colonies/m2 conducted benthic surveys of algae, habitats are the primary sample units. (Wheaton and Jaap, 1988). Of the 23 sponges, and cnidarians, including oc- A second-stage sample unit is defined octocoral species observed, Eunicea tocorals, in a broad range of hard-bot- as a belt transect of fixed area within a flexuosa, E. succinea,and Plexaura tom and coral reef habitats from the primary sample unit. The size of an in- homomalla comprised nearly 55% of northern extent of Biscayne National dividual primary sampling unit allowed the octocoral fauna. Additional species Park to the Dry Tortugas (Miller et al., divers to swim to the location of any recorded in relatively high densities 2002; Smith et al., 2011). Originally given second-stage sampling unit from were Antillogorgia americana, Gorgo- designed to document the status and a moored or anchored vessel. During nia ventalina, and Muricea atlantica trends of no-take management zones 1999–2004, octocorals were surveyed (Wheaton and Jaap, 1988). throughout the FKNMS, the program for colony densities in replicate 25 m The CREMP, spearheaded by the expanded to include multiple habi- x 0.4 m belt transects. From 2005–14, Florida Fish and Wildlife Research In- tats from nearshore patch reefs to the the belt transect survey area was re- stitute, has monitored the status and deeper reef slope. By broadly sam- duced to replicate 8 m x 1 m samples trends of 37 sites in the Florida Keys pling populations among multiple per site based upon results from pre- National Marine Sanctuary (FKNMS) habitat types across the south Florida vious surveys that indicated slightly since 1995 (Ruzicka et al., 2009) and shelf, inside and outside of the no-take smaller sample sizes were sufficient to in 2012 added an octocoral demo- management zones, and throughout maintain targeted variance (measured graphic survey component at a subset the Florida Keys from south of Mi- as coefficient of variation). Within belt of sites from the upper Keys to the ami to the Dry Tortugas, over a 15-yr transect areas, octocorals are identified Dry Tortugas. In 2003, additional sites period (including 2014 surveys of oc- to the lowest taxonomic level possible were added in Miami-Dade, Broward, tocorals), this effort documented the (usually to the species level) and col- and Palm Beach Counties as part of distribution, abundance, and chang- onies (ramets) are counted. All visual the Southeast Florida CREMP (SEC- es over time of coral reef organisms surveys are conducted using SCUBA, REMP). This effort currently encom- and communities in the region. In the pre-formatted data slates, pencils, and passes 17 sites (Gilliam et al., 2004b). section below, data are summarized rulers. Results are transcribed the same Using video and still photography, related to octocoral surveys, includ- day into a database to help facilitate benthic cover is assessed by identify- ing recently conducted surveys dur- quality control and quality assurance. ing the major functional cover groups ing June–October 2014. The surveys To control for spatial variation in the (macroalgae, sponges, stony corals, provide information on distribution, benthic variables assessed, the Florida and octocorals) within replicate belt density, and population abundance es- Keys survey domain was partitioned transects. Benthic cover by octocorals timates of Florida Keys octocorals. into strata based upon: 1) habitat is similar throughout the Florida Reef The survey sample design is de- type and depth, 2) geographic region Tract from Palm Beach County to the scribed in Smith et al. (2011) and par- (along-shelf position), and 3) manage- Dry Tortugas (Gilliam, 2012). Per- allels efforts for reef fish surveys in ment zones comprising Biscayne Na- centage cover by octocorals increased the south Florida region since the late tional Park (BNP), areas outside of significantly since 2010 in parts of Mi- 1990s. For octocorals, data have been Florida Keys National Marine Sanctu- ami-Dade County (12.8% in 2010 vs. collected on species presence-absence, ary no-take zones, and Sanctuary no- 19.7% in 2011) and Broward County colony densities, and colony heights. take zones. Grid cells (sites) 200 m x (5.4% in 2010 vs. 7.6% in 2011). In Briefly, a geographic information sys- 200 m in dimension are randomly se- Palm Beach County, however, sites tem (GIS) containing digital layers for lected from the combination of habitat within each of the three reef tracts benthic habitat, bathymetry, and man- type, regional sector, and management have shown a decrease in octocoral agement zone boundaries in the Florida zone factors. Habitats are designated cover since the project’s inception. Keys is used to facilitate delineation of using regional benthic habitat maps. Gilliam (2012) reported that several the Florida Keys study area. Existing The habitat classification scheme ac- Palm Beach County study sites buried resolution of benthic habitat mapping counts for features that correlate with by sand, possibly from beach renour- is such that the survey domain was di- benthic faunal distributions, includ- ishment activities and/or storm events, vided into a grid of individual cells 200 ing cross-shelf position, topographic may have contributed to the decreased m x 200 m (40,000 m2) that serve as complexity, and the proportion of sand octocoral cover. primary sampling units. A two-stage interspersed among hard-bottom struc- sampling scheme was adapted (Smith tures. A geographic regional stratifica- Geographic Distribution et al., 2011) following Cochran (1977) tion variable (i.e., upper, middle, and and Density Patterns and is employed to control for spa- lower Florida Keys) was used to ac- in the Florida Keys tial variation in population metrics at count for oceanographic and geologi- From 1999–2014, the Sanctuary scales smaller than the grid cell mini- cal features in the Florida Keys that Coral Reef Evaluation and Assess- mum mapping unit. Grid cells con- influence the distribution and com-

40 Marine Fisheries Review Table 8.—Sampling effort for octocorals in the upper Florida Keys from northern Key Largo to Upper Matecumbe • 2005: 195 sites from northern Key during June–October 2014. The number of sites available represents the number of 200 m x 200 m mapping grid cells based upon existing habitat mapping and bathymetry data. Two 8 m x 1 m belt transects were surveyed Biscayne National Park to Key for octocoral densities and colony sizes. West; No. sites Habitat area No. sites Area sampled • 2006: 46 sites in the Dry Tor­- Benthic habitat type available (m2) sampled (m2) tugas; Inshore patch reefs 2 80,000 2 32 • 2008: 145 sites from northern Key Mid-channel patch reefs 622 24,880,000 17 272 Offshore patch reefs 940 37,600,000 26 416 Largo to Key West and 43 addi- Back-reef rubble 62 2,480,000 7 112 Inner line reef tract 97 3,880,000 11 176 tional sites in the Dry Tortugas; High-relief spur and groove 111 4,440,000 16 256 • 2009: 160 sites from northern Low-relief hard-bottom (< 6 m) 513 20,520,000 3 48 Low-relief spur and groove (6–15 m) 367 14,680,000 10 160 Key Largo to Key West; and Low-relief hard-bottom (6–15 m) 480 19,200,000 4 64 • 2014: 102 sites from northern Key Patchy hard-bottom (6–15 m) 98 3,920,000 6 96 Total 3,292 131,680,000 102 1,632 Largo to Upper Matecumbe Key (see Table 8). Data presented in this section rep- munity composition of hard-bottom nate site was sampled instead. Once resents a small proportion of the over- and reef habitats. Management zones on-site, a two- or three-person dive all sampling effort, but illustrate three are incorporated as a third stratifica- team deployed two transect tapes. For important aspects of octocorals in the tion variable that delineates areas open octocorals, an 8 m x 1 m belt area Florida Keys, with a particular focus and closed to consumptive activities. was surveyed along two transects. on those taxa (species, genera, and The power of the stratified random Statistical analyses using this two- color groups) targeted by the octo- sampling approach is two-fold: 1) the stage sampling design are based coral fishery. First, most octocoral taxa habitats comprising the most area are upon Cochran (1977) and Smith et al. (genera and species) are broadly dis- initially allocated more sites than those (2011). Density and abundance cal- tributed on the south Florida shelf. The with less area (i.e., a proportional de- culations are based upon the number availability of hard substratum appears sign); and 2) habitats exhibiting more of organisms recorded within the belt to be one of the most important factors variability with respect to particular transect stations. First, density (no. of affecting the spatial distribution of oc- metrics (e.g., coral species richness, octocoral colonies/m2) is calculated for tocorals. Second, changes in octocoral octocoral density) are allocated more each transect. Next, mean density and densities from 1999–2014 indicate ei- sites than those with less variability. variance are calculated for each site, ther no significant temporal patterns or The habitats sampled during 1999– using the densities of the two stations. increases for many taxa. Finally, popu- 2014 incorporate most of the hard- The mean site-level densities and vari- lation abundance estimates, integrat- bottom and coral reef habitat types ances are then used to calculate mean ing density estimates with habitat area, from the island platform (e.g., in- stratum-level (habitat, management indicate that octocoral populations are shore patch reefs such as Tavernier zones, and habitat by management either stable or have increased during Rocks) inshore of Hawk Channel to zone) densities and variances. Finally, the past 15 years. Population sizes for ~15 m depth along the Florida Keys, stratum-level and domain abundance many taxa are quite large (tens to hun- including the Dry Tortugas. Benthic estimates are calculated based upon dreds of millions of colonies). habitats surveyed for octocorals in- the stratum-level densities and vari- It is noteworthy that this sampling clude inshore patch reefs, mid-chan- ances, as well as the proportional ar- program does not specifically target nel patch reefs, offshore patch reefs, eas of each stratum within the domain nearshore hard-bottom or ma- reef rubble, shallow (< 6 m) hard- (Smith et al., 2011). trix habitats, where, for example, some bottom, shallow (< 6 m) spur and Octocoral survey data (species rich- octocorals such as Pterogorgia anceps groove, and deeper fore-reef habitats ness, octocoral densities, and in some and P. guadalupensis can be abun- (low-relief spur and groove, consoli- cases colony heights) from this pro- dant (Voss and Voss, 1955; Voss et al., dated hard-bottom, patchy hard-bot- gram were derived from the following 1969; Chiappone and Sullivan, 1994). tom) from 6–15 m depth. In certain efforts from 1999–2014: Deeper-water (> 20 m) habitats are years, deeper habitats were sampled also excluded for most years, which to 27 m depth. The underwater sur- • 1999: 80 sites from northern Key means that taxa in the “red” collec- veys occurred at randomly selected, Largo to Key West and 23 addi- tor group are usually not encountered pre-determined coordinates, located tional sites in the Dry Tortugas; (e.g., Iciliogorgia schrammi and Swif- with a differential global positioning • 2000: 45 sites from Big Pine Key tia exserta). In addition, the Flori- system. If the original waypoint was to Key West and 36 additional da Keys surveys are limited to just a not the intended habitat type, based sites in the Dry Tortugas; small portion of the Florida coastline on visual assessment by a snorkeler • 2001: 86 sites from northern Key and thus do not include certain taxa or boat observers, the closest alter- Largo to Key West; (e.g., Leptogorgia) that are distributed

80(3) 41 Figure 11.—Density (no. colonies/m2) distribution maps A) for all encountered branching octocoral species combined and B–P) alphabetically by species in the upper Florida Keys from Carysfort Reef to Alligator Light, based upon surveys of replicate 8 m x 1 m belt transects sampled per site at 102 hard- bottom and reef sites during June–October 2014. Figure 11 continues on next eight pages.

further north in warm-temperate en- tocorals the authors are aware of for ing population estimates for species vironments on both the Atlantic and the area. that are patchy or rare requires add- Gulf of Mexico coasts. Figure 11 illustrates the density dis- ing more sites (possibly hundreds) to An example of the fishery indepen- tribution patterns of octocorals in the the sample design, increasing the costs dent octocoral data are presented be- upper Florida Keys (northern Key Lar- and amount of in-water effort for the low for the upper Florida Keys from go to Upper Matecumbe), based upon overall survey. Carysfort Reef to inshore of northern surveys from June–October 2014 (Ta- Figure 12 and Table 9 show densi- Key Largo (Carysfort Reef), south- ble 9). The density distribution maps ty estimates combined for all habitats westwards to Alligator Light to in- indicate that for all octocoral species surveyed in the upper Florida Keys shore of Upper Matecumbe Key. This combined, including those that are area for the following sample peri- represents a small proportion of the targeted in the octocoral fishery, most ods: 1999–2002 (combined), 2005, Florida Keys reef tract, much less species are broadly distributed across 2009, and 2014. For the collector the southeastern Florida coast, but il- the south Florida shelf. A few species, color groups, overall mean densities lustrates some of the basic patterns such as Antillogorgia kallos, Ptero- for individual taxa and for combina- in octocoral distribution, density, and gorgia anceps and P. guadalupensis, tions of taxa in color categories either population sizes that help to place the appear to be relatively rare, but this have shown non-significant temporal octocoral fishery and historical and may be due to the exclusion of near- changes or have increased (Table 9). current collection levels into perspec- shore hard-bottom and seagrass-hard- Colony density increases occurred in tive. These data include the longest bottom matric habitats in the survey nearly all of the taxa under the pur- time series available and the most design and/or the extremely patchy ple color category, in addition to non- current data available (2014) for oc- nature of some octocorals. Improv- targeted taxa. The relative density of

42 Marine Fisheries Review Figure 11.­—Continued.

80(3) 43 Figure 11.­—Continued.

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80(3) 45 Figure 11.­—Continued.

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80(3) 47 Figure 11.­—Continued.

48 Marine Fisheries Review Figure 11.­—Continued.

80(3) 49 Figure 11.­—Continued.

Figure 12.—Octocoral densities ( in the upper Florida Keys from Carysfort Reef, northern Key Largo to Alligator Light, Upper Matecumbe Key, based upon surveys during A) 1999–2002, B) 2005, C) 2009, and D) 2014. Species are ranked along the x-axis based upon 2014 densities. Error bars represent 95% confidence intervals. Octocoral species or genera targeted in the octocoral fishery are indicated with an asterisk (*).

50 Marine Fisheries Review Figure 12.­—Continued.

80(3) 51 Table 9.—Octocoral density estimates (mean no. colonies per m2 ± 95% CI) for sampled hard-bottom and coral identification guides for the western reef habitats in the upper Florida Keys from Carysfort Reef, northern Key Largo to Alligator Light, Upper Mate- cumbe Key during 1999–2014. Asterisk (*) indicate the most highly valued species or genera targeted by the octo- Atlantic that included an illustrated coral fishery. Octocorals are further divided by color group (i.e., purple, red, other). nd = not encountered/no data. key to over 100 species within four Note that octocorals in the red collector group (Diodogorgia, Ellisella, Iciliogorgia, Leptogorgia, Nicella, and Swiftia) were not encountered during the benthic surveys. orders: Telestacea, Alcyonacea, Gor-

Octocoral taxa/collector group 1999–2002 2005 2009 2014 gonacea, and Pennatulacea. In 1977, Cairns (1977) published a follow-up Other species group Diodogorgia nodulifera* nd nd nd nd identification guide of common oc- Muricea spp.*1 1.24 ± 0.53 0.43 ± 1.80 0.29 ± 1.43 0.50 ± 1.01 M. elongata 0.33 ± 0.16 0.16 ± 0.56 0.13 ± 0.85 0.26 ± 0.85 tocorals that included photographs of M. muricata 0.92 ± 0.38 0.15 ± 1.00 0.16 ± 0.57 0.23 ± 0.16 dried colonies from Florida, the Ca- M. pinnata nd nd nd nd Plexaura homomalla 0.33 ± 0.14 0.19 ± 0.13 0.16 ± 0.11 0.29 ± 0.12 ribbean, and Gulf of Mexico. Sán- Plexaurella dichotoma/fusifera 0.25 ± 0.87 0.36 ± 0.16 0.29 ± 0.13 0.68 ± 1.24 chez and Wirshing (2005) improved P. nutans* 0.28 ± 0.41 0.37 ± 0.45 0.13 ± 0.25 0.30 ± 0.13 Pterogoria citrina* 0.23 ± 0.11 0.30 ± 0.26 0.28 ± 0.20 0.25 ± 0.86 upon these and other earlier works, Total density (other) 2.33 ± 2.06 1.65 ± 2.81 1.14 ± 2.12 2.02 ± 3.36 with their field guide to zooxanthel-

Purple species group late octocorals in the western Atlantic. Antillogorgia spp.*2 3.05 ± 1.21 4.64 ± 2.42 5.41 ± 2.21 7.86 ± 5.01 High-resolution in-situ photographs Briareum asbestinum* 0.29 ± 0.17 1.14 ± 0.58 0.93 ± 0.57 0.33 ± 0.13 Eunicea spp.3 2.30 ± 3.16 3.59 ± 2.49 1.58 ± 2.72 3.22 ± 3.29 and flow charts based on basic mor- E. flexuosa* 0.29 ± 0.59 0.53 ± 0.16 0.35 ± 0.12 0.54 ± 0.16 Muriceopsis flavida* 0.39 ± 0.26 0.45 ± 0.23 0.53 ± 0.21 1.33 ± 0.60 phological characters were meant to Pseudoplexaura spp.4 0.93 ± 0.74 0.59 ± 0.35 0.98 ± 0.89 1.44 ± 1.30 be used as a quick field reference for P. porosa 0.13 ± 0.14 0.12 ± 0.90 0.33 ± 0.25 0.48 ± 0.23 Pterogorgia anceps* 0.28 ± 0.32 0.12 ± 0.11 0.75 ± 0.58 0.36 ± 0.42 identification. In 2011, Dr. C. Mess- P. guadalupensis 0.66 ± 0.60 0.13 ± 0.11 0.13 ± 0.25 0.27 ± 0.32 ing of Nova Southeastern University Total density (purple) 8.32 ± 7.19 11.32 ± 7.36 10.97 ± 7.79 15.82 ± 11.47 (NSU) spearheaded a workshop to Not targeted develop an online interactive guide A. americana 3.97 ± 0.92 3.52 ± 0.61 4.42 ± 0.88 4.40 ± 0.73 Gorgonia ventalina 1.78 ± 0.49 1.67 ± 0.57 2.87 ± 0.80 2.99 ± 0.89 to South Florida octocorals funded as P. kuna4 nd 0.32 ± 0.55 0.14 ± 0.11 0.65 ± 0.60 part of this overall octocoral fishery P. grisea 0.32 ± 0.13 0.85 ± 0.40 0.68 ± 0.46 0.12 ± 0.47 Total density (not targeted) 6.07 ± 1.53 6.37 ± 2.13 8.10 ± 2.25 8.16 ± 2.69 investigation. With assistance from leading octocoral experts, samples Total octocoral density (all groups) 16.72 ± 10.79 19.33 ± 12.29 20.22 ± 12.17 26.00 ± 17.52 that had been photographed in-situ, 1Includes the taxa M. elongata, M. laxa, and M. muricata 2Includes the taxa A. acerosa, A. bipinnata, A. elisabethae, A. kallos, and A. rigida. Although A. acerosa can be yellow- and then preserved, were positively golden in color and is included in both the purple and other categories, > 99% of colonies encountered are purple and identified using microscopic examina- are thus included in the purple category. 3Includes the taxa E. calyculata, E. fusca, E. knighti, E. mammosa, E. palmeri, E. succinea and E. tourneforti tion of sclerites. Large dried colonies 4Includes the taxa P. flagellosa, P. wagenaari, and unidentifiable Pseudoplexaura colonies 5Plexaura kuna was not distinguished from Eunicea flexuosa, then known as Plexaura flexuosa, during 1999-2002. from the NSU invertebrate collec- tion were also photographed. The ma- jor outcome of the workshop was a web-based octocoral guide (www. octocoral species, comparing the top based upon surveys in 1999–2002, cnso.nova.edu/ncri/sofla_octocoral_ ten densest species in the upper Flor- 2005, 2009, and 2014 (Fig. 13). Table guide/index.html/). Users can search ida Keys, remained mostly unchanged 11 lists octocoral abundance estimates through a list of octocoral species or (Fig. 12). for the Florida Keys, excluding the click through character menus re- Marquesas and Dry Tortugas, from lated to growth form, color, or aper- Florida Keys Population 1999–2009, which are referenced be- Abundance Estimates ture shape to narrow species searches. low to help evaluate the impact of Once a species is selected, a page is Estimates of octocoral abundance octocoral fishery landings from FWC linked that includes information on its for the upper Florida Keys are illus- trip ticket data relative to estimated habitat and ecology, along with pho- trated in Figure 13 and Table 10. Pop- population sizes. tographs. Currently, the South Florida ulation abundance estimates are based octocoral identification guide includes upon colony densities and habitat area Fishery Information Needs and Management Recommendations 47 species. The guide, plus other tax- and demonstrate the large population onomic records, were used to help sizes (tens to hundreds of millions Assessment of inform the production of a fishery- of colonies) for most gorgonian taxa. Identification Aides specific identification guide that was Total colony abundances in the upper General knowledge about octo- developed as part of this octocoral Florida Keys either suggest non-sig- fishery project. nificant changes over time or increas- corals targeted in the marine life trade is probably sufficient to address many To supplement the online interac- es (Table 10). The relative abundance tive South Florida octocoral guide of octocoral species, comparing the of the fundamental questions related to species identifications, life history, and described above, a waterproof field top ten densest species in the upper guide has been suggested as a poten- Florida Keys, is mostly unchanged population abundance. Bayer (1961) published one of the first octocoral tially useful tool for octocoral collec-

52 Marine Fisheries Review Figure 13.—Octocoral abundance (total colonies) in the upper Florida Keys from Carysfort Reef, northern Key Largo to Alligator Light, Upper Matecumbe Key, based upon surveys during A) 1999–2002, B) 2005, C) 2009, and D) 2014. Species are ranked along the x-axis from high-to-low based upon 2014 abundances. Error bars represent 95% confidence intervals. Octocoral species or genera targeted in the octocoral fishery are indicated with an asterisk (*). Figure 13 continues on next page. tors, with a broader intended audience species may be known by several corals. To assist in identification, basic to include law enforcement, hobbyists, common names (Bayer, 1961; Voss et habitats where octocoral species occur, and the public. A guide is currently be- al., 1969; Cairns, 1977; Cairns et al., collection depths, and distinguishing ing developed that will be organized 2002). This complicates and confuses diagnostic characteristics will be noted by the color headings currently used to the tracking and marketing of species for each species. Figure 14 presents a report collection activities in the FWC sold within the fishery. Furthermore, draft page from the field guide. If on- Trip Ticket reports (i.e., other, purple, species misidentification is an ongo- going interest exists, workshops could and red categories). Under each head- ing problem for both the fishery and be held to assist collectors in the use ing, color photographs taken in situ of scientific communities. A goal of the of these guides. the octocoral species commonly col- field guide is to provide a tool that lected in the fishery will list both the can be used by all participants within Life History Traits scientific name and the official com- the marine life fishery (e.g., collector, There is nothing unusual or notable mon name. A major discrepancy with- wholesaler, hobbyist, and law enforce- about octocoral life histories that sug- in the marine life fishery is the misuse ment) to help create consistency with gest that the current level of collec- of common names such that a single the identification and naming of octo- tion, based upon collector interviews,

80(3) 53 Figure 13.­—Continued.

FWC trip ticket data, and available the aquarium trade (23 species and 15 ment, settle immediately after detach- fishery independent assessments, is genera), reproductive mode and tim- ing from the parental colony (Brazeau a threat to the sustainability of oc- ing of reproduction is known for 12 and Lasker, 1990). This is significant tocoral populations. The majority of species. The majority of these (10 of because brooding colonies can con- wider Caribbean octocorals repro- 12 species) are broadcast spawners tribute directly to the population dy- duce sexually, with male and female that release gametes during the sum- namics of the immediate reef area. If colonies that are distinct. There is no mer months. Antillogorgia elisabe- substantial quantities of adult colonies indication that octocoral collectors se- thae and Muriceopsis flavida, on the of any of these brooding species were lectively target colonies based upon other hand, release gametes during the removed from a single area, similar to gender. Many species also reproduce winter months (Kahng et al., 2011). forest clear-cutting, then one would asexually through fragmentation and Brooding species, where males release expect a significant decrease in repro- vegetative propagation, supporting gametes that fertilize eggs retained by ductive output (Goffredo and Lasker, high population growth rates (Lasker, females, include A. elisabethae, Bria- 2008). The collectors interviewed for 1988; Kahng et al., 2011). Many oc- reum asbestinum, Muriceopsis flavida, this report stated that clear-cutting for tocoral colonies spawn over several and Pterogorgia anceps (Ritson-Wil- octocorals is not practiced. days over consecutive months in a year liams, 2010; Kahng et al., 2011). The Octocorals generally grow faster (Fitzsimmons-Sosa et al., 2004; Guti- planula of brooders, which form from than most stony corals, with linear errez-Rodriguez and Lasker, 2004). the fertilized egg and have cilia or branch extensions of 4 cm/yr in thin- Of the octocoral species collected for hair-like projections that aid in move- ly branched species of Antillogorgia,

54 Marine Fisheries Review gonia ventalina, Plexaura kuna, P. homomalla), gaps in life history data exist for several targeted genera and species. Besides distribution data and one study on reproduction (Ritson- Williams, 2010), for example, the only well-studied aspect of the genus Ptero- gorgia are the secondary metabolites found within its tissues (Schmitz and Lorance, 1971; Lorance, 1977; Paw- lik and Fenical, 1992; Rodriguez and Ramirez, 1994; Jensen et al., 1996). A brief observation in October 2009 of surface brooding in colonies of P. anceps in Belize was noted by Ritson- Williams (2010). This paucity of life history data is somewhat unusual for a common, easily identifiable, shallow- water genus. As noted above, surveys focusing on size and mode of reproduc- Figure 14.—Draft page for the Octocoral Field Identification Guide. tion would be relevant in Pterogorgia to determine how size and morphol- ogy are related to reproductive output. Muricea, and Muriceopsis. However, Colonies of A. elisabethae are collect- However, this interest might be largely thicker branched species such as Plex- ed by cropping colonies that are 5–15 academic, because there is no evidence aurella barely grow 1 cm/yr (Yoshioka cm in height and collectors rotate sites to suggest that population numbers and Yoshioka, 1991). Faster-growing on a 2–3 yr basis to allow recovery are being adversely impacted at cur- species tend to be found at higher den- before re-collecting. Castanaro and rent collection levels in the U.S. South sities (Yoshioka, 1997). Octocorals Lasker (2003) measured accelerated Atlantic. There is also a paucity of life typically reach sexual maturity when growth rates among collected colo- history information on octocorals of they are over 20 cm in height. An ex- nies, particularly those that retained the genus Muricea; only information ception is Pterogorgia citrina, a small at least 12 branches in size. However, on the breeding period of M. atlantica bushy octocoral that rarely grows because collected colonies are small- (gametes are present June–September) above 30 cm in height that may reach er, fewer zygotes are produced, re- is available (Fitzsimmons and Hallock sexual maturity sooner; however, little ducing recruitment at collection sites. Mueller, 1996). Even though it is a is known about this species. To provide As a result, collectors have switched commonly requested octocoral for the adequately sized octocorals to home to collecting at 5-yr intervals to give aquarium trade, additional studies are aquarists, collectors usually take col- colonies greater time to reproduce be- not considered a priority because ex- onies < 20 cm in height. In doing so, fore the next collection (Goffredo and isting population numbers appear to be they leave larger colonies behind that Lasker, 2008). Octocoral collectors large enough in relation to current col- are the major contributors of gametes in Florida who clip 20-cm fragments lection levels. to the population (Beiring and Lasker, from adult colonies for Eco-Gorgs® Studies of octocorals at deeper (30+ 2000). The octocoral collectors inter- leave behind over 80% of the colony. m) depths (e.g., Diodogorgia nodu- viewed indicated that they rotate col- The impact on reproduction is prob- lifera and Swiftia exserta) have primar- lection sites and are selective in which ably minimal to individual colonies ily described their presence or absence colonies they collect. As aesthetics is a and insignificant to the larger popu- at various locations throughout the wid- major factor in marketability of octo- lation because only a small fraction er Caribbean (Bayer, 1961; Veronique, coral colonies, many ‘imperfect’ colo- of the larger population is clipped for 1987; Humann, 1993). Density data nies are passed over by collectors. fragments, but this requires additional for these two deepwater species are not In the Bahamas, colonies of Antil- study to confirm. available from the literature. Anecdot- logorgia elisabethae are clipped and ally, information from collectors sug- collected for the cosmetics industry. Information Gaps gests that these species are found in The bioactive compound pseudopter- in Life History high densities in Palm Beach County, osin found in this species is an anti- Though some species of octocorals with colonies spaced a few centime- inflammatory agent that is added to are relatively well-studied (e.g., Antil- ters apart, and at relatively high colony cosmetic products (Look et al., 1986). logorgia spp., Eunicea flexuosa, Gor- densities in Miami–Dade County, with

80(3) 55 Table 10.—Octocoral abundance estimates (total colonies ± 95% CI) in sampled hard-bottom and coral reef habi- tors have observed morphological dif- tats in the upper Florida Keys from Carysfort Reef, northern Key Largo to Alligator Light, Upper Matecumbe Key during 1999–2014. Asterisks (*) indicate the most highly valued octocoral species or genera targeted by the fish- ferences between S. exserta colonies in ery. Octocorals are further divided by color group (i.e. purple, red, and other). nd = no data. Note that octocorals Palm Beach County, where colonies are in the red collector group (e.g. Diodogorgia, Ellisella, Iciliogorgia, Leptogorgia, Nicella, and Swiftia) were not en- countered during the upper Florida Keys benthic surveys. short and rigid, with those in Miami–

Octocoral taxa/collector group 1999-2002 2005 2009 2014 Dade County, where colonies are tall and flexible. Growth and regeneration Other species group Diodogorgia nodulifera* nd nd nd nd rates of S. exserta would be interesting Muricea spp.*1 16,797,660 ± 34,960,192 ± 39,246,900 ± 67,777,312 ± 7,211,207 20,322,052 19,373,360 25,963,333 to obtain related to collection methods ® M. elongata 4,359,698 ± 14,376,154 ± 17,839,500 ± 35,989,570 ± (e.g., fragmentation for Eco-Gorgs ), 2,111,128 7,629,469 11,605,665 11,493,995 but there is no evidence that current M. muricata 12,437,962 ± 20,420,673 ± 21,407,400 ± 31,787,742 ± collection levels are negatively affect- 5,100,079 12,368,587 7,767,695 14,469,338 M. pinnata nd nd nd nd ing populations of these two species. Plexaura homomalla 4,487,925 ± 25,811,731 ± 21,917,100 ± 38,912,581 ± 1,945,163 18,330,061 15,532,634 16,817,943 Population Abundance Estimates Plexaurella spp.* 28,209,811 ± 49,826,442 ± 39,586,700 ± 64,397,581 ± 12,326,630 22,861,624 18,595,991 33,864,573 Benthic surveys for octocorals con- Pterogorgia citrina* 30,902,566 ± 39,534,423 ± 37,887,700 ± 27,859,946 ± ducted in the Florida Keys during the 15,140,642 35,182,956 26,772,620 11,742,691 past 15 years provide habitat-specific Total abundance (other) 80,397,962 ± 150,132,788 ± 138,638,400 ± 198,947,419 ± density and population abundance es- 36,623,642 96,696,692 80,274,605 88,388,542 timates. Time series data from 1999– Purple species group Antillogorgia spp.*2 374,677,585 ± 478,170,481 ± 688,264,900 ± 923,580,054 ± 2014 for octocoral taxa are available 132,298,971 176,419,729 258,903,714 412,160,297 for the upper Florida Keys, repre- Briareum asbestinum* 39,237,283 ± 155,197,115 ± 125,726,000 ± 44,758,602 ± senting the longest time series avail- 23,377,355 79,368,560 68,905,724 18,241,600 able for density (Fig. 12, Table 9) and Eunicea spp.3 99,631,925 ± 219,399,712 ± 170,579,600 ± 227,355,430 ± 33,934,826 157,299,336 96,285,956 149,445,948 abundance data (Fig. 13, Table 10). E. flexuosa* 38,980,830 ± 72,207,500 ± 47,062,300 ± 73,166,613 ± Total abundance for collected spe- 8,071,085 22,209,603 15,861,707 21,973,470 cies in the Florida Keys is presented Muriceopsis flavida* 40,904,226 ± 61,752,115 ± 71,358,000 ± 181,226,667 ± 27,271,051 30,872,926 28,896,058 80,268,316 for 2009, the most recent sampling Pseudoplexaura spp.4 12,694,415 ± 34,960,192 ± 13,252,200 ± 36,811,667 ± that included the entire Florida Keys 8,835,166 20,849,658 12,104,263 17,624,464 from northern Biscayne National Park P. porosa 1,795,170 ± 15,029,615 ± 45,023,500 ± 65,859,086 ± 1,406,924 12,218,382 34,347,016 31,928,218 to southwest of Key West (Table 11). Pterogorgia anceps* 384,679 ± 1,633,654 ± 1,019,400 ± 4,841,237 ± The surveys were conducted as part 431,954 1,529,584 783,155 5,711,015 P. guadalupensis 897,585 ± 1,797,019 ± 169,900 ± 365,376 ± of a Florida Keys-wide monitoring 815,207 1,550,504 337,117 437,514 and assessment program designed to Total abundance (purple) 609,203,698 ± 1,040,147,404 ± 1,162,455,800 ± 1,557,964,731 ± evaluate benthic community structure 236,442,539 502,318,282 516,424,711 737,790,843 nearshore to offshore, among different Not targeted habitat types, and inside and outside of A. americana 539,192,075 ± 478,170,481 ± 600,766,400 ± 597,573,011 ± 125,201,041 82,948,559 119,118,372 95,566,345 marine protected areas (Miller et al., Gorgonia ventalina 241,835,019 ± 227,241,250 ± 390,090,400 ± 370,948,333 ± 2002; Smith et. al., 2011). However, 66,173,864 77,682,730 107,499,242 96,755,309 P. kuna4 nd 4,410,865 ± 1,868,900 ± 822,098 ± many nearshore locations used by ma- 7,415,976 1,541,617 813,253 rine life collectors, such as tidal chan- P. grisea 4,103,245 ± 10,945,481 ± 9,174,600 ± 16,715,968 ±x nels, bridge pilings, and Florida Bay or 1,712,921 5,411,117 1,541,617 813,253 Total abundance (not targeted) 785,130,340 ± 720,768,077 ± 1,001,900,300 ± 986,059,409 ± Biscayne Bay octocoral patches, were 193,087,825 193,087,825 234,466,637 199,487,414 not sampled as part of this program. Total octocoral abundance 1,474,732,000 ± 1,911,048,269 ± 2,302,994,500 ± 2,742,971,559 ± Among the 47 species recorded in the (all groups) 466,154,007 772,473,357 831,165,953 831,165,953 Keys-wide surveys, population den- 1Includes the taxa M. elongata, M. laxa, and M. muricata 2Includes the taxa A. acerosa, A. bipinnata, A. elisabethae, A. kallos, and A. rigida. Although A. acerosa can be yellow- sities and abundances are presented golden in color and is included in both the purple and other collector categories, > 99% of colonies encountered are for 15 species (Fig. 11) that represent purple and are thus included in the purple category. 3Includes the taxa E. calyculata, E. fusca, E. knighti, E. mammosa, E. palmeri, E. succinea and E. tourneforti those collected in the octocoral fishery 4Includes the taxa P. flagellosa, P. wagenaari, and unidentifiable Pseudoplexaura colonies 5Plexaura kuna was not distinguished from Eunicea flexuosa, then known as Plexaura flexuosa, during 1999-2002. (Table 12). Species are also divided for comparison into the “other, purple, and red” groups reported by collec- colonies generally 7 m apart. Several tion exists. Studies of S. exserta have tors. Species from the “red” group are hundred colonies can be collected in a focused on their associated microbiota found at depths outside of the range single dive. Populations of these two (Bruck et al., 2007), morphology (Men- targeted in Florida Keys-wide benthic species may serve as important habitat zel et al., 2012), and biology (Olano surveys (< 15 m depth), so data are for other marine life, but little informa- and Bigger, 2000). Interestingly, collec- unavailable for these species.

56 Marine Fisheries Review In the upper Florida Keys, total oc- Table 11.—Octocoral abundance estimates (total colonies ± 95% CI) for sampled hard-bottom and coral reef habitats from northern Biscayne National Park to east of the Marquesas Keys, Florida Keys, during 1999–2009. tocoral population numbers for both The most highly valued octocoral species or genera targeted are indicated with an asterisk (*) and are further “other” and “purple” groups have in- divided by color group (i.e. purple, red, and other). nd = no data. Note that octocorals in the red collector group creased significantly during the last 15 (e.g. Diodogorgia, Ellisella, Iciliogorgia, Leptogorgia, Nicella, and Swiftia) were not encountered during these surveys. years (Table 10), from over 80 million Octocoral taxa/collector group 1999-2002 2005 2009 colonies in 1999–2002 (combined) to Other species group Diodogorgia nodulifera* nd nd nd almost 200 million in 2014 for octo- Muricea spp.*1 114,466,278 ± 229,332,813 ± 270,783,127 ± corals in the “other,” category, and 23,345,596 44,812,261 54,530,487 from over 600 million to more than M. elongate 54,896,155 ± 129,844,550 ± 149,545,171 ± 1.5 billion in 2014 for octocorals in the 13,416,666 23,802,005 30,493,430 M. muricata 59,403,547 ± 98,981,708 ± 121,237,956 ± “purple category.” In 2009 in the upper 10,601,445 20,419,537 24,037,057 Florida Keys, totals for the “other” and M. pinnata nd nd nd “purple” categories were approximate- Plexaura homomalla 51,131,235 ± 90,685,145 ± 71,650,109 ± ly 140,000 and just over 1.4 billion, re- 13,974,304 24,536,639 23,179,909 Plexaurella spp.* 91,941,210 ± 133,700,718 ± 61,457,283 ± spectively. Total numbers for colonies 22,984,308 35,218,576 28,670,931 Florida Keys-wide in 2009 for “other” Pterogorgia citrina* 47,353,292 ± 58,926,251 ± 61,457,283 ± and “purple” was estimated at just over 16,364,273 36,218,576 28,670,931 Total abundance (other) 304,892,015 ± 512,644,926 ± 502,066,548 ± 500 million and nearly 2 billion colo- 77,668,481 140,749,768 130,939,795 nies, respectively (Table 11). While the Purple species group threshold number of 70,000 colonies Antillogorgia spp.*2 531,244,061 ± 717,169,257 ± 834,535,525 ± allowed for collecting has never been 145,690,832 188,139,742 274,982,791 Briareum asbestinum* 76,833,389 ± 256,057,667 ± 202,025,444 ± exceeded, this threshold represents 28,820,534 91,398,237 77,658,165 0.004% of total octocoral numbers for Eunicea spp.3 389,162,980 ± 547,578,643 ± 416,116,429 ± collected species as determined in 2009. 81,019,210 198,535,609 124,077,575 E. flexuosa* 185,228,376 ± 266,279,655 ± 190,583,527 ± The actual number of colonies reported 23,125,728 40,101,745 35,674,286 by collectors in 2009 was 16,053 for Muriceopsis flavida* 62,159,541 ± 91,947,753 ± 91,947,753 ± “other” and 19,228 for “purple.” When 27,764,515 31,754,005 32,270,055 Pseudoplexaura spp.4 69,971,429 ± 179,075,503 ± 35,070,418 ± compared to the total numbers of octo- 25,084,018 36,586,142 14,600,796 corals in the natural population for each P. porosa 51,464,030 ± 81,975,495 ± 155,932,898 ± group, the percentages are 0.003% and 16,649,265 24,831,817 51,472,259 Pterogorgia anceps* 2,954,689 ± 2,801,050 ± 1,964,400 ± 0.001%, respectively for “other” and 1,725,127 2,158,399 1,092,191 “purple” categories (Fig. 13). Popula- P. guadalupensis 20,421,201 ± 4,565,968 ± 1,247,261 ± tion estimates at the individual species 35,917,649 2,816,245 824,147 level are potentially more relevant to Total abundance (purple) 1,389,439,696 ± 2,147,450,990 ± 2,147,450,990 ± 385,796,878 616,321,940 612,652,265 management interests, but these group Not targeted estimates provide a first-order assess- A. americana 1,112,059,940 ± 1,137,085,998 ± 1,392,367,210 ± ment that documents the large number 147,609,769 141,090,463 154,299,778 of octocorals in the Florida Keys and Gorgonia ventalina 437,122,788 ± 472,620,974 ± 679,304,335 ± the relatively miniscule numbers col- 75,248,774 89,265,986 118,314,731 P. kuna4 nd 5,501,538 ± 2,469,016 ± lected in the octocoral fishery. Since 7,493,196 1,651,251 genera or species level data are not re- P. grisea 17,072,178 ± 34,026,481 ± 20,671,706 ± ported by collectors, estimates of rela- 3,994,974 8,087,015 7,161,260 Total abundance (not targeted) 1,566,254,906 ± 1,649,234,991 ± 2,094,812,268 ± tive percentages collected by species 226,853,517 245,936,660 281,427,021 are not explicitly possible, although Total octocoral abundance 2,094,812,268 ± 4,759,804,279 ± 4,896,380,423 ± there is now a better understanding of (all groups) 273,610,977 377,877,699 406,830,115 the species collected in each species 1Includes the taxa M. elongata, M. laxa, and M. muricata 2Includes the taxa A. acerosa, A. bipinnata, A. elisabethae, A. kallos, and A. rigida. Although A. acerosa can be yellow- color group (Table 2). golden in color and is included in both the purple and other collector categories, > 99% of colonies encountered are purple and are thus included in the purple category. For the “red” species found in deep- 3Includes the taxa E. calyculata, E. fusca, E. knighti, E. mammosa, E. palmeri, E. succinea and E. tourneforti er water (> 20 m depth), anecdotal 4Includes the taxa P. flagellosa, P. wagenaari, and unidentifiable Pseudoplexaura colonies 5 evidence from collectors indicates that Plexaura kuna was not distinguished from Eunicea flexuosa, then known as Plexaura flexuosa, during 1999-2002. large patches of Diodogorgia nodu- lifera and Swiftia exserta exist in Palm that he returns to the same general ed States indicate that S. exserta is Beach County at depths of 18–45 m area several times per year and easily broadly distributed along the edge of and in Miami-Dade County at 25–30 find hundreds of colonies on a single the continental shelf, and deeper. Al- m. The patches are apparently dense dive. In fact, surveys along much of though limited information is available and persistent. One collector reported the southeastern coast of the Unit- from the literature on deeper water oc-

80(3) 57 Table 12.—Octocorals surveyed in fishery independent surveys that are also targeted by octocoral collectors. Center (NMFS/SEFSC). Specifically, Species goup Scientific name Collector common name the SAFMC asked for an “evaluation Other Antillogorgia acerosa golden plume gorgonian of octocoral landings and fishery im- Muricea elongata orange spiny sea rod/rusty gorgonian pacts” because octocoral catch data Muricea muricata spiny sea fan Plexaura homomalla black sea rod were being used to set Acceptable Plexaurella fusifera brown tree gorgonian Biological Catch limits and Overfish Plexaurella nutans giant slit pore/large polyp gorgonian Pterogorgia citrina yellow ribbon/yellow sea whip gorgonian limits rather than population data. An

Purple Antillogorgia acerosa purple or golden willow gorgonian additional concern with the octocoral Antillogorgia elisabethae purple frilly gorgonian fishery is that a substantial amount of Antillogorgia kallos purple plume/bi-pinnate gorgonian Briareum asbestinum corky sea finger/sea stalk gorgonian taxonomic uncertainty exists among Eunicea flexuosa purple candelabra/swollen knob candelabrum gorgonian collectors, wholesalers, and law en- Muriceopsis flavida purple brush/bottle brush gorgonian Pseudoplexaura porosa purple tree gorgonian forcement. When colonies are identi- Pterogorgia anceps purple ribbon/angular sea whip gorgonian fied and lumped by color, rather than Pterogorgia guadalupensis purple blade/flat-blade/grooved-blade sea whip gorgonian by species, information is lost relat- ed to life-history traits or ecological tocoral taxa, Goldberg (1973) reported million colonies). These octocorals are function. These are factors that that D. nodulifera comprising approximate- abundant and grow quickly, two traits could potentially affect management ly 5% of the total colonies on the outer that suggest that current collection strategies. fore reef off Palm Beach County. S. levels probably do not negatively af- A goal of this study was to provide exserta was observed in relatively high fect these species. In contrast, all three the SAFMC and FWC with the best abundance at 30% in the same loca- species in the genus Pterogorgia (P. available information to help manage tion and lower in abundance (1.25%) citrina, P. anceps, and P. guadalupen- the octocoral fishery, with the least inshore. sis) and Plexaurella fusifera are less economic impact on the collectors While collectors are not currently abundant (though ranging from hun- who depend upon this resource for required to report at the species level, dreds of thousands to tens of millions income. The management recommen- population data for each of the 15 spe- of colonies), but little is known about dations are summarized as follows. cies targeted is available (Tables 10 whether or not these species are target- Based on interviews with collectors and 11). At the species level, octocoral ed by collectors. If one or more of the and a synthesis of FWC Trip Ticket abundance has increased since 1999 in Pterogorgia species are brooders, local results, the long-term stability of the the upper Florida Keys for all species collection at “hot spots” may affect re- octocoral fishery is not likely to sig- in the “other” group, except Pterogor- production and successful recruitment, nificantly change, so there is there- gia citrina, which appears relatively as Castanaro and Lasker (2003) docu- fore little concern to be had about the stable over the period considered. For mented with collection of A. elisabe- fishery exceeding the 70,000 colonies/ the “purple” group, most species have thae in the Bahamas. For example, P. yr threshold. The social dynamics of increased or remained stable. Popula- citrina has a relatively small maximum the aquarium industry to seek color- tion sizes of Pterogorgia anceps and colony size, and if collectors target the ful, rare, and exotic marine species for P. guadalupensis increased between larger fecund colonies or perhaps a home aquaria places octocorals on the 1999 and 2005, then fell in 2009 and color morph (e.g., yellow vs. purple), low end of the desired list of marine increased again in 2014. It is difficult there may be at least a local negative life species. Although some octocoral to identify the causes of these fluc- effect on populations. Despite uncer- species are vibrantly colored (e.g., An- tuations; however, natural changes tainties about population estimates for tillogorgia acerosa, Muriceopsis flavi- in population dynamics can occur, as some species, there is nothing in the da, Leptogorgia spp.), they are neither large colonies break or become de- abundance estimates to suggest that rare nor exotic. Species that are col- tached because of high wave action the current collection levels for octo- lected from deeper habitats (e.g., Swif- due to storms. In addition, B. asbesti- corals is adversely affecting octocoral tia exserta and Diodogorgia spp.) num is more susceptible to bleaching- populations in the Florida Keys. are difficult to keep in good health in related mortality. home aquaria and are marketed as ap- Currently, octocoral abundance in Fishery Management propriate for advanced aquarists, thus the upper Florida Keys is dominated Recommendations resulting in minimal demand. Octo- by Antillogorgia species (Fig. 12, Ta- Information assembled for this corals also do not contribute ecosys- ble 10). Three species are particularly study aimed to address management tem services in home aquaria, such as prized by marine aquarium hobbyists concerns about the octocoral fish- control and removal of algae and detri- due to their vibrant purple color: A. ery that were raised by the South At- tus, which also limits demand. In addi- elisabethae (~416 million estimated lantic Fishery Management Council tion, during the last 20 years, octocoral colonies), A. kallos (~392 million col- (SAFMC) in a 22 June 2010 memo landings have fluctuated slightly or ac- onies), and Muriceopsis flavida(~181 to NOAA’s Southeast Fishery Science tually decreased.

58 Marine Fisheries Review Distribution and abundance in- levels), natural populations are prob- they might benefit the collectors by formation obtained from an exhaus- ably not threatened. These results also helping them to better market octo- tive literature search, along with age, suggest that if demand for octocorals corals. Additional taxonomic resolu- growth, population and habitat data, were to increase substantially, that the tion for the Trip Ticket program is not suggests that current information is 70,000 colony quota could reasonably being proposed, as it is not necessary adequate to describe the populations be increased, assuming the reason for for ensuring sustainability of the octo- and the octocoral fishery, making a the increased demand can be rationally coral fishery at current levels. While formal stock assessment probably a evaluated. For example, if demand was collectors may benefit from the addi- low priority. The list of 12 targeted to increase, is it based upon one spe- tional data, voluntary participation is species in original scope of work for cies, one group of species, or spread recommended. this study was expanded to 24 species across all species? In addition, bet- Information obtained and compiled based on interviews with collectors. ter understanding of collecting meth- for this study suggests that addition- Considerable lumping of taxa based ods would be relevant. For example, al research is not necessarily needed upon colony color still occurs. Scien- was demand being met through Eco- to better manage the octocoral fish- tifically, there are genera where addi- gorgs®, other fragmentation methods, ery. While the population biology of tional information on population status rotating sites, or something else? deepwater species remains largely un- and life history information would Data collected by FWC through the known, there is no evidence that pop- be academically interesting (e.g., for Trip Ticket program is adequate to ulations are being adversely affected the genera Muricea, Pterogorgia, and understand and manage the octocoral by the relatively low numbers of colo- Diodogorgia), but based on the cur- fishery, although FWC has proposed nies collected. Octocoral collection in rent state of knowledge and collecting refinements to the reporting criteria the Bahamas for the cosmetics indus- levels, this additional information is beyond the three current octocoral try and the artisanal jewelry trade, for probably a low priority for informing categories. Regional distinctions that example, previously prompted studies management needs related to species- separate Florida Keys landings from on growth and reproduction of tar- specific, seasonal, or depth-driven reg- others in Atlantic Ocean waters would geted octocoral species to understand ulation of annual quotas. be beneficial for understanding what the effects of repeated and extensive The collection of octocorals, below parts of the Florida Reef Tract are uti- clear-cutting, sometimes to the point the 70,000 colony per year quota, a lized for collecting octocorals, since that only holdfasts remained (Morel level that has never been reached, does management reviews are underway in et al., 1996; Castanaro and Lasker, not appear to adversely impact octo- Monroe, Miami-Dade, Broward, and 2003; Cadena and Sanchez, 2010). In coral populations. For multiple sam- Palm Beach Counties. Collectors who Florida, collection for the aquarium pling periods spanning over a decade, were interviewed indicated that they trade dramatically contrasts with that the Florida Keys abundance estimates would like clarification in the report- conducted by commercial collectors presented for 15 species illustrate that ing methodology for clipped frag- for the pharmaceutical industry. The octocoral populations are large (tens ments versus whole colonies collected. selective removal of adult colonies, of millions to hundreds of millions, For example, should three clippings sub-adult colonies, or branches from per species) and despite some fluctua- from a single colony be recorded as adult colonies at current levels does tions that are likely due to natural vari- three colonies or one colony? not appear to warrant any change to ation (along with sampling issues and Collectors and aquarium hobbyists the current fishery management plan taxonomic questions) are generally in- stated that they would welcome and for octocorals. creasing. This result provides evidence use a field guide to help with octo- that current and projected octocoral coral identification, which would ul- Acknowledgments collection levels are acceptable (< timately provide better taxonomic Funding for this project was provid- 0.01% of estimated population sizes) resolution for collected species. A ed by the NOAA Coral Reef Conserva- and that no octocoral species is over- potential benefit to collectors who re- tion Program’s Cooperative Agreement collected. The scaling of quotas and port at higher taxonomic resolution with the South Atlantic Fishery Man- landings at finer taxonomic resolution would be that they could document agement Council (2011–13). The au- may be desirable to provide further that their collections are eco-friendly, thors especially thank J. Schull (NOAA data resolution. Further, the collectors by linking scientific results presented Fisheries, SEFSC-Miami Laboratory) interviewed rotate their collection sites here that indicate collecting represents and A. David (NOAA Fisheries, SEF- and are selective about removing indi- an insignificant percentage of the to- SC-Panama City Laboratory) for proj- vidual colonies that are size-appropri- tal population. None of these sugges- ect guidance and support. M. Bademan, ate and aesthetically pleasing for the tions related to recording additional or N. Sheridan, and V. Brinkhuis with the aquarium trade. As long as octocoral better information as part of the Trip Florida Wildlife Research Institute, collection does not increase substan- Ticket program are needed to benefit Fish and Wildlife Conservation Com- tially (e.g., 500 to 1,000 times current or protect the fishery itself. Instead, mission, provided helpful reviews. The

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