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Home , Acanella, Acanthogorgiidae, Alcyonacea, Anomocora, Antipathes, Antipathes dichotoma

STATE OF DEEP ECOSYSTEMS IN THE PACIFIC REGION

STATE OF DEEP CORAL ECOSYSTEMS IN THE U.S. PACIFIC ISLANDS REGION: AND THE U.S. PACIFIC TERRITORIES

Frank A. Parrish1 and Amy R. Baco2

I. INTRODUCTION While trace coral samples from anecdotal dredging and suggest a wide distribution The U.S. Pacific Islands Region consists of of deep throughout the Pacific, the only more than 50 oceanic islands, including two detailed assessment of deep corals within the U.S. (Hawaii and ), parts waters of the Pacific has been in the Hawaiian of four other archipelagos (, , . Antipatharians were first reported , and ), and from Hawaiian waters more than 75 ago numerous in proximity to each of these (Verrill 1928). The earliest descriptions of deep groups. These islands include the State of Hawaii, octocorals in Hawaii are recorded by Dana (1846), the Commonwealth of the Northern Mariana with Nutting (1908) reporting 68 . Other Islands (CNMI), and the territories of and significant contributions to the species lists of this ISLANDS PACIFIC , as well as nine sovereign region include Muzik (1979) and Grigg and Bayer Federal territories—Midway , , (1976) for octocorals, as well as Vaughan (1907) , , Jarvis , and (1984, 2006) for scleractinians. Wells , , Atoll, and (1954) provides data on the Marshall Islands. ). This also encompasses the Pacific deep coral research has expanded greatly Pacific Island States in free association with the over the last four decades, primarily as a result (former U.S. trust territories also of the establishment of commercial for known as the Freely Associated States) including black, pink, and coral off the main Hawaiian the Republic of , the Federated States Islands, and subsequent development of of (Chuuk, Pohnpei, , and management plans for these resources by the Yap), and the Republic of the Marshall Islands. State of Hawaii and the Western Pacific Fishery This region includes some of the most remote, Management Council. Deep corals are harvested unpopulated islands in the Pacific, as well as as raw material for the jewelry trade. The coral many densely populated islands, and it extends supports a portion of a $70 million Hawaii-based from the South Pacific (e.g., American Samoa; industry that employs roughly 650 people in its 14º S latitude) to the North Pacific ( 28º manufacturing facility and 50 retail stores (Carl N latitude) (Figure 4.1). The punctuated of Marsh— Divers pers. comm.) the Pacific Region distinguishes deepwater coral communities biogeographically and ecologically Commercial beds of were first from other areas in the United States. Because discovered at a depth of 30-75 m off Lahaina, of the isolated of these islands (especially Maui in 1958. Some of the earliest ecological Hawaii and the Northwestern ), work on black corals was carried out in the 1960s they possess some of the highest levels of marine in the channel waters off Maui using SCUBA endemism recorded anywhere on earth. (Grigg 1965). The Maui black coral bed has remained the focus of coral harvesters throughout 1Pacific Islands Fisheries Science Center, the fishery’s history and has been periodically NOAA resurveyed over the last 40 years. These studies have provided the longest data sets available 2570 Dole St. , HI 96822 worldwide on the status and trends of black coral populations and the effects of the commercial 2 Woods Hole Oceanographic Institution fishery and other natural and anthropogenic Biology Department stressors. In the mid 1960s, isolated patches MS#33, 250 Redfield of pink (Corallium spp.), gold (Gerardia sp.) Woods Hole, MA 02543 and bamboo ( sp., sp.) corals

155 156 PACIFIC ISLANDS equipped with a multibeam bottom mapper and mapper bottom multibeam a with equipped and ( submersibles 3-person deep-diving has Undersea the facility replacing expanded, NOAA’ssince The of (NURP). Program node Research established an (HURL), Laboratory Research Undersea Hawaii and became the centerpiece of the newly formed the renamed was submersible the 1980 In 1979. and 1974 between fishery the for coral 1993) the sub was leased to commercially harvest (Grigg research periodic from Aside Dynamics. submersible person two- the to access was infrastructure research gear. AHawaii’sin coral advancement deep key nonselective other and dredges net tangle used Many of the earliest precious surveys of precious on coral beds 1970. in Hawaii of the at began corals focused program research coral deep long-term A1993). (Grigg Bed) (Makapuu north of off and Banks) (Milwaukee Island mdepths Midway 300–500 at identified were the PacificIslandsRegionforNationalMarineFisheriesService. Figure 4.1. REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF ics V Pisces MapofthePacificBasinshowingU.S.islandsandtheirExclusiveEconomicZonethatcomprise ad ddctd upr vessel support dedicated a and ) ep tr 2 Star Deep Makalii rm General from ics IV Pisces with two, with Makalii

r te esrs ht ae en mlyd to employed been have that measures the are targets for the commercial fishery. Also discussed are that taxa coral the on focused is work this of Much corals. deep of dynamics and distribution the on and organisms, other and corals between the region’s of coral community, composition ecological relationships genetic and taxonomic the the on focused from have Studies conducted. been have are corals deep Hawaiian Archipelago, where most the of the other surveys of and Most corals, precious corals, black included. on available be information will corals black shallower However, m. 50 than deeper corals on focus mostly will chapter Pacific the report, national the within corals this of intent the with keeping In Region. Islands is deep what about of summary known a provides chapter This and research coral of increased participationbymoreresearchers. focus infrastructure the new This expanded 1998). Malahoff and ( vehicle operated remotely a RCV-150 (Chave ) STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION protect deep coral ecosystems and to manage Equatorial Current and the South Equatorial the commercial fishery. Current and travels eastward at slightly more than 2 km h-1 (Thurman 1981). The boundaries of these water masses overlap and contribute II. GEOLOGICAL SETTING to long-distance dispersal of pelagic larvae, particularly in the western Pacific. The northern The U.S. Pacific Islands lack the shelf area that Hawaiian seamounts fall in the northern portions typically defines the deep- benthic of the north gyre; Hawaii, Wake and Johnston are of the continental United States. Instead, the in the center of the ; Kingman submerged slopes of volcanic pinnacles that rise and Palmyra are in the equatorial/eastern Pacific; steeply from abyssal depths of 4–7 km provide Jarvis, Howland, and Baker are in the equatorial the hard substratum that deep corals colonize. system; American Samoa is in the equatorial The region has endured a long history of plate portion of the southern gyre; and the Mariana drift, subsidence, and rise, and many Islands are affected by the north central gyre, of the volcanic islands have drowned creating the equatorial systems, and the Kuroshio current numerous submerged banks and seamounts. A from Asia. striking feature of the Pacific Basin is the linear nature of the island chains and seamounts. They While deep water masses originate from surface are aligned in a north-northwesterly direction, currents, no deep water masses form in the a consequence of the northwesterly drift of the Pacific Basin. Deep water migrates in from PACIFIC ISLANDS PACIFIC Pacific plate over stationary volcanic hotspots the Atlantic the with a (Kennett 1982). The resulting islands and uniform temperature and salinity below about seamounts are progressively older in proportion 2000 m (Knauss 1996). The deep water flows to distance from a . For example, the northward at depths below 2500 m and southward island of Hawaii lies above the plume and above 2500 m. Seamounts, pinnacles, and other is the only island in the Hawaiian Archipelago that structures obstruct current flow and can generate is volcanically active. To the northwest, eddies of varying intensity, depending on the on Oahu ceased about three million years ago; current velocity, depth or of the , about five million years ago; and Midway slope of the side, and strength of the Island about 27 million years ago (Grigg 1988a). stratification. Both cold and warm water eddies This geologic process defines the Pacific plate are formed as a result of a seamount obstructing and, as a result, the Pacific Islands region has current flow in the deep . Typically, some of the youngest (main Hawaiian Islands) anticyclonic (cold water) eddies are formed above and oldest (Line Islands) volcanic archipelagos the seamount and remain tightly associated in the world. with the top of the seamount, while the cyclonic (warm water) is formed downstream behind the seamount (Kamenkovich et al. III. OCEANOGRAPHIC SETTING 1986). Deep corals are thought to benefit from the flow acceleration, larval retention, and high The Pacific is composed of two large gyres, nutrient waters from deep caused by the northern and southern central gyres. In the the presence of the seamount and the generated South Pacific, southeast drive the eddies (Genin et al. 1986; Mullineaux and Mills South Equatorial Current westerly between 15o 1997; Coutis and Middleton 2002). S and 3o N latitude. Within the South Equatorial Current is the Cromwell Current, or Pacific Oxygen in the deep water of the Pacific has been Equatorial Undercurrent. This current exists at depleted to very low levels as a result of the length depths of 70–200 m, and is approximately 450 of time it takes for water to move into and across km wide and flows with velocities of up to 5 km -1h the Pacific Basin. Having aged thousands of for a distance of up to 13,000 km in the opposite years, oxygen averages 0.5–4.5 ml l-1 versus the direction of the South Equatorial Current (Tchernia significantly higher Atlantic average of 3.0–6.5 1980; Thurman 1981). In the North Pacific, the ml l-1 (Thurman 1981). The effect of low oxygen North Equatorial Current flows westward at 1 km levels on deep corals is poorly documented; h-1 between 8o and 20o N latitude. The Equatorial however, Wishner et al. (1990) attributed patterns Counter Current is located between the North in the abundance and distribution of ,

157 158 PACIFIC ISLANDS 4.1 deep of records Published explored. been yet waters around other U.S Pacific Islands have not deep because largely Archipelago Hawaiian the around However, only identified been region. have species Pacific most U.S. the in exist to known are groups coral deep major the of Most IV. specific patternsinoxygenlevels. depth- to organisms benthic other and pens, sea REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF water Isidids indeeper laauense Corallium secundum Corallium beds in preciouscoral antipatharians octocorals and Other deeper water antipatharians in octocorals and Other antipatharians Shallower Gerardia scleractinians Other rostrata Enallopsammia DEEP CORALS STRUCTURE AND HABITAT-FORMING Taxa

sp. Structure-formingattributesofdeepcoralsinHawaii. Size (widthorheight) Relative Abundance Spatial Overall Rating Reef-Building Associations Morphology Attribute Building Reef- No No No No No No No No No Abundance Medium Medium High High High High High Low Low Maximum Medium Medium Medium Medium Colony Large Large Small -Low Size Med Med Small (<30cm)/Medium(30cm-1m)/ Large(>1m) Table Key Morphology Branching Branching Branching Branching Branching Branching Branching Branching Branching Non- e seis f cooas to e seis of species new two octocorals, of species new three least at collected cruise Islands Hawaiian 1 main 2004 a and addition, In species. zoanthid 2005), new (Cairns species stylasterid new three antipatharians, of species new several and identified eight new species of octocorals, two new genera data) unpublished Hawaiian Baco, Northwestern (A. the Islands in cruise 2003 A appendix). chapter See 2006, (Cairns Hawaii to endemic be to thought scleractinians the of 21% with scleractinians, azooxanthellate of species than 137 species of gorgonian octocorals and 63 corals from the Hawaiian Archipelago include more None/ Few(1-2)/Many(>2) Branching/ Non-branching Low/ Medium/ High Low/ Medium/High Solitary/ Clumped Associations Measure Yes/No with Other Structure- Forming Many Many Many Many Many Many Many Many Few Dispersion Clumped Clumped Clumped Clumped Clumped Clumped Clumped Clumped Colony Solitary Spatial Importance Structural Rating of Medium Medium Medium Medium Overall High High High High Low STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION antipatharians, and provided range extensions for Hawaii can be found in Cairns (2006) and is for several genera and species of corals that also included in the Appendix to this chapter. were not previously known from Hawaii. Thus, although an extensive species list exists for the b. Black corals (Class , Hawaiian Archipelago, the high rate of discovery Antipatharia) of new species and new records implies the Fourteen genera of black corals are reported from Archipelago is also largely undersampled. the Hawaii-Pacific region with species found in both shallow and deep habitats. The shallowest In general, the deep corals in the Pacific Islands genera (Cirripathes spp. and spp.) do not form the extensive reef structures observed prefer shaded or low light areas and can occur in the Atlantic and South Pacific. Instead, corals underneath ledges and in in shallow water grow attached directly to the exposed (e.g., anguina can occur in 4 m depth) carbonate, or substratum. where surge is minimal, or in the open on steep Octocorals and antipatharians have been found walls at deeper depths. Antipathes spp. appears to grow in high densities at numerous sites, to settle predominantly in depressions, cracks or particularly on summit areas of seamounts or other rugged features along steep ledges, with other topographic highs, where they often form few colonies found on smooth basaltic substratum extensive coral gardens or “beds” with abundant (Grigg 1965). Shallower antipatharians in associated invertebrates. The coral taxa listed in Hawaii also appear to prefer substrates that are this chapter are those that present a conspicuous encrusted with calcium carbonate from coralline PACIFIC ISLANDS PACIFIC relief profile that could serve as a source of , bryozoans, and corals. The highest habitat (Table 4.1). At death, these taxa decay densities are found on hard sloping substratum, in from physical and bioerosion forces until they areas with 0.5–2 knot currents (Grigg 1965). The detach from the substratum and are swept away. best studied black corals are the commercially Cemented reefs from accumulated dead material harvested species and have not been observed. Antipathes grandis. Recent taxonomic work (D. Opresko pers. comm.) on the Hawaiian A. a. Stony corals (Class Anthozoa, Order dichotoma suggests it is a new species and is ) currently being referred to as Antipathes cf. Enallopsammia rostrata is an arborescent curvata. The A. cf. curvata and A. grandis exhibit scleractinian coral in the Family Dendrophyllidae. similar growth rates (6.42 cm yr-1 and 6.12 cm yr-1, The full depth range for this species is listed as respectively) and reach reproductive maturity at 229–2165 m in Cairns (1984), but it has been ages 12–13. Fertilization takes place externally observed in Hawaii primarily at depths of 500– in the , and light and temperature 600 m. In some areas it forms bushy colonies, influence larval settlement patterns. The larvae with dead coral accumulating near the base of A. cf. curvata and A. grandis are negatively of the colony much like that observed among phototactic, and the lower depth limit coincides Lophelia reefs in the Atlantic. Further exploration with the top of the (~ 100 m) in the and characterization of this species is needed to main Hawaiian Islands (Grigg 1993). determine its abundance throughout the region and its potential role in forming habitat. Much less is known about deeper genera of black corals. They have similar morphologies to kauaiensis and M. oculata also the shallower corals, including colonies shaped occur in Hawaii and have the potential to form as whip-like filaments Stichopathes ( spp.) and reef structures, however, little is known of their as branching, sometimes “feathery” colonies abundance or distribution in the Archipelago. (Myriopathes, Bathypathes spp., Stauropathes Besides these examples, scleractinians that and ). However, the life history of have been observed are primarily solitary cup these deeper genera is likely to be much different corals. They can occur in abundance, e.g., on than their shallower relatives. Radiometric dating (A. Baco pers. obs), but many of A. cf. cruvata and Leiopathes indicates the species are small and not observable with a deeper Leiopathes genera grow 10 to 70 times submersible, preventing a true determination of slower than the shallower A. cf. curvata (Roark their distribution. A complete species list (to date) et al. 2006).

159 160 PACIFIC ISLANDS sp. inthe Atlantic (Druffel et al.1995). consistent on is aging radiometric from findings with which 2006), al. et 2700 (Roark and years 450 between samples coral gold of the Hawaiian species has estimated the life span on work radiometric Recent 2002). (Grigg years have provided an estimated lifespan of around 40 growth bands assumed to be annual in periodicity of span life The cause. other some or urchins predatory by decorticated been has coral host the of portion a after spreads and it colonizes if opportunistically or host the of tissue living the outcompete if known not is It completely. zoanthids eventually replace the host colony of as case the well In as bottoms. hard to species coral observed living been other colonize have Hawaii in Zoanthids of 350–600m. depths at Chain Seamount Emperor the into and Archipelago Hawaiian the throughout distributed m in height (Figure 4.2 B). 2–3 as tall as observed been have colonies and an arborescent morphology similar to gorgonians, observed has It species. these of largest the certainly and been have coral, species six and,ollected (Baco, unpublished data). The gold least at from deep water in described Hawaii been although yet taxonomically,has zoanthid of species No Zoanthidea) c. Goldcoral REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF Gerardia A— Figure 4.2. Corallium Gerardia (Class Anthozoa, Order sp., is probably the most common Photos ofthetwoprimaryGenerathatcompriseHawaiianpreciouscoralfishery sp.andB— sp. is uncertain. Counts of Counts uncertain. is sp. Gerardia Gerardia Gerardia Gerardia sp. is widely sp.PhotocreditF. Parrish,NOAA Fisheries. Gerardia sp., the sp., sp. can sp.

the 2005 Line Island surveys and it was present was it and surveys Island Line 2005 the Gold coral was one of the few corals seen during press) and are estimated to reach reproductive reach to estimated are and press) in Baco and Waller 1993; (Grigg gonochoristic in beds. co-occuring the scleractinians sometimes and with an abundance of other octocorals, zoanthids, invertebrates of diversity high a support usually (Figure and beds large in considerably found often are They vary A). 4.2 can “fan” colony’s the of shape and height the and shades, color height in cm the 30 that more to Growing 1956). (Bayer Chain data) Seamount Emperor unpublished the into Baco, and 1993; 1974, (Grigg seamounts throughout the Hawaiian Archipelago and islands on m 350–600 of depths at occur to and (red coral; formely identified as most Hawaiian the species, the Two inArchipelago. far corals diverse by and abundant are octocorals Gorgonian Gorgonacea) d. Gorgonians region (FrankParrishpers.obs.). the in abundance low in been always has coral gold suggesting colonies, fallen of accumulation no found attached were colonies gold individual the where spot the cliffsthe below of base the at were infested colonies with All large other “bed.” unidentified a patches zoanthids. called be no Surveys to enough with sparse, were colonies the However, Kingman. and Palymra, Jarvis, at Corallium Corallium secundum Corallium . secundum C. spp. occur in a variety of red/pink of variety a in occur spp. (Class Anthozoa, Order and (pink coral) are known are coral) (pink oalu laauense Corallium . laauense C. Corallium regale are ) STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION maturity at 12–13 years (Grigg 1993). These these species in this region. A complete species species are relatively long lived, with some of the list (to date) is included in the Appendix to this oldest colonies observed within Makapuu Bed chapter. about 0.7 m in height and approximately 80 years old (Grigg 1988b, Roark 2006). Populations of g. Stylasterids (Class , C. secundum appear to be recruitment limited, Order Anthoathecatae) although in favorable environments (e.g., Four species of stylaserids are present in Makapuu Bed) populations are relatively stable, Hawaii, but they are distributed very patchily suggesting that recruitment and mortality are in a throughout the Archipelago. An extreme example steady state (Grigg 1993). is Disticophora anceps. It has a very wide depth range but has only been found on the northwest More than 130 other species of octocorals are slope of Island in densities of several known from the Archipelago and they represent colonies per square meter in some areas (Cairns a diverse array of families and genera. Most 2005; A. Baco unpublished data). Again, little abundant are the Families , Isididae, else is known about the biology and ecology of , , and this group in this region. . To our knowledge, besides the species discussed above, there isn’t any Hawaiian stylasterids are discussed in Cairns information on the biology and ecology of these (2005) and a complete species list (to date) is groups in this region. Further discussion of their included in the Appendix to this chapter. PACIFIC ISLANDS PACIFIC depth distributions are in the following sections and a complete species list (to date) is included in the Appendix to this chapter. V. SPATIAL DISTRIBUTION OF CORAL SPECIES AND HABITAT e. True soft corals (Class Anthozoa, Order ) General distribution The Alcyonacea are represented in this region by Our knowledge of the spatial distribution of deep only 12 species in three families. Of these, the corals in the U.S. Insular Pacific is limited to is the most widely distributed. Hawaii. Even in Hawaii, very little of the deep It is often observed in beds in sea has been explored and every research patches with large number of small individuals expedition is yielding large numbers of new surrounding a larger individual (A. Baco pers. species. Until 2003, the majority of studies in obs.). The bright purple grandiflora Hawaii came from sparse trawl data or had has been observed growing on Gerardia at a concentrated on the harvested black, gold, and number of sites throughout the Archipelago (A. pink corals. Often referred to as “precious corals,” Baco pers. obs.) these are the primary deep coral taxa harvested for the jewelry trade. Most of these are found Like the gorgonians, little else is known about between depths of 300 and 500 m and have the biology and ecology of these species in been collected by dredge or submersible. Often, this region. A complete species list (to date) is black coral (Antipathes spp.) is distinguished included in the Appendix to this chapter. from the rest of the precious corals because the Antipathes taxa that are used for jewelry occur f. Pennatulaceans (Class Anthozoa, Order much shallower (<100 m) and are harvested Pennatulacea) by scuba divers. Beside precious corals, many Pennatulaceans seen in Hawaii tend to be more other taxa of deep corals have not been studied abundant in areas high in , although because they were not one of the management Kophobelemnon sp. has occasionally been unit species of the precious coral fishery. In 2003 observed on adjacent hard bottoms near the and 2004, the Pisces submersibles were used Cross Seamount deep coral bed. Near the for the first studies of non-precious corals, thus Keahole deep coral bed on the island of Hawaii, extending the exploration of corals well below a single species (as yet unidentified) occurs in previously surveyed depths. In 2005, the first high abundance in patches of sediment at depths surveys conducted outside of Hawaii using the of about 400 m (A. Baco pers. obs.). Again, little Pisces submersibles were at Rose Atoll and the else is known about the biology and ecology of U.S Line Islands (Jarvis, Palmyra, and Kingman).

161 162 PACIFIC ISLANDS Islands at depths of about 30–110 m. The largest Black coral beds are found off the main Hawaiian Black coralbeds flow for corals for flow thelaminar different (e.g., for regimes flow preferred reflect may differences habitat These press). in relief areas on pinnacles, walls, and cliffs (Parrish relief of outcrops; and laauense Corallium pavement; carbonate over flow high of areas in secundum example, For taxa. coral among differ to likely is growth and colonization for suitability environmental The etc.). is load, particulate of (flow, presence there the certainly premium substrate and environmental conditions with and of associated taxa patchiness distribution coral the different influences clearly Depth U.S. Pacificisunknown. the of rest the in corals deep of distribution The Inset showsadiverconductingcoralsurvey. Mapcreidt:F. Parrish,NOAA Fisheries. Figure 4.3. REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF Gerardia Topographic mapofthemainHawaiianIslandswiththreeknownblackcoralbedsmarked. per t go i lre numbers large in grow to appears sp.). rw i a intermediate an in grows Gerardia C. secundum C. sp. grows in high , alternating , Corallium end of the chain. Other species of black corals black of species Other chain. the of end shoaling of the thermocline towards the northwest Hawaiian Islands (Grigg 1974), perhaps due Northwestern (see the to in the found been has coral black 1977) Eldridge chapter appendix). and Little commercially important (Devaney depths shallower from absent antipatharians of species other with along Islands, Hawaiian main the off ulex kg. 40,000 contained Kauai at bed m, to be 166,000 kg or 84,000 colonies, while the 70 and 40 between area, Channel Auau the for of crop standing total a estimated by followed are locations these in found species dominant The southern end of the island of Hawaii (Figure 4.3). Gig t l 20) salr lc crl bed coral km (0.4 black smaller A 2002). al. et m (Grigg 30-90 of depths at bridge islands two land the between drowned a encrusting , and in the middle of the Auau Channel, between Maui bed covers an estimated area of 1.7 km Antipathes s on i dee lctos 1055 m) (110–565 locations deeper in found is 2 ) is located off Kauai and another at the at another and Kauai off located is ) niahs rni. grandis. Antipathes cf. curvata (95% of the population) A. rg (1976) Grigg Myriopathes cf. 2 and lies curvata

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION occur in the Northwestern Hawaiian Islands and and Gerardia) are usually found, but the ratio include taxa such as Cirripathes, Stichopathes, of abundance can vary greatly (Parrish in Stauropathes, Bathypathes, Myriopathes ulex, press)(Figure 4.4). It is difficult to estimate the Trissopathes, Umbellopathes, Dendropathes, size of coral beds, so only relative differences in and Leiopathes. bed size (based on impressions of coral density and the area covered by the submersible track) Between Black and Precious Coral Beds were shown in Figure 4.4 to determine the size of The depth zone between the black coral beds pie diagrams. Most precious coral sites also have and the precious coral beds has had less study. a number of other noncommercial taxa; these Corals have been observed in this zone; for include various octocorals (e.g., , example, an abundance of octocorals occur at Paracalyptrophora, Acanthogorgia, Lepidisis, the Makapuu coral bed on the island of Oahu, , , Kereoides, , and shallower than the precious corals, but they have various paramuriceids) and antipatharians (e.g. not been sufficiently sampled to comment on Leiopathes, Trissopathes, and Bathypathes) (A. diversity or species composition. Baco, unpublished data).

Precious coral beds Of the known coral beds, the Makapuu coral bed Probably the most abundant of Hawaii’s known is the best studied and most diverse. It is located deep corals are the precious corals, including between 375 and 450 m depth in the channel octocorals Corallium laauense (red coral) and between the islands of Oahu and . The PACIFIC ISLANDS PACIFIC Corallium secundum (pink coral), and the bed comprises an area of about 3.6 km2, with zoanthid Gerardia sp. (gold coral). These species the most abundant coral C. secundum, at a are known to occur in significant abundance in mean density of 0.22 colonies per square meter at least 16 locations in the Hawaiian Archipelago between 365 and 400 m (Grigg 1988b). Other at depths of 350–600 m (Grigg 1974, 1993; corals found at Makapuu include Baco, unpublished data) and into the Emperor (Lepidis olapa, 0.041 colonies m-2; Acanella spp., Seamount Chain (Bayer 1956). Within a given 0.001 colonies m-2), gold coral (Gerardia sp., coral bed, the two primary genera (Corallium 0.0005 m-2), as well as three genera of gorgonians

Figure 4.4. Topographic map of the Northwestern Hawaiian Islands with coral survey dive sites of the sub- mersible Pisces V (inset). Pie charts represent the relative amount of coral among sites and the within-site fraction of the two primary precious coral genera. 3D map and Pisces photo credit: Hawaii Undersea Research Laboratory.

163 164 PACIFIC ISLANDS Antipathes. cf. curvata Figure 4.5. many, of distribution depth The Bathypathes. Corallium Paragorgia Acanella, Isidella, Keratoisis, Candidella, Calyptrophora, , Iridigorgia, genera are: octocoral common observed more the Among data). unpublished Baco, (A. appear depth this to around begin antipatharians isidids, and primnoids, coralliids, chrysogorgiids, of species communities, to a different suite of species. Many from m, 600 appears to be a transition in species below about Although there limited, very is observations of number the sites. between sponges and corals both of composition species in occur to appears as ridges and pinnacles, a fair amount of overlap such areas current high in depths deeper these At Point. Keahole and Seamount, Necker Cross Island, of east seamount unnamed an Island, Laysan of southeast seamount small the Ridge, Pioneer a depths; coral only precious below at sites few conducted been have Explorations Beyond PreciousCoralBeds undescribed of octocorals. number a and (antipatharian) Paramuriceidae, hilgendorfi including interest are also present in or adjacent to the bed ( gilberti ( REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF Leiopathes Narella ), a ( sp., nlosmi rostrata, Enallopsammia Two speciesofblackcoral trees acanthogorgiids, , a wl a te niahra genus antipatharian the as well as , n. sp. Corallium Psuedothesea ) hyooga Metallogorgia, , rsoahs pseudotristicha Trissopathes . Many taxa of no commercial Stylatula (right).Photo creditF. Parrish,NOAA Fisheries. - and - sp and sp sp.), and black coral Gerardia Paragorgia

Callorgorgia -dominated Thouarella Antipathes grandis and , sp.,

of essential habitat, protected species, and species, protected habitat, fish essential of mandate NOAA the address to conducted been uncertain ecological consequences. Studies have with habitat fishery bottom the to effect direct a is target a as corals deep a of Take of submersible. manipulator the using or divers scuba by using collection hand as such methodologies selective allowed is corals dredging, precious of The harvesting and waters. Pacific U.S. in banned are coral-associated which from to come communities. effectsspecies coral most deep Nationally, species with the on focused associations studies prompted of has number corals a deep of harvesting The V. below 1800m(Baco,unpublisheddata). continue to appears species these of all, not but observations. Even less is known about the deep underwater infrequent making in and ranges depth overlap the evaluations comparing to these limited been However, have targets. fishery other and corals precious between association harvests precious corals, there is little evidence of a direct that fishery the of exception the With Commercial fisheryspecies expanded toincludeinvertebrates. historically have been focused on fish and only These recently have been concerns. ecosystem CORAL COMMUNITIES SPECIES ASSOCIATIONS WITHDEEP (left)and h cmeca bottom commercial of most the but range depth this in lives also snapper), fish commercially sought bottom the of One 2005). Parrish and and Boland 2004; Parrish Boland 1989; al. et (Moffitt be m) (30–100 found can (antipatharians) corals black where depths into range trade aquarium shallow the and fishers recreational by the targeted fish reef coral of Some stocks. to fishery commercial contribute may these of corals any that ecological effects indirect any or fishery, coral precious the in targeted not corals sea pin virescens Aprion (grey STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION fish reside at depths below antipatharians and Heterocarpus sp. is the focus of an intermittent above the precious corals (<300 m) (Uchida main Hawaiian Island trap fishery that targets and Tagami 1984). The shallowest members of depths of 500 to 900 m (Moffitt and Parrish 1992) this group, such as Pristipomoides filamentosus overlapping the lower limit of precious coral (pink snapper), Pristipomoides zonatus (oblique- depths, but in the depth range of many other banded snapper), and Epinephelus quernus deep coral species. (Hawaiian ), have been seen in the vicinity of deeper black coral trees (Moffitt et al. Noncommercial species 1989). Similarly, the deeper members including Fish Etelis carbunculus ( snapper) and Etelis Studies of fish associations with deep corals have coruscans (flame snapper) have been seen focused almost exclusively on the inventory of fish among the shallower precious corals (Kelley et taxa and appraisal of the obligate or facultative al. 2006). The groundfish, Pseudopentaceros roles corals play in fish assemblages. The fish wheeleri (armorhead) and Beryx sp. () community of the Auau Channel black coral bed occur throughout this depth range (250–350) was recently surveyed (Boland and Parrish 2005) but are more common on the seamounts at the and 95% of the fish found in and around the northern end of the Hawaiian Archipelago (Uchida black corals were known to commonly occur on and Tagami 1984). There is no information on shallower reefs where black corals do not grow. the degree of overlap of these fish with (the longnose hawkfish), was corals. Heterocarpus sp. (deep-water shrimp) found exclusively within the black coral trees. PACIFIC ISLANDS PACIFIC has been seen among the precious corals but at Behavioral data indicated that although most of densities consistent with other bottom relief types. the reef fish routinely passed through the coral

Figure 4.6. Examples of invertebrates found associated with deep corals (A-basket star, B-ophiuroid, C-asteroid. Photo credit A.Baco, WHOI.

165 166 PACIFIC ISLANDS u saitcl vlain sgetd ht this that suggested evaluations statistical but density, fish greater supported often corals with Areas species. dominant of couple lownumbers a only in with observed were taxa without. fish those Most and corals with sources. areas between relief abiotic Species richness of fish was not with observed to differ interchangeably seen use were to observed taxa fish 42 the of Many fish 2006). few (Parrish corals precious with associations indicated depths subphotic deeper at communities fish of surveys The 1994). Mundy and (Chave species noncommercial are depths subphotic and slope deep the of fish the of Most movement. deep fish for corridors the improving perhaps and the reef of aspects vertical the enhancing by community fish the to thecontribute colonies coral, coral black and fish between relationship 4.5). Although there is little or (Figure no known obligate threat a evading the when cover used for corals reliably species four only branches, REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF Gerardia and Corallium p. s shelter as spp. Invertebrates coral colonies. the on rely stages juvenile if or ecology fish the if the corals play some seasonal or diurnal role in unknown is It fish. adult on exclusively focused and day the during summer, in conducted were appeared fish the by incidental, it is corals important to consider these of studies use Although frequently used most that community fish the hoverers” of segment the as “benthic revealed groups functional two the than taller significantly is it because probably species, fish oriented mostly around coral commercial three the Of beds. coral the of were not seen in the mean size of differences Also, fish in corals. or on fish outside of dependency a on based not and flow high and relief of areas in coral and fish of co-occurrence on based was soitd ih ep oas Bak oas are corals Black species corals. deep with of associated array wide a is There Corallium species. Classifying the fish into fish the Classifying species. Gerardia sp.asshelter. A. Baco,WHOI. colony.Photo credit on a diflora Clavularia gran purple octocoral Figure 4.7. Gerardia growing Gerardia The sp. - sp. STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION known to be colonized by oysters, bryozoans, A more generalized commensal is the unbranched and shrimp (Hoover 1998). The most common basket star, Asteroschema. Asteroschema sp. invertebrates associated with the deeper precious has been observed in a number of different species corals include zoanthids, anemones, galathaeoid of octocorals at a number of sites, but not on the , sponges, ophiuroids, and basket stars surrounding substratum. Dead coral skeletons (Figure 4.6). Gerardia sp. is perhaps the best also appear to provide good recruitment habitat known coral that overgrows the skeletons of for many invertebrate species. Many types of other coral species. Examination of Gerardia sp. sessile fauna have been observed as well as skeletons and many submersible observations several types of young corals. In particular, young suggests it has an obligate need to start its colonies of the red coral, Corallium laauense colony over the skeleton of other corals. Bamboo have been observed growing on dead skeletons corals seem to be the most frequent target, of Gerardia sp. particularly Isidella trichotoma, but Gerardia has been observed growing on quite a number of Besides the galathaeoid crabs that inhabit the species. Several other species of unidentified branches of coral colonies, a number of larger zoanthids also appear to prefer to grow on other crabs are routinely encountered patrolling the corals, including Gerardia sp. At least one of bottom around deep corals (e.g., crabs in the these zoanthids was observed growing on basalt families , ). substrate as well as encrusting other corals (A. Baco pers. obs.). Octocorals also can grow on deep corals by resident invertebrates PACIFIC ISLANDS PACIFIC on the skeletons of other coral species. The also occurs. Seastars feed on coral colonies bright purple Clavularia grandiflora has been by everting their gut, leaving behind patches observed growing on Gerardia at a number of of bare coral skeleton. Cidarid urchins are sites throughout the Archipelago (Figure 4.7). also known to feed on deep corals and these In all of these cases, it is not clear whether the urchins have been observed on deep corals overgrowing corals are actually killing or injuring in Hawaii. However, an absence of bare coral the coral whose skeleton they are growing on, or skeleton around the urchin’s location suggests it if they have simply colonized a dead area of the is unlikely they are consuming the coral tissues skeleton. (A. Baco, unpublished data). A single species of crinoid was observed in 2004 in very high Many coral colonies have a number of commensal densities at the Makapuu coral bed (A. Baco, invertebrates such as galathaeoid crabs, unpublished data). Grigg 2002 comments on an barnacles, sponges, anemones, , abundant crinoid observed during night dives at crinoids, ophiuroids, and basket stars (Chave and this site. The crinoids now cover many octocoral Malahoff 1998; A. Baco and Shirley, unpublished colonies and anything else that sticks up more data). Casual observations suggest some of these than a few millimeters off the bottom (A. Baco, commensals may be specific to their host while unpublished data), suggesting their abundance others are more generalized. The relationships has increased over Grigg’s observations. Their between the coral and the commensal, e.g., origin and the reason for their recent increase in symbiotic, parasitic, obligate, facultative, etc., population density, as well as their potential for have not been determined in most cases and competing with corals for food, are unknown. many of the commensal species have yet to be identified. One of the more notable commensal Monk seals relationships is the general association of In the lower Northwestern Hawaiian Islands, polychaetes with species in the genus Corallium. the endangered Hawaiian has been Each Corallium species appears to have its own documented to routinely visit deep corals as species of polynoid . In Corallium part of its foraging activities. However, there secundum and Corallium laauense, these have been no reported interactions between polychaetes can reach fairly high densities. The monk seals and the precious coral harvesting polychaetes live in tunnels under the coral soft in the main Hawaiian Islands. Telemetry and tissue with the skeleton often growing over the scatological analysis indicate seals prey on polychaete tunnels. This relationship has also bottom-dwelling fish (Goodman-Lowe 1998; been observed in other corals such as Candidella Parrish and Abernathy 2006; Longnecker et helminthophora. al. 2006). Video cameras fitted to seals in the

167 168 PACIFIC ISLANDS a b afce b a ubr f aua and has natural mortality Natural of stressors. number anthropogenic a by affected be may Deep coral communities within the Pacific region VI. holes andcracksofthebottom. and into look to sub the from field light the using then the submersible over the looking shows briefly encounter seals the of video The pers. obs.). Baco (A. surveys coral conducting were scientists while m 500 than more at submersible seals have also been observed from the Pisces V Monk 2006). al. et (Stewart depths subphotic to patterns feeding similar shows Islands Hawaiian of seals at the northern extent of the Northwestern (Parrish et al. corals 2002). gold and More red recent satellite revealed tagging have with surveys where locations subphotic specific at focused foraging their of weeks spent seals eels. certain indicated seals on to attached tags Satellite feed they where m) of (100 corals beds whip to commuting seals recorded have Islands Hawaiian Northwestern Squirrelfish snapper, HawaiianGrouper, Ruby-coloredsnapper, Blue-greensnapperfish. of species two include: shrimp Deepwater . Pomfret, Opah, Dolphinfish, Wahoo, spearfish, Shortbill Sailfish, Marlin Black Marlin, Pacific Blue Marlin, Striped Swordfish, , Skipjack Tuna, , Bigeye Albacore, includes: Swordfish Table4.2 REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF Mid-water Trawl Pelagic longline Hook andline Bottom Trawl Bottom-set Bottom-set Gear Type COMMUNITIES STRESSORS ONDEEP CORAL Longline Dredge Gillnet Traps Potential effects of gears on deep coral habitat in Hawaii. Ratings detailed in table key. table in detailed Ratings Hawaii. in habitat coral deep on gears effectsfishing Potential of Tuna/ Deepwater Snappers/ swordfish Current Fishery Region Shrimp Use in Tuna/ N/A N/A N/A N/A N/A Heterocarpus, Cirripathes of Impact Potential Severity High High Med Med Med Low Low Low Snappers and Groupers include: Pink snapper, Flower snapper, Flower snapper, Pink include: Groupers and Snappers sp. Extent of Potential Fishing Impact Gear from High Med Med Med Low Low Low Low emut Clhn emut ad oe of Hawaiian the of end north the some at seamounts the and Seamount) Colahan Seamount, ofthe end Emperor Chain the south (e.g., Coco at Seamount, Milwaukee seamounts the fished From 1967 Region.to Islands 1975, Soviet Pacific and Japanese U.S. trawlers the in banned are Mobile bottom-tending gear (e.g., trawls, dredges) Fishing effects 1993). (Grigg lost be can growth coral accumulated decades of gear, damaging bottom using fishing indiscriminate of periods intensive During time. same the at effects to exposed are classes damage Many 4.2). (Table fisheries coral habitat unmanaged in to vulnerable associated with fishing gear and overexploitation highly them makes corals deep of attributes history life The to able rarely reattach. are colonies Detached 1993). (Grigg colonies of toppling to leads which site, attachment the at substrata the of bioerosion by and by smothering to attributed been Geographic of Usein Current Region Extent High Med Low Low N/A N/A N/A N/A with coral Overlap habitat of use Low Low Low N/A N/A N/A N/A N/A of Gear Overall Impact Rating Low Low Low N/A N/A N/A N/A N/A STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

Archipelago, including seamounts that were later has landed 680 metric tons since the fishery’s designated as part of the U.S. exclusive economic inception in 1984 (PIFSC IR-06-010). The shrimp zone (e.g., , Seamount 10, trapping overlaps the depth range of deep corals Seamount 11, Ladd Seamount) (Uchida and (Ralston and Tagami 1992; Moffitt and Parrish Tagami 1984). The primary fishing target was 1992), but actual impacts to deep corals have not Pseudopentaceros wheeleri (armorhead) and been documented. Shrimp have been observed Beryx spp. ()(Uchida and Tagami 1984). associated with hard bottom features (Moffitt and In 2003, submersible dives at Seamount 11 Parrish 1992) and if fishers seek hard bottom to found a large area with coral stumps and no new set their traps, there is potential for damage to colonization (A. Baco pers. obs). The destruction deep corals. In Table 4.2, the overall gear impact was confined to a large swath within a fairly large rating of shrimp trapping was classified “Low” precious coral bed (previously unknown). It could because of the small size and localized nature not be determined if this affected area was the of the fishery. However if the fishery expands result of mobile bottom-tending gear used in the the potential impacts to deep corals would be early 1970s but if so, recovery clearly requires an important consideration. There are no other decades. recognized trap fisheries operating in the U.S. Western Pacific Islands and if any recreational Longline fishing or artesianal trapping is happening, it is at a very Bottom longlining is not permitted in the Pacific small scale and in shallow depths. Islands Region. Pelagic longlining for tuna and PACIFIC ISLANDS PACIFIC swordfish is permitted and is the region’s largest Other fishery. Longlines must be set at least 25 miles, Fishing for reef species and bottom fish typically and in most cases 50–75 miles from emergent rely on spearing and hook and line fishing. parts of the Hawaiian Archipelago (WPFMC Spearfishing is largely constrained to the 1991). This regulation was adopted to prevent shallowest depths and is unlikely to have an effect conflicts with the coastal trolling fishery butit on deep corals. Corals might be damaged by also reduces the possibility that the gear will the 3-kg bottom weight used to lower handlines affect deep corals on the slopes and seamounts for bottom fishing or might be snagged by the of the Hawaiian Ridge. One exception is attached hooks. However, visual surveys from Cross Seamount located ~ 100 miles south of submersibles have inspected popular bottom Oahu. A popular fishing site for monofilament fishing sites in the main and Northwestern handline fishing and some longline activity, it Hawaiian Islands for fishing impacts and have has accumulated numerous large fragments of found little or no derelict gear (Kelley et al. monofilament line draped over the summit (F. 2006), and there are no reports of coral bycatch Parrish pers. obs). Some of these line fragments (WPFMC 2005). have been seen entangled in Gerardia sp. colonies (A. Baco pers. obs.) and other coral Effects of other human activities trees appear to have been damaged. As this is the only location that impacts to deep coral from Coral Harvesting monofilament fishing have been documented the The commercial harvest of coral is the best assigned impact rating is “Low.” documented effect to black corals and precious corals (pink, red, and gold) within the Hawaiian Traps Archipelago. Coral harvesting has been subject Bottom-set traps have been used to catch lobster to management under both federal and state and shrimp in the Hawaiian Archipelago. The regulations since the 1980s. Commercial harvest Northwestern Hawaiian Islands trap fishery for of black coral has always been selective, collected Panulirus marginatus (Hawaiian spiny lobster) by scuba divers using hand tools. The deeper and Scyllarides squammosus (slipper lobster) is precious coral beds were fished initially using now closed, but had always operated in waters nonselective tangle net dredges, but regulations shallower than deep coral habitat (Polovina now require the use of selective methodologies 1994, Dinardo and Moffitt 2007). Trap fishing for such as a submersible (Figure 4.8). Commercial the deepwater shrimp Heterocarpus laevigatus harvesting of black and precious corals has not and Heterocarpus ensifer is a small-scale pulse been reported elsewhere in the U.S. Pacific fishery limited to the main Hawaiian Islands that outside of the Hawaiian Archipelago.

169 170 PACIFIC ISLANDS o h aalblt o dtie btyerc maps bathymetric detailed of availability the to due fishing in efficiency improved to attributed (WPRFMC 2006). Much of bins this mt 7-year increase prior has the 22 double been than at more is which bin 7-year 1999-2005 the with to 2005 1985 from increased 7-year landings into showed bins data Aggregating disclosure data. prevents annual of thanthree confidentiality so fewer fishers with operates often fishery This 2004). (Grigg harvesting in increase an by accompanied demand in increase an was there 2002 and 1999 Between 2001). (Grigg 1997 to yr kg 1014 at coral black of harvest and Kauai. State records indicate a mean annual Hawaii off beds other two black in occurred also coral of harvest commercial Limited the present. to continuously levels varying at operating fishery the kept has black and 1960 small in industry the coral established Inc. Divers, Maui credit: American DeepwaterEngineering. WHOI). InsetistheDeepworkersubmersiblewhichwasmostrecentharvestingtoolemployed.Photo Figure 4.8. REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF A derelictcoraldredgelostduringtheearliestdaysoffishery(Dredgephotocredit: A.Baco, -1 from 1981 from tangle nets (Grigg 1993). Maui Divers of Hawaii of Divers Maui 1993). (Grigg nets tangle using bed the dredged fishermen of group small a and Oahu, off Bed Makapuu the in discovered Seamount Emperor 1966, In the Chain. in the Banks on depth Milwaukee m 400 about at discovered was coral pink grade commercial of bed a 1965, In (Grigg 1993). waters and deeper are not considered in to be found of commercial are grade region this in exist curvata. c.f. with >90% of the harvested coral consisting of been commercially harvested in Hawaiian waters grandis Antipathes cf. international trade, only three species ( in reported been have antipatharians of genera 11 Although positioning. GPS of adoption and curvata {formerly Other black coral species known to known species coral black Other Corallium secundum Corallium and niahs dichotoma Antipathes Myriopathes ulex) Myriopathes Antipathes was also was have A. } ,

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION began using a manned submersible to selectively in the Northwestern Hawaiian Islands (Grigg harvest pink, gold, and bamboo coral from this 2002). Because the fishery is made up of a single bed. They removed a total of 8227 kg of pink company, confidentiality prevents reporting of coral and 2097 kg of gold coral between 1973 landings data. However the permitted quota was and 1978 and then discontinued fishing as a not filled at either of the two beds where corals result of high operating costs (Grigg 1993). Pink were harvested. Grigg (2002), working closely corals were also harvested in 1988 from Hancock with the industry, reported removal of 60% of the Seamount using nonselective gear, although only allowed coral quota (1,216 kg) at the Makapuu 450 kg of C. secundum were obtained, most of Bed and 20% (211 kg) at the Keahole Bed. The which were considered poor quality (Grigg 1993). precious coral fishery remains dormant today. In 1978, an undescribed species of Corallium was discovered at Midway Grounds (Emperor Illegal coral dredging Seamounts) at depths of 900–1500 m. This Currently, the threat of illegal coral dredging is resulted in a “coral rush,” with more than 100 thought to be remote. It is included here because boats from and operating in this foreign fishing vessels were documented illegally area. Total yield exceeded 300 metric tons from coral dredging in the remote Northwestern 1979 to 1984 and then dropped off because of Hawaiian Islands in the early 1970s (Grigg resource depletion (Grigg 1993). 1993). Currently, there is no evidence or even rumors of such illegal activity. However, much of In 1999, a Hawaii-based marine salvage and the Pacific region is remote and unpopulated and PACIFIC ISLANDS PACIFIC engineering company bought two deep-worker any such activity could go undetected. Given the submersibles and began commercial harvesting slow growth of deep corals and low recruitment of deep corals at the Makapuu and Keahole coral rates, even brief periods of illegal dredging could beds. Operations targeted pink, red, and gold have lasting effects. corals. Harvesting ended in 2001, when their first coral indicated the price of the coral was too low to make submersible operations cost In 1972, the alien soft coral riisei (Family effective, and potential harvesting grounds in the ) was found in the fouling community Northwestern Hawaiian Islands were eliminated of (Englund 2002). Originally as a result of Presidential Executive Order 13196, thought to have colonized from the tropical which formed the Ecosystem Reserve Atlantic, recent genetic work (Samuel Kahng pers. comm.) suggests it arrived from elsewhere. It has and continues to spread to other suitable areas in Hawaii with high flow and low light (Figure 4.9). In 2001 deepwater surveys of the Auau Channel black coral beds using submarines revealed that more than 50% of the black coral, particularly the deeper, large reproductive colonies, were overgrown and killed by Carijoa (Kahng and Grigg 2005). However, Carijoa was rare on black coral trees in waters shallower than 75 m (Boland and Parrish 2005). Light levels are thought to be too high for Carijoa to colonize the shallower black coral trees. This invasive coral has been identified as a risk to the black coral fishery. Historically, black coral trees that were too deep to be harvested by divers were thought to serve as a de facto reserve for the fishery. With the recent discovery that many of the deep Figure 4.9. The invasive gorgonian octocoral , colonies have been killed by Carijoa, current that infests the deeper black corals of the Auau Channel management practices are being reviewed beds.Photo credit: F. Parrish, NOAA Fisheries. (Grigg 2004). Preliminary surveys of black

171 172 PACIFIC ISLANDS ige em oa mpig fot, multibeam efforts, mapping and beam system) single (GLORIA sonar sidescan earlier multibeam sonar mapping efforts. Supplementing Future coral research will have the benefit ofMapping Research recent coral deep ecosystem. the monitor and growth validate to remeasurement future for colonies marked and thermographs, deployed sites, study established research this all supported that expeditions The work. the continuing in interest expressed have harvesters coral “replanting” and 2002), (Montgomery coral preliminary Some research has been conducted with Hawaiian black harvesting. from stock coral black the to impacts mitigate to means a as discussed been has corals Replanting 2002; 2002). Grigg and Kahng (Grigg stock coral black threat the to the presents on focusing now are Parrish 2006; Baco and Shirley in 2005; prep.). Parrish Studies and Boland fauna 2002; al. other et (Parrish and corals ecological between potential associations and prep.) in Baco 2006; al. et Baco 2005; Shank and (Baco Archipelago Hawaiian the across the taxa coral assessing of work connectivity include to expanded has 1993, research management years, 1988b, recent In 1974, 2001). 1965, manage (Grigg to fishery needs coral the information the 1976), support to Bayer growth and age of estimates and on and assessments focused (Grigg been has research Historically, Management offisheryresourcesandhabitats September 1983(48FR39229). in implemented was plan The 1980). (WPFMC as part of the national network of established fishery councils was Council Management Pacific plan Fishery Western the management when fishery NMFS by first approved the was plan its with The Plan. associated Management Fishery Coral Precious corals deep managing of history 25-year a has Region Islands Pacific The VII. been reported. not has Pacific the in elsewhere corals deep on of occurrence of infestation an Hawaii found of not island have the and Kauai on beds coral REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF RESOURCES AND HABITATS MANAGEMENT OFFISHERY Carijoa or other invasive species invasive other or Carijoa Carijoa aio riisei Carijoa n coral and The .

many Pacific Islands, deep corals also occur corals deep Islands, Pacific many of relief steep the of Because Region. Islands U.S. (EEZ) of the Pacific management plans (FMPs) for the fisheries in the (WPFMC) has responsibility for preparing fishery The Western Pacific Fishery Management Council Fisheries ManagementCouncil corals. deep of abundance colony the surveying directly of means promising more a as Islands Hawaiian main the of beds coral black the on technology survey scan laser-line test to underway currently are Efforts corals. deep of likelihood the infer to bathymetric context fundamental that future coral a surveys will be provide able to use will efforts Miller et al. 2003; Parke and Wang 2005). These HURL; at Smith John by (Products Pacific U.S. the of portions other and the archipelago Hawaiian for and made been have imagery, backscatter detailed including products, M icue poiin fr xlrtr fishing exploratory for the provisions includes but FMP Archipelago Hawaiian be the in can are beds that coral known the corals All bed. of each from harvested theharvesting amount and designate method and western region the within Pacific regulations beds coral its known and classify FMP Coral Precious The Directed Harvest least at within thelast25years. trawling, from impacts spared largely the throughout Thus, been have corals deep and shallow both region, Refuges. Wildlife and is not allowed within the Fish and Atoll are managed by the Department Fish of Defense. U.S. Wildlife the Wildlife Service, while Wake Island and Johnston National by are administered Atoll Refuges Rose and Midway Island, Island, Baker Island, Howland Island, Thesovereign Palmyra Reef, Kingman of territories Jarvis Atoll, waters. their within trawls fish American Samoa all prohibit the use of demersal and CNMI, Guam, Hawaii, of laws territorial and State The 1983. in EEZ Island Pacific U.S. the throughout gillnets bottom-set and longlines, set bottom- trawls, fish fishing demersal of prohibited gears, impacts habitat managing early in an leader WPFMC, regulations. and laws own bytheir of isgoverned EEZs management their within fisheries and countries sovereign are States Associated Freely The regulations. and laws territory fisheries and state by governed be also can and waters, territory and state within STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION in other areas of the U.S. Pacific. The beds are Closed areas classified as: 1) Established Beds, 2) Conditional As noted above, all U.S. State and Federal Beds, 3) Refugia Beds, and 4) Exploratory Permit waters in the Pacific Islands are closed to Areas. Established beds have a history of harvest trawling and dredging—the for which maximum sustainable yield (MSY) has most destructive to deep corals. Additional been determined. Makapuu is the only designated restrictions on fishing and other potentially Established Bed although the Auau black coral harmful activities are in place in the National bed is in the process of being designated as Wildlife Refuges, Papahānaumokuākea Marine established. Conditional beds have MSYs National Monument, and in marine protected estimated based on their perceived size relative areas within state or territory waters. to established beds. There are four conditional beds: , , Brooks Banks The only area that was specifically closed to and 180 Fathom Bank. The WESTPAC Bed is protect deep corals was WESTPAC Bank (located designated as a refugia bed, where no harvest N.W. of ) in the Northwestern Hawaiian is permitted. Exploratory permit areas include Islands. It was set aside by the precious coral four unexplored portions of the EEZ around FMP as a refuge from coral harvesting. Despite Hawaii, Guam and CNMI, American Samoa, some interest, domestic precious coral fishing and all remaining U.S. Island Possessions. The has never occurred in the Northwestern Hawaiian FMP, as amended in 2002, prohibits the use of Islands. On determining that monk seals nonselective gear (e.g., tangle nets, dredges) were visiting precious coral beds, the WPFMC PACIFIC ISLANDS PACIFIC throughout the management area. Black coral proposed expanding the refuge to include areas is primarily found in State waters and the State where seals were visiting. Superseding this move, and the WPFMC jointly manage the resource. the Northwestern Hawaiian Islands Coral Reef Quotas and minimum size limits are monitored Ecosystem Reserve was established in 2001 by through mandatory reporting to NMFS and the Executive Order (No. 13178 and No. 13196) and Hawaii State Division of Aquatic Resources using prohibited most commercial fishing, including coral landing logs and buyer reports. all harvesting of deep corals in the Reserve. In 2006, while the Northwestern Hawaiian Islands Currently, two precious coral issues are Reserve was undergoing the designation process progressing through the WPFMC process. The to become a national marine sanctuary, it was first is reconciling coral lifespan estimates derived proclaimed a national monument by Presidential from radiometry studies (Roark et al. 2006) with Order under the Antiquity Act of 1906 and prior estimates made from the size structure renamed Papahānaumokuākea Marine National distribution of coral colonies and ring counts Monument. Within the main Hawaiian Islands from basal stem cross sections. Of the three and elsewhere in the Pacific, marine protected commercial corals, the black coral (Antipathes cf. areas have not been established specifically for curvata) radiometric estimates were consistent the purpose of protecting deep coral communities. with growth rates estimated from size structure However, there is interest from managers and data (Grigg and Bayer 1976). The radiometric coral harvesters to establish a closed area off life span of pink coral was twice prior estimates, Maui specifically for black coral to serve asa and gold coral (Gerardia sp.) was estimated at reproductive reserve and a biological reference more than an order magnitude longer lived than site. prior growth estimates (Grigg 2002). This has prompted the WPFMC to put a 5-year moratorium Minerals Management Service on the fishing of gold coral until the conflicting Oil or gas exploration does not occur in the lifespan data can be resolved. The second Pacific Islands Region. Historically, some issue is concern that the Maui black coral bed research has focused on the prospect of may be experiencing reduced recruitment (Tony mining manganese nodules that are formed at Montgomery, State of Hawaii, unpublished data; abyssal depths. Recently, interest in -rich WPFMC 2006). This uncertainty combined with manganese mining has resurged and large areas the loss of a portion of the stock to Carijoa riisei of the Pacific , some of which include leaves today’s biomass at least 25% lower than U.S. Pacific Islands and seamounts, are part of assessments in 1976 (Grigg 2004). the potential mining areas (International Seabed Authority www.isa.org.jm/en/seabedarea/default.

173 174 PACIFIC ISLANDS   1) previously sites at gear surveyed usingvisualmethods. such using of focus initial an with stocks coral assess to tools new promising validate to needed is research Also, more the to Region. Islands Pacific U.S. out the of areas remote extend This and continue should infrastructure. submersible and (ROV), vehicles operated remotely mapping, available using achieved been has success Good goal. primary a is found thatare they means where corals determining deep of nature patchy and of fragile inherently The components ecosystems. coral deep ecological and physiological important the determining 2) and corals, deep of inventory species and mapping 1) following: the into divided be can work future for priorities The VIII. high a considered priority. be should impacts mining to determine deep-coral abundance and potential asp). REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF Highestmappingandassessmentpriorities of deepcorals. abundance ,and ranges, species survey deeper habitats to better determine the depth to need a is There m. 1800 below corals none and precious of outside few occurred have corals studies, deep of some assessments baseline had Hawaiian has the Archipelago surveys—Although Deeper deep coral . on theories current to test invaluable an provide and corals, deep work, of systematics and connectivity taxonomy improve to for samples provide material DNA conservation the and provide also will assessments These effort. enforcement more focused permit will resources coral with extensive areas or Documenting mining activities. fishery cobalt-manganese by affected be may that those particularly areas, other unstudied.these Pacific for needed are assessments western Baseline the of rest the Archipelago, Hawaiian and CNMI, Guam, Samoa, leaving American Pacific— the occurred around and has remote in research the most Historically, in Assessments

Further studies of these cobalt-rich regions DEEP CORAL COMMUNITIES UNDERSTAND AND CONSERVE REGIONAL PRIORITIESTO ) hsooia ad clgcl opnns of components ecological and Physiological 2)       the deepcoralecosystem er ao hud e uvyd o sgs of coral recovery. signs for surveyed be should ago 30 years than more harvesting illegal bottom or to trawling subjected were The that recovering. seamounts are taxa coral the not or periodically reassessed to determine whether be should harvesting coral and gear fishing have that been commerciallyharvestedorimpactedby studies—Beds recovery Coral can completeintheirlifetimes. species taxonomists these new than description needing and revision needing corals deep of groups more currently are there and world the in experts coral deep few very are people trained to identify these corals. There of number the increase to and corals, deep of systematics and taxonomy the improve to and ecology studies will be a dedicated effort assessment the of all Taxonomy—Criticalto present threat to the stock of black coral and coral black of stock the to threat present of riisei Carijoa impact ecological The corals. and may be more tightly associated with deep Pacific the of regions remote the in different tobe likely attention, are communities fish Subphotic colonies. greater coral the on dependent be to likely needs particularly the invertebrates, which are more associated ecological their to community corals of associations—The contribution Species other parts ofthePacific. to biogeographically and genetically connected are Pacific U.S. the of seamounts chains and islands the how and them, between and are island/seamount populations within the connected how corals; deep of abilities dispersal reproduction, and history, growth, life recruitment, the about more and connectivity, biogeography—There is population a need to understand history, Life where they mightoccurinunexploredareas. predicting as well as be record climate will a as corals distributions deep evaluating to fundamental coral influence factors that oceanographic the Understanding factors. influenced environmental or biological other is by distribution their that implies in even areas withsimilarsubstrate,relief,anddepth corals, deep of nature patchy The corals— deep for parameters Environmental ersns h ms cer and clear most the represents aio riisei Carijoa — STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

has important implications to the black coral the impacts to deep corals from derelict fishery. Research is needed to identify and handlines/longlines at Cross Seamount understand possible ecological changes to and shrimp trapping in the main Hawaiian the community and develop means to limit Islands should be assessed. the spread of Carijoa riisei to other black coral habitats. Some research should be  Many of the new coral beds that have conducted to determine the feasibility of been identified in the Northwestern remediation efforts for black coral through Hawaiian Islands are protected as part replanting programs. of the Papahānaumokuākea Marine National Monument. Being remote from the anthropogenic influences of the main IX. CONCLUSIONS Hawaiian Islands make them important biological reference sites for future  Deep corals occur throughout the U.S. research. Pacific but only the Hawaiian Archipelago and Line Islands have been the subject of any surveys. Coral habitat is patchy, suggesting at least a basic need for suitable bottom type and conditions of rapid flow. The gradients in dissolved oxygen,

temperature, suspended particulates, etc., ISLANDS PACIFIC are less understood and are a priority for future work. Available surveys indicate coral beds dense with colonies that cover large areas are the exception. Given the region has little history of trawling and mobile bottom-tending gear, it is reasonable to assume this is the natural condition.

 Hawaiian fish are known to opportunistically use the corals as shelter and to some degree they co-occur with corals in high flow habitats. It is not known how the fish behave with deep corals in the other parts of the Pacific. Invertebrates are largely unstudied and the degree of their association with deep corals is unknown but likely to be greater.

 Currently, the greatest threat to corals is the potential for spread of the invasive species Carijoa riisei from the Auau channel to other black coral beds on Kauai and Hawaii. Following that, the harvesting in the 3 beds where the coral fishery operates needs to be closely monitored. For the black coral bed in the Auau channel, attention is needed because of the unanticipated loss of black coral to Carijoa riisei. At Makapuu, the regrowth of pink coral has been documented once and should be checked for continued resilience. The Keahole bed was targeted for red and gold coral, and its prospects for resilience are as yet unproven. Finally,

175 176 PACIFIC ISLANDS ae F (95 Cnrbtos o the to Contributions (1955) FM Bayer re-description and Description (1952) FM Bayer of corals primnoid new Two (1951) FM Bayer deep- the in flow Gene prep) (In AR Baco and Distribution prep) (In TS Shirley AR, Baco Six (2006) TM Shank AM, Clark AR, Baco genetic Population (2005) TM Shank AR, Baco deep-sea the on Report (1902) A Alcock deep-sea the of account An (1898) A Alcock X. REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF REFERENCES tutr o te aain precious Hawaiian the of structure Siboga Expedition16a Expedition. Siboga the of Madreporaria Calcutta: Trustees IndianMuseum Investigator. Royal ship survey the marine Indian by collected Madreporaria opooy f h . and the Museum 105:207-220 of Proceedings of the United States systematics, National morphology nomenclature, Telstacea) , PacificScience6:126-136 Steamer (1.Alcyonacea, Commission “” Fish collected US octocorals the by Hawaiian the of 41:40-43 Sciences of WashingtonAcademy the of Journal Calyptrophorinae. subfamily the the Archipelago. from Hawaiian the of seamounts and islands Coralliidae) (Octocorallia: coral precious sea Hawaiian of precious corals. coral invertebrates deep-sea associated of preference habitat and islands Molecular EcologyNotes6:147-149 the archipelago. Hawaiian the of seamounts from Coralliidae) sea deep the coral from loci microsatellite ecosystems. and Springer-Verlag BerlinHeidelberg corals Cold-water 663-678 in Freiwald A, Roberts JM (eds.), Pages microsatellites. using Corallidae) coral oalu laauense Corallium laauense Corallium oalu laauense Corallium (Octocorallia: (Octocorallia:

ae F (95 A e seis f the of species new A (1995) FM Bayer octocoral Isidid new A (1990) FM Bayer of species old and new Some (1982) FM Bayer Bayer FM (1956) Descriptions and re-descriptions Berntson EA, Bayer FM, McArthur AG, SC Primnoidae (1989) J Stefani FM, Bayer of species new A (1988) J Stefani FM, Bayer (1997) FM Bayer of species new Three (1996) FM Bayer ognca genus gorgonacean Biological the Society ofWashington 103:205-228 of Pacific Proceedings the Ocean. in elsewhere from New species from Caledonia, with descriptions of other new Gorgonacea) (Anthozoa: 95:110-160 of Society Biological the of Proceedings genus related Fanelia the of revalidation with genus primnoid the Pacific Science10:67-95 . Gorgonacea: 2. steamer Albatross Commission Fish collected U.S. Octocorals the by Hawaiian the of 138:235-246 Anthozoa) sequences. rRNA 18s nuclear on (: based Octocorallia the within relationships Phylogenetic (2001) Paris 10:449-518 Nouvelle-Caledonie. de Naturelle d’Histoire Musee du Bulletin de (Gorgonacea) the Biological of Society ofWashington 101:257-279 Proceedings Pacific. Western the from species other some of descriptions with Caledonia, New from Chrysogorgia the Biological of Society ofWashington 110:511-519 Proceedings Pacific. family the eastern the from (Anthozoa) Primnoidae of octocoral gorgonacean Washington 109:205-228 of Society Biological the of Proceedings Genus Gorgonacea, (Anthozoa: coral precious waters. Washington 108:147-152 of Society Hawaiian Biological the of Proceedings from Octocorallia) ry Ceetrt: Anthozoa). (Coelenterata: Gray Corallium (Octocorallia: Gorgonacea) ael nuttingi Narella ) from Pacific waters. Pacific from ) Narella Callorgorgia (Anthozoa: a new a , Gray, STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

Boland RC, Parrish FA (2005) Description of fish Dana JD (1846) Zoophytes. U.S. exploring assemblages in the black coral beds off expedition during the years 1838-1842, Lahaina, Maui, Hawaii. Pacific Science under the command of , 59: 411-420 U.S.N. 7: i-iv + 1-740. Lea and Blanchard, Philadelphia. Brook G (1889) Report on the Antipatharia. Report of the Scientific Results of the Devaney DM, Eldridge LG (1977) Reef and shore Voyage of HMS Challenger 32:1-222. pls. fauna of Hawaii, Section 1: 1-15 through Ctenophora. Bishop Musuem Special Publication 64(1) p 278 Cairns SD (1978) Distichopora (Haplomerismos) anceps, a new Stylasterine coral DiNardo GT, Moffitt RB (2007) The northwestern (Coelenterata: Stylasterina) from Hawaiian Islands Lobster Fishery: A deep water off the Hawaiian Islands. Targeted Slipper Lobster Fishery. In Micronesica 14:83-87 Lavalli KL, Spanier E (eds) The biology and fisheries of the slipper lobster CRC Cairns SD (1982) Antarctic and subantarctic Press pp 243-261 Scleractina. Antarctic Research Series 3: 74 pp. Druffel ER, Griffin MS, Witter A, Nelson E, Southon J, Kasgarian M, Vogel J (1995) Cairns SD (1984) New records of ahermatypic Gerardia: Bristlecone pine of the deep- ISLANDS PACIFIC corals (Scleractinia) from the Hawaiian sea? Geochima et Cosmochimica Acta and Line Islands. Occasional Papers 59:5031-5036 of the Bernice Pauahi 25(10):1-30 Duchassaing P, Michelotti J (1860) Memoire sur les coralliaires des Antilles. Memorie Cairns SD (1999) Cnidaria Anthozoa: Deep- della Reale Accademia delle Scienze di Water azooxanthellate Scleractinia from Torino19: 279-365, pls. 1-10 , and Islands. Memoires du Museum National d’histoire Duncan PM (1872) On the structure and affinities Naturelle, 180:31-167 of Guynia annulata Duncan, with remarks on the persistence of Palaeozoic types of Cairns SD (2005) A revision of the Hawaiian Madreporaria. Philosophical Transactions Stylasteridae (Cnidaria: Hydrozoa: of the Royal Society of London 162:29- Athecata). Pacific Science 59:439-451 40

Cairns SD (2006) New records of azooxanthellate Duncan PM (1873) A description of the Scleractinia from the Hawaiian Islands. Madreporaria dredged up during Occasional Papers of the Bernice Pauahi expeditions of H.M.S. Porcupine in Bishop Museum 87:45-53 1869 and 1870. Part 1. Philosophical Transactions of the Royal Society of Coutis PF, Middleton JH (2002) The physical and London 8:303-344, pls. 39-49 biological impact of a small island wake in the deep ocean. Deep Sea Research I Duncan PM (1876) Notice of some deep-sea and 49:1341-1361 littoral corals from the , , Indian, , Persian Chave EH, Mundy BC (1994) Deep-sea benthic Gulf, and Japan &c. . Proceedings fish of the Hawaiian Archipelago, Cross of the Zoological Society of London Seamount, and Johnston Atoll. Pacific 1876:428-442, pls. 38-41 Science 48:367-409

Chave E H, Malahoff A (1998) In deeper waters. University of Hawaii Press, Honolulu, HI. p 125

177 178 PACIFIC ISLANDS Grigg RW (1974) Distribution and abundance of abundance and Distribution (1974) RW Grigg Grigg RW (1965) Ecological studies of black coral forms new two of Description (1866) JE Gray Goodman-Lowe G (1998) Diet of Hawaiian monk FN Spiess PF, Lonsdale PK, Dayton A, Genin from Learning (1998) SD der Cairns N, Feinstein Fortsetzungen (1794) EJC Esper Pflanzenthiere. Die pp. (1792) 2, EJC Vol. Esper Englund RA (2002) The loss of native biodiversity Ellis J, Solander D (1786) The natural history of the REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF au Hwia Ilns Islands. Hawaiian Oahu, estuarine, and freshwater,communities of , in species introductions nonindigenous continuing and Sons; P. Elmsly, London and White Benjamin 1-63. pls. 1-208, + systematically Ellis, John arranged by the late Daniel Solander. i-xii late the by globe the of parts various from collected many curious and uncommon zoophytes, 235-240 pp Symposium Reef Coral International 2 Proceed. Hawaii. in corals precious in Hawaii.PacificScience19:244-260 27 1866:24- London, of Society Zoological the of Proceedings corals. gorgoniid of during 1991-1994. MarineBiology132:535-546 Islands Hawaiian Northwestern ( seal over acceleration deep-sea topography. Nature,322:59-61 current of evidence (1986) Corals on seamount peaks provide Nautilus, 112:73-832 and The patterns. analysis attachment of discussion an with Xenophoridae), corals affixed by azooxanthellate of survey a collector: the 1-64, Nurnberg pp. 1-2, parts 1, Volume Pflanzenthiere. 133-164, Nurnberg 25:418-430 oahs schauinslandi Monachus Xenophora (; fo the from ) nd

Grigg RW (1988b) Recruitment limitation of a deep Grigg RW (1988a) Paleoceanography of the coral Order Antipatharia (1977) D RW,Opresko Grigg knowledge Present (1976) FM RW,Bayer Grigg commercial The (1975). LG RW,Eldridge Grigg ie I (89 Te ognca olce by collected Gorgonacea The (1899) IL Hiles Grigg RW (2004) Harvesting impacts and invasion Grigg RW, Grossman EE, Earle SA, Gittings SR, Hawaii: in corals Precious (2002) RW Grigg a of history coral: Black (2001) RW Grigg of fisheries coral Precious (1993) RW Grigg benthic hard-bottom octocoral population octocoral hard-bottom benthic chain. Hawaiian-Emporer Science 240:1737-1743 the in reefs Vol. 1:242-261 LG (eds) Reef and shore fauna of Hawaii. (black corals). In: Devaney DM, Eldredge Science 30:167-175 Pacific Hawaii. in Gorgonacea order the of zoogeography and systematics the of 75-01 Publ. Guam University Marine Laboratory Grant Sea Islands. Mariana The 1: Part potential of precious corals in Micronesia. Part 2.Cambridge. Results, Zoological Dr.Willey’sWilley: A. Science 58:1-6 black coral yield off Maui, Hawaii. Pacific by an alien species decrease estimates of Coral Reefs21:73-82 Islands. Hawaiian S.E. Channel, Au,au topography, karst antecedent and reefs Drowned (2002) J, McDonaugh D, Lott Marine FisheriesReview64:13-20 revised and bed management new measures for existing beds. a of Discovery Pacific Science 55:291-299 Hawaii. in fishery sustainable Islands. Pacific S. Marine FisheriesReview55:50-60 U. the and Hawaii Progress Series45:121-126 Ecology Marine Islands. Hawaiian the in STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

Hoover JP (1998) Hawaii’s Sea Creatures, A Knauss JA (1996) Introduction to physical guide to Hawaii’s marine invertebrates Prentice Hall p 338 Mutual Publishing, Honolulu Hawaii p 366 Kolliker A (1870) Beitrage zur kenntnis der polypen. Verhandlungen der Physikalisch Hubrecht AA (1885) On a new pennatulid from – Medicinischen Gesellschaft in Wurzburg, the Japanese Sea. Proceedings of the 2:11-29, pls. 3-4 Zoological Society of London Kolliker A (1880) Report on the Pennatulida Kahng SE, Grigg RW (2005) Impact of an alien dredged by H.M.S. Challenger during the octocoral, Carijoa riisei, on black corals in years 1873-1876. London Hawaii. Coral Reefs 24:556-562 Kukenthal W (1907) Versuch einer Revision Kamenkovich VM, Koshlyakov MN, Monin AS der Alcyonacean. 2. Die Familie der (1986), Synoptic eddies in the ocean. D. Nephthyiden. 3. Teil. Die Gattungen Reidel Publishing Company Eunepthya Verrill und Marenzelleria Zoologische Jahrbucher Keller NB (1974) New data about some species Systematik 24:317-390 of Madreporarian corals of the genus . Trudy Instituta Okeanologii Kukenthal W (1910) Zur kenntnis der gattung 98:199-212 Anthomastus verr. In Doflein, F. (Ed.) PACIFIC ISLANDS PACIFIC Beitrage zur naturgeschichte ostasiens. Kelley C R, Ikehara W (2006). The impacts of math.-phys. Klasse K. bayer. Akad. bottomfishing on Raita and West St. Wissenschaft., Suppl. Bd. 1:1-16 Rogatien Banks in the Northwestern Hawaiian Islands. Atoll Research Bulletin Kükenthal W (1919) Gorgonaria. In 543:307 Wissenschaftliche Ergebnisse der deutschen Tiefsee Expedition auf dem Kelley CR, Moffitt RB, Smith JR (2006) Mega to Dampfer “” 1898-1899 13:1-946, micro-scale classification and description pls. 30-89 of bottomfish essential fish habitat on four banks in the Northwestern Hawaiian Lesson RP (1831) Illustrations de zoologie. Islands. Atoll Research Bulletin 543: 321 Paris.

Kennett JP (1982) , Prentice- Linnaeus C (1758) Systema naturae per regna Hall, Inc. p 813 tria naturae. Editio decima, reformata: 1- 824. Holmiae Kinoshita K (1907) Vorlaufige Mitteilung uber einige neue japanische Primnoidkorallen. Longenecker K, Dollar RA, Cahoon MK (2006) Annotationes zoologicae Japonenses Increasing taxonomic resolution in dietary 6:229-237 analysis of the . Atoll Research Bulletin 543:101 Kinoshita K (1908) Primnoidae von Japan. Journal College of Science, Imperial University of Miller JE, Hoeke RK, Appelgate TB, Johnson PJ, Tokyo23: 1-74, pls. 1-6 Smith, J, Bevacqua S (2003). Bathymetric Atlas of the Northwestern Hawaiian Kinoshita K (1909) On the Primnoidae, a family Islands, Draft.May 2003, National of the Gorgonacea. Dobutsugaku zasshi Oceanic and Atmospheric Administration 21 (243):1-100 and Hawaii Mapping Research Group, p 65 Kinoshita K (1913) Studien uber einige Chrysogorgiiden . Journal College of Science, Imperial University of Tokyo33:1-47, pls. 1-3

179 180 PACIFIC ISLANDS ulnax S Mls W 19) ts o the of test A (1997) SW Mills LS, Mullineaux Hydriod, certain on Report (1881) HN Moseley Moseley HN (1880) Description of a new species to report Preliminary (1876) HN Moseley of feasibility The (2002) AD Montgomery of assessment An (1992) FA Parrish RB, Moffitt JJ(1989) FA, Polovina Parrish RB, Moffitt Recherches (1848b) J Haime H, Edwards Milne Recherches (1848a) J Haime H, Edwards Milne REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF Naturelles series3,10:65-114, pl.1 des Eupsammides. Annales des Sciences Monographie 3. Mem. Polypiers. les sur des 4-6 Annales Sciences Naturelles series general. 3, 9:37-89, pls en des development Polypiers le et structure la sur Observations 1. Mem. Polypiers. les sur 44:745-770 I Research Sea Deep flows, generated seamount- in hypothesis retention larval HMS Challenger2:127-208 of the Voyage of Results Scientific the of 3. On the deep-sea Madrepararia. Report Part 1873-1876. years the in Challenger corals procured during the voyage of the H.M.S. Madreporarian and Alcyonaria, the of Proceedings Zoological SocietyofLondon1880:41-42 coral. simple of Biological Sciences24:544-569 of the Royal Society of London. Series B: 31 Aug. and 1870, 30 Dec. dates the between water deep in Challenger H.M.S. by dredged corals true the on Thompson Wyville Professor (Order coral black Antipatharia). Hydrobiologia 471:157-164 transplanting Fishery Bulletin90:476-482 submersible. a from observations 2: Part laevigatus of biomass exploitable the Science Marine 44:616-630 of Bulletin and Hawaii. in reefs artificial biomass deepwater of structure, Community in the main Hawaiian Islands. Hawaiian main the in st 17. Proceedings 1875. , Heterocarpus th , Nutting CC (1910) The Gorgonacea of the Siboga Nutting CC (1908) Descriptions of the Alcyonaria of revision systematic A (1979) KM Muzik gorgonian, bioluminescent A (1978) K Muzik Opresko DM (2005) New genera and species of species and genera New the (2005) DM Opresko of Revision (2003) DM the Opresko of Revision (2002) DM Opresko classification the of study A(1974) DM Opresko Nutting CC (1912) Descriptions of the Alcyonaria 11 1- pls. 13b:1-39, Monographs Expedition expedition. VI. The Gorgonellidae. Siboga Museum 34:543-601 National States United the of Proceedings 1902. in Islands Hawaiian the of vicinity the in Albatross Steamer Fisheries of Bureau U.S. the by collected 227 and p Miami. of University Dissertation. PhD Paramuriceidae Octocorallia). (Coelenterata: Plexauridae Hawaiian the Bulletin ofMarineScience28:735-741 olapa (Coelenterata: Octocorallia), from Hawaii. Lepidisis from the North Pacific. Zoologische Pacific. North Mededelingen 79-2p129 the from Antipatharian corals (Cnidaria: Anthozoa) Zoologische (Cnidaria:Anthozoa). . Mededelingen Leiden77:495-536 III: Part Antipatharia Zoologische Anthozoa). . Mededelingen Leiden76:411-442 II: (Cnidaria: Part Antipatharia 1-149. CoralGables,FL (Coelenterata: Univ.Miami: Dissertation, Ph.D. species. Anthipatharia eleven of redescriptions with Anthozoa), the of 1-21 pls. 43:1-104, Museum National States United the of Proceedings waters, 1906. during Japanese in mainly Albatross, Steamer Fisheries U.S. the by collected nw species new , STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

Pallas PS (1766) Elenchus zoophytorum sistens Pourtalès LF (1878) Report on the corals of the generum adumbrationes generaliores “Blake” expedition. Bulletin of the Museum et specierum cognitarum succinctas of Comparative Zoology at Harvard descriptions cum selectis auctorum University 5:197-212 synonymis. i-xvi, I 28, I 451. Hagae- Comitum Ralston S, Tagami DT (1992) An Assessment of the exploitable biomass of Heterocarpus Parke MF, Wang HY (2005). Bathymetric laevigatus in the main Hawaiian islands. Atlas of the main Hawaiian Islands – Part 1: Trapping surveys, depletion Information for Ecosystem Assessment experiment, and length structure. Fishery and Monitoring. National Oceanic and Bulletin 90:494-504 Atmospheric Administration, PIFSC Data Report DR-05-002 Ridley SO (1884) Alcyonaria. In Report on the zoological collections made in the Indo- Parrish FA, Boland RC (2004) Habitat and reef- during the voyage of the fish assemblages of bank summits in the H.M.S. Alert 1881-2: 327-365, pls. 36-38. Northwestern Hawaiian Islands. Marine British Museum, London. Biology 144:1065-1073 Roark EB, Guilderson TP, Dunbar RB, Ingram Parrish FA, Abernathy K, Marshall GJ, Buhleier BL (2006) Radiocarbon based ages and

BM (2002). Hawaiian monk seals growth rates: Hawaiian deep sea corals. ISLANDS PACIFIC (Monachus schauinslandi) foraging in Marine Ecology Progress Series 327:1- deep-water coral beds. 14 Science 18:244-258 Sars GO (1872) On some remarkable forms of Parrish FA (2006). Precious corals, and subphotic life from the great deeps off the fish assemblages. Atoll Research Bulletin Norwegian coast. Christiana. Brogger 543:425-438 and Christie

Parrish FA, Abernathy K (2006). Movements of Schultze LS (1896) Beitrag zur systematic monk seals relative to ecological depth antipatharia. Abhandlungen der zones in the lower northwestern Hawaiian Senckenbergishcen Naturforschenden Islands. Atoll Research Bulletin 543:113 Gesellschaft 23: 1-40

Parrish FA (In press) Density and habitat of three Stewart BS, Antonelis GA, Baker JD, Yochem deep-sea corals in the lower Hawaiian PK (2006) Foraging biogeography of chain. Bulletin of Marine Science Hawaiian monk seals in the Northwestern Hawaiian Islands. Atoll Research Bulletin Polovina JJ (1994) The lobster fishery in the 543:131 Northwestern Hawaiian Islands. In: Phillips BF, Cobb JS, Kittaka J (eds) Spiny Stiasny G (1947) De gorgonien-familie lobster management. Blackwell Scientific Acanthogorgiidae Kukenth. & Gorz. (Met publications Oxford pp 83-90 bijzondere Inachtneming van het material der Siboga-Expeditie). Verh. kon. Nederl. Pourtalès LF (1868) Contributions to the fauna Akad. Wetensch., Afd. Naturelle 43: 1-93, of the at great depths pls. 1-3. Amsterdam (second series). Bulletin of the Museum of Comparative Zoology at Harvard Studer T (1894) Notes preliminaire sur les University 1:121-141 alcyonaires. Bulletin of the Museum of Comparative Zoology at Harvard Pourtalès LF (1871) Deep-sea corals. Ill. University 25:53-69 Catalogue of the Museum of Comparative Zoology at Harvard University 4:1-93 Tchernia P (1980) Descriptive Regional Oceanography. Pergamon Press p 253

181 182 PACIFIC ISLANDS el J (94 Rcn crl o te Marshall the of corals Recent (1954) JW Wells Reproductive Press) (In AR Baco , RG Waller Siboga- der Gorgoniden Die (1906) J Versluys water shallow Hawaiian (1928) AE Verrill and Anthozoa, the on Report (1883) AE Verrill Vaughan TW (1907) Recent Madreporaria of the the of Antipatharia The (1914) AJ Pesch van Groundfish (1984) DT Tagami RN, Uchida Oceanography.Introductory (1981) HV Thurman (1906) WD Henderson account JA, An Thomson REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF ihre ad eerh n h vcnt of vicinity the in research and Fisheries Company. p129 3 the IndianMuseum,Calcutta.p132 the deep sea. XVI:10 pls. The Trustees of of alcyonarians The I. Ocean. Indian the Indian Marine Survey Ship Investigator in of the alcyonarians collected by the Royal sad. .. elgcl Survey Geological U.S. Professional Paper260-I: 385-486 Islands. Bulletin ofMarineScience secundum Archipelago: deep- Hawaiian the from corals of precious water species three of morphology 187, pls.1-10 13a:1- dition. Monographs Expedition Primnoidae. Siboga Die II. Expedition Bishop Pauahi Museum Bulletin49:1-30 Bernice Anthozoa. Harvard at Zoology University 11:1-72 Comparative Museum the of of Bulletin 1880-1882. in Hawk” “Fish steamer Commission Fish U.S. the by and 1877-1879, in Blake the by dredged species additional some on 59:1- 222, pls.1-96 Museum National States United Bulletin. Laysan. and Islands Hawaiian Expedition Monographs Amsterdam, 17:1-258 Siboga Expedition. Siboga Marine FisheriesReview46:1-17 Ocean. Pacific North the in seamounts rd d Cals . erl Publishing Merrill E. Charles ed. and Gerardia oalu laauense Corallium sp., Corallium . Western Pacific Regional Fishery Management Fishery Regional Pacific Western Management Fishery Regional Pacific Western Management Fishery Regional Pacific Western Indo-Pacific on Notes (1982) JW Wells rgt P Sue T 18) eot n the on Report (1889) T Studer EP, Wright of Report In: Alcyonaria. The (1885) EP Wright Mullineaux M, Gowing L, Levin K, Wishner Management Fishery Regional Pacific Western NOAA Admin Award NAO7FC0680 of WESTPAC Region. Fishery Environmental Pacific Western the Pelagic Final Statement, Impact (1991) Council region. Honolulu, HI Pacific western the of fisheries corals precious the for plan management 1and environmental assessment for the fishery Amendment and western region. the Pacific of fishery the corals for precious regulations proposed management and plan Fishery (1980). Council Pacific Islands. Galapagos Science 36:211-220 the from corals New 9. Part corals. scleractinian Zool. 31:1-314,pls.1-43 Alcyonaria collected by H.M.S. Challenger. of HMS ChallengerNarrative1:689-693 the Voyage of Results Scientific the oxygen seamount. Nature346:57-59 the deep a on zonation of benthic in minimum Involvement (1990) L 18-19 48p. Apr report workshop management and science coral Black 2006 (2006) Council WESTPAC NOAA Admin Region. Award NAO7FC0680 of the and Pacific Western Fishery Bottomfish Groundfish Seamount Environmental Statement, Final Impact (2005) Council STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION References Cairns, 1999 1907, Cairns, 1984 Vaughan, Pourtalès, 1871, Cairns, 1984 Cairns, 1984 Duncan, 1873, Cairns, 1984 1907, Cairns, 1984 Vaughan, Cairns, 1984 1907, Cairns, 1984 Vaughan, 1881, Cairns, 1984 Moseley, Alcock, 1898, Cairns, 1984 1907 Vaughan, Cairns, 1984 Alcock, 1902, Cairns, 1984 1881, Cairns, 1984 Moseley, Alcock, 1898, Cairns, 1984 ** (m) *** Depth Range 369 205-296 201-271** 274-336** 1602** 44-388** 331-337** 457-627** 137-439** 56-206** 583-678** 244-322** 165-600** 197-634** 274-518** PACIFIC ISLANDS PACIFIC All species listed are found in Hawaii except one Distribution HI Islands Atoll, HI Islands & RMI HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands & Christmas Island, Line Islands HI Islands D. andamanicus Species C. ambrosia A. fecunda sp. cf. sp. cf. sp. cf. and several antipatharins as noted. List for octocorals scleractinians in Hawaii based primarily on Anthemiphyllia macrolobata Anthemiphyllia pacifica Anomocora Bourneotrochus stellulatus Caryophyllia atlantica Caryophyllia hawaiiensis Caryophyllia marmorea Caryophyllia octopali Caryophyllia Caryophyllia ”Ceratotrochus” laxus Coenosmilia inordinata Conotrochus funicolumna Crispatotrochus rubescens Deltocyathus List of known species ceep corals from the U.S. Pacific Islands. Keroeides koreni Higher Taxon Cnidaria Anthozoa Class Subclass () Order Scleractinia Anthemiphylliidae Family Family Appendix 4.1. octocoral, A. Baco and Stephen Cairns, Smithsonian Institution, with additions from recent Pisces cruises led by unpublished list compiled by Dr. additions for non-Hawaiian islands based on cited literature. List antipatharians from Hawaii and Guam unpublished list compiled Dennis Opresko. by and courtesy of Dr. * = depth range known from full for species, including outside chapter region ** = depth range known from 2 or more specimens Hawaii, Christmas, Line Islands, etc *** = depth from a single individual Hawaii, usually the holotype

183 184 PACIFIC ISLANDS STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF Depth Range Higher Taxon Species Distribution (m References Desmophyllum dianthus HI Islands Esper, 1794 Paracyathus molokensis HI Islands 161-260** Vaughan, 1907 Trochocyathus aithoseptatus HI Islands 371-454** Cairns, 1984 Trochocyathus burchae HI Islands 64*** Cairns, 1984 Trochocyathus gardineri HI Islands 274-470** Vaughan, 1907, Cairns, 1984 Trochocyathus mauiensis HI Islands 174-278** Vaughan, 1907 Trochocyathus oahensis HI Islands 75-571** Vaughan, 1907, Cairns, 1984 Trochocyathus patelliformis HI Islands 1020*** Cairns, 1999 Trochocyathus rhombocolumna HI Islands 110-530* Alcock, 1902 Family Balanophyllia desmophyllioides HI Islands 143-406** Vaughan, 1907 Balanophyllia diomedeae HI Islands 110-307** Vaughan, 1907, Cairns, 1984 Balanophyllia gigas HI Islands 90-640* Moseley, 1881 Balanophyllia laysanensis HI Islands 238-271** Vaughan, 1907 Cladopsammia echinata HI Islands 295-470** Cairns, 1984 Cladopsammia eguchii HI Islands Wells, 1982 Eguchipsammia gaditana HI Islands 244-470** Duncan, 1873, Cairns 1984 Eguchipsammia fistula HI Islands Alcock, 1902 Eguchipsammia serpentina HI Islands 269-362** Vaughan, 1907 Enallopsammia rostrata HI Islands 362-583** Pourtalès, 1878, Cairns, 1984 Milne-Edwards & Haime, 1848a & b, Endopachys grayi HI Islands 37-274** Cairns, 1984 HI Islands & NW of Family Flabellum marcus Wake Island 1261-1602** Keller, 1974, Cairns, 1984 HI Islands 183-517** Lesson, 1831, Cairns, 1984 Flabellum vaughani HI Islands 232-369** Cairns, 1984 Javania exserta HI Islands 400*** Cairns, 2006 Javania fuscus HI Islands 13-271** Vaughan, 1907

HI Islands & Christmas Javania insignis Island, Line Islands 52-825** Duncan, 1876, Cairns, 1984 STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION References Moseley, 1880, Cairns, 1984 Moseley, Feinstein and Cairns, 1998, Cairns 2006 Cairns, 2006 Cairns, 1984 Sars, 1872, Cairns, 1984 1907, Cairns, 1984, 2006 Vaughan, Duncan, 1872, Cairns, 1984 1876, Cairns, 1984 Moseley, 1907, Cairns 2006 Vaughan 1907, Cairns, 1984 Vaughan, Cairns, 1984 Duncan, 1876, Cairns 1984, 2006 Pourtalès, 1868, Cairns, 1982, Cairns, 1984 1928 Verrill, van Pesch, 1914 Pallas, 1766 Pallas, 1766 Schultze, 1896 Brook, 1889, Grigg and Opresko, 1977 Dana, 1846, Grigg and Opresko, 1977 Linnaeus, 1758, Grigg and Eldridge, 1975 (m Depth Range 244-322** 119-291*** 440-858** 212-503** 1762-2056** 369-541** 64-384** 109-470** 362-538** 362-538** 627-750** 439-494** 439** 347-366*** 25-40*** PACIFIC ISLANDS PACIFIC Distribution HI Islands & Johnston Atoll HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands & S. Pacific Seamounts HI Islands HI Islands HI Islands Guam Guam HI Islands HI Islands Guam Guam HI Islands dichotoma

spinuilosa A. . Species A. curvata A. flabellum A sp., cf. n. sp., cf. sp. cf. sp., cf. Javania lamprotichum Placotrochides minuta Polymyces wellsi Fungiacyathus fissilis Fungiacyathus fragilis Gardineria hawaiiensis Guynia annulata Letepsammia formosissima Madracis kauaiensis Madrepora kauaiensis Deltocyathoides orientalis Stenocyathus vermiformis Antipathes grandis Antipathes Antipathes Antipathes Antipathes Antipathes intermedia Cirrhipathes anguina Cirrhipathes contorta Cirrhipathes propinqua Cirrhipathes spiralis Higher Taxon Family Fungiacyathidae Family Gardineriidae Family Guyniidae Family Micrabaciidae Family Pocilloporidae Family Oculinidae Family Turbinoliidae Family Stenocyathidae Antipatharia Order Family

185 186 PACIFIC ISLANDS STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF Depth Range Higher Taxon Species Distribution (m References

Stichopathes echinulata HI Islands 305-565*** Brook, 1889, Grigg and Opresko, 1977 Family sarothamnoides Guam Brook, 1889 van Pesch, 1914, Grigg and Opresko, Acanthopathes undulata HI Islands 110-490** 1977 HI Islands & Channel Family Cladopathidae Trissopathes pseudotristicha Islands 326-4539** Opresko, 2003 Trissopathes tetracrada HI Islands 375-425** Opresko, 2003 Family Leiopathidae HI Islands Esper, 1792, Opresko, 1974 Prelim. ID by Opresko, Pisces Cruise Leiopathes n. sp. HI Islands 403-471** 2003 &2004 Ellis and Solander, 1786, Grigg and Family Myriopathes ulex HI Islands & Guam Eldridge, 1975 Myriopathes sp., cf. M. japonica HI Islands Brook, 1889 Cupressopathes abies Guam Linnaeus, 1758 Ellis and Solander, 1786, Grigg and sp., cf. A. subpinnata HI Islands 455-460*** Opresko, 1977

Family Schizopathidae Bathypathes alternata HI Islands 1195-1744** Brook, 1889, Pisces Cruise 2003 Bathypathes conferta HI Islands 380*** Brook, 1889, Grigg and Opresko, 1977 Bathypathes patula HI Islands Brook, 1889, Unpubl record at USNM HI Islands & Johnston 220-441, 1400- Stauropathes staurocrada Atoll 1700** Opresko, 2002 Prelim. ID by Opresko, Pisces Cruise Stauropathes sp. HI Islands 604*** 2003 Umbellapathes helioanthes HI Islands 1205-1383** Opresko 2005 Prelim. ID by Opresko, Pisces cruise Umbellapathes, new species B HI Islands 742-744*** 2004 Dendropathes bacotaylorae HI Islands 408*** Opresko 2005 Order Zoanthidea Zoanthid blue HI Islands 352-415** Chave and Malahoff, 1998 STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION References Chave and Malahoff, 1998 Chave and Malahoff, Ryland and Baco in prep., Chave 1998 Malahoff, 1998 Chave and Malahoff, 1998 Chave and Malahoff, 1998 1952, Chave and Malahoff, Bayer, 1906, fide Williams Versluys, 1952 Kukenthal 1910, Bayer, fide Williams And Mich., 1860 Duch. 1952 Nutting, 1908, Bayer, 1952 Nutting, 1908, Bayer, 1952 Nutting, 1908, Bayer, 1952 Bayer, fide Williams 1952 Nutting, 1908, Bayer, 1952 Nutting, 1908, Bayer, 1976 Grigg and Bayer, Muzik, 1979 1976 1947, Grigg and Bayer, Stiasny, Berntson et al., 2001 1976 Nutting, 1908, Grigg and Bayer, 1884, Nutting, 1908 Ridley, 1976 1956, Grigg and Bayer, Bayer, (m Depth Range 500-1910** 343-577** 343-460** 332-1025** 356-462** 20-201** 966* 914* 518-616* 223-283* 144* 215-564** 350-396** 1295*** 71-396** 237-2533* 340-465, 1387- 1820** PACIFIC ISLANDS PACIFIC Distribution HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands Invasive, HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands A. striata Nutting, 1911 A. paramuricata Species sp. cf. sp. cf. sp. n. sp. sp. 1 sp. 2 sp. sp. sp. Zoanthid tan Gerardia Parazoanthus Anthomastus fisheri Anthomastus (Bathyalcyon) robustus Anthomastus granulosus Inflatocalyx Carijoa riisei Clavularia grandiflora corrugata Telestula spiculicola Telestula spiculicola robusta Telestula Nidalia alexanderi Siphonogorgia collaris Acanthogorgia Acanthogorgia Acanthogorgia Acanthogorgia Cyclomuricea flabellata Muricella tenera Anthothela nuttingi Higher Taxon Subclass Octocorallia Alcyonacea Order Family Family Clavulariidae Family Order Gorgonacea Acanthogorgiidae Family Family

187 188 PACIFIC ISLANDS STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF Depth Range Higher Taxon Species Distribution (m References Antothela n. sp. 1 HI Islands 1319*** Prelim. ID by Cairns, Pisces Cruise 2003 Antothela n. sp. 2 HI Islands 1804*** Prelim. ID by Cairns, Pisces Cruise 2003 Family Chrysogorgiidae Chrysogorgia arborescens HI Islands 722-914* Nutting, 1908 Bayer and Stefani, 1988, Berntson et al., Chrysogorgia chryseis HI Islands 732*** 2001 Chrysogorgia delicata HI Islands 536-1463* Nutting, 1908 Chrysogorgia elegans HI Islands 433-634* Verrill, 1883, Nutting, 1908 Chrysogorgia flavescens HI Islands 1688-1977* Nutting, 1908

Chrysogorgia geniculata HI Islands 146-616* Wright & Studer, 1889, Nutting, 1908 Wright & Studer, 1889, Grigg and Bayer, Chrysogorgia sp. cf. C. japonica HI Islands 750-1050** 1976 Kinoshita, 1913, Grigg and Bayer, 1976, Chrysogorgia paillosa HI Islands 704-1858* Nutting, 1908 Bayer and Stefani, 1988, Chave and Chrysogorgia scintillans HI Islands 580-2050** Malahoff, 1998 Chrysogorgia stellata HI Islands 649-678* Nutting, 1908 646-675, 830- Nutting, 1908, Grigg and Bayer, 1976, Chrysogorgia sp. cf. C. stellata HI Islands 922* Bayer and Stefani 1988 Chrysogorgia n. sp. (1/3R) HI Islands 1204*** Prelim. ID by Cairns, Pisces Cruise 2003 Chrysogorgia n. sp (2/5L) HI Islands 691-742** Prelim. ID by Cairns, Pisces Cruise 2004 Iridogorgia superba HI Islands 704-914* Nutting, 1908, Grigg and Bayer, 1976

Iridogorgia bella HI Islands 750-1925** Nutting, 1908, Chave and Malahoff, 1998 Iridogorgia. n. sp. HI Islands 1443*** Prelim. ID by Cairns, Pisces Cruise 2003 Metallogorgia melanotrichos HI Islands 183-1385* Wright and Studer, 1889, Nutting, 1908 Metallogorgia n. sp. HI Islands 1805*** Prelim. ID by Cairns, Pisces Cruise 2003 Pleurogorgia militaris HI Islands 2142* Nutting, 1908

Radicipes spiralis HI Islands 258** Nutting, 1908, Grigg and Bayer, 1976 STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION References Bayer, 1956, Bayer unpubl ms Bayer, 1955, Bayer unpubl ms Bayer, 1996, Bayer unpubl ms Bayer, 1976 1956, Grigg and Bayer, Bayer, 1956, Bayer unpubl ms Bayer, 1976 1956, Grigg and Bayer, Bayer, 1956, Grigg and Dana, 1846, Bayer, 1976, Pisces Cruise 2003 Bayer, 1955, Pisces Cruise 2003 Bayer, Prelim. ID by Cairns, Pisces Cruise 2003 Prelim. ID by Cairns, Pisces Cruise 2003 Prelim. ID by Cairns, Pisces Cruise 2003 1976 1956, Grigg and Bayer, Bayer, 1976 Grigg and Bayer, 1990 Bayer, Nutting, 1910 1990 Bayer, Muzik museum id Prelim. ID by Cairns, Pisces Cruise 2003 1976 Nutting, 1908, Grigg and Bayer, Nutting, 1908 1976 Grigg and Bayer, 1889, Grigg and Bayer and Studer, Wright 1976 Muzik, 1978 1889, Nutting 1908, and Studer, Wright Muzik, 1978 Berntson et al., 2001 (m Depth Range 1829-2403*** 1145*** 365-580** 232-282*** 365-719** 231-576** 1096*** 167-408** 275-495** 275-445** 1920*** 1808*** 346-465** 1344-1582* 305-565** 215-665** 539-631** 1425*** PACIFIC ISLANDS PACIFIC Distribution HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands Species A secundum cf. n. sp. “5” n. sp. n. sp. sp. sp. n. sp. (lyrate) Corallium abyssale Corallium ducale Corallium kishinouyei Corallium laauense Corallium niveum Corallium regale Corallium secundum Corallium imperiale Corallium Corallium laauense x halmahera Corallium Paracorallium tortuosum Acanella dispar Acanella weberi Isidella trichotoma Isidella Isidella Keratoisis flabellum Keratoisis grandis Keratoisis Lepidisis nuda Lepidisis olapa Lepidisis paucispinosa Lepidisis Higher Taxon Family Coralliidae Family Family Isididae

189 190 PACIFIC ISLANDS STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF Depth Range Higher Taxon Species Distribution (m References Family Keroeididae Keroeides fallax HI Islands 238-245*** Bayer, 1956 Keroeides koreni Marshall Islands Wright & Studer, 1889, Bayer, 1956 Keroeides mosaica HI Islands 167-465** Bayer, 1956, Grigg and Bayer, 1976 Keroeides pallida HI Islands 146*** Hiles, 1899, Bayer, 1956 Family Paragorgia dendroides HI Islands 490-1910** Bayer, 1956, Chave and Malahoff, 1998 Paragorgia sp. cf. P. regalis Nutting, 1912 HI Islands 350-396** Grigg and Bayer, 1976 Paragorgia n. sp. HI Islands 350-396** Grigg and Bayer, 1976 428-531, 581- Family Plexauridae Anthomuricea tenuispina HI Islands 688** Nutting, 1908, Grigg and Bayer, 1976 Anthomuricea sp. cf. A. divergens HI Islands 381-426** Kükenthal, 1919, Grigg and Bayer, 1976 Anthomuricea sp. cf. A. reticulata HI Islands 362-421** Nutting, 1910, Grigg and Bayer, 1976 Anthomuricea n. sp. A HI Islands Muzik, 1979 brunnea HI Islands 167-396** Nutting, 1908, Grigg and Bayer, 1976 Bebryce n. sp. HI Islands Muzik, 1979 Muriceides sp. A HI Islands Muzik, 1979 Muriceides sp. B HI Islands Muzik, 1979 Filigella n. sp. Thesea n. sp. HI Islands Muzik, 1979 Muriceides tenuis HI Islands 232-362* Nutting, 1908, Muzik, 1979 Muriceides n. sp. A HI Islands Muzik, 1979 Muriceides n. sp. B HI Islands Muzik, 1979 New genus, n. sp. HI Islands Muzik, 1979 Thomson & Henderson, 1906, Muzik, Paracis horrida HI Islands 1979 Paracis miyajimai HI Islands 362-531** Kinoshita, 1909, Grigg and Bayer, 1976 Paracis n. sp. A HI Islands Muzik, 1979 Paracis spinifera HI Islands 350-396** Nutting, 1912, Grigg and Bayer, 1976 350-396, 924- HI Islandsensis HI Islands 1241** Nutting, 1908, Grigg and Bayer, 1976 STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION References Bayer unpubl ms Muzik, 1979 Muzik, 1979 1976 Nutting, 1910, Grigg and Bayer, 1976 Grigg and Bayer, 1976 Nutting, 1910, Grigg and Bayer, Thom & Hend, 1906, Grigg and Bayer, 1976 1976 Grigg and Bayer, 1976 Grigg and Bayer, Muzik 1979 1976 Nutting, ??, Grigg and Bayer, 1889, Grigg and Bayer, & Studer, Wright 1976 1976 Grigg and Bayer, 1976 Grigg and Bayer, Muzik, 1979 Muzik, 1979 Muzik, 1979 1982 Kukenthal, 1907, Bayer, 1976 Nutting, 1908, Grigg and Bayer, 1976 Grigg and Bayer, 1976 1894, Grigg and Bayer, Studer, 1976 Nutting, 1908, Grigg and Bayer, 1951 Bayer, 1976 1866, Grigg and Bayer, Gray, 1976 Grigg and Bayer, (m Depth Range 350-396** 335-375** 73, 182** 147, 350-396** 340-365** 350-396** 313-399** 315-412** 350-396** 350-396** 215-960** 350-396** 781-1145** 1207-3292* 12-1275 216-432** 344-454** PACIFIC ISLANDS PACIFIC Distribution HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands Ps. orientalis Species P. dendritica P. Ps. placoderma sp. cf. n. sp. T. ramosa T. n. sp. A sp. n. n. sp. B sp. cf. sp. sp. cf. n. sp. n. sp. 2 n. sp. C n. sp. 1 A n. sp. n. sp. B sp. cf. n. sp. 1 n. sp. 2 Placogorgia Placogorgia Placogorgia Placogorgia Placogorgia Placogorgia Pseudothesea Swiftia Swiftia pacifica Thesea arbuscula Villogorgia Villogorgia Villogorgia Villogorgia Villogorgia Callogorgia formosa Callogorgia gilberti Callogorgia Calyptrophora agassizii Calyptrophora angularis Calyptrophora clarki Calyptrophora japonica Calyptrophora Higher Taxon Family Primnoidae

191 192 PACIFIC ISLANDS STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF Depth Range Higher Taxon Species Distribution (m References Calyptrophora wyvillei HI Islands 744-823* Wright, 1885, Nutting, 1908 Calyptrophora n. sp. 1 (lyrate) HI Islands 1078*** Prelim. ID by Cairns, Pisces Cruise 2003

Calyptrophora n. sp. 2 HI Islands 1807*** Prelim. ID by Cairns, Pisces Cruise 2003 Wright & Studer, 1889, Pisces cruise Candidella gigantea HI Islands 1720-1815** 2003

Candidella helminthophora HI Islands 38-1820** Nutting, 1908, Grigg and Bayer, 1976 Fanellia euthyeia HI Islands Bayer and Stefani, 1989 Fanellia medialis HI Islands Bayer and Stefani, 1989 Fanellia tuberculata HI Islands Versluys, 1906, Bayer, 1982 Nutting, 1908, Berntson et al., 2001, Narella bowersi HI Islands 1344-1937* Grigg and Bayer, 1976 Narella dichotoma HI Islands Versluys, 1906, Bayer ms: 27 Narella sp. cf. N. megalepis HI Islands 215-564** Kinoshita, 1908, Grigg and Bayer, 1976 Narella nuttingi HI Islands 1350*** Bayer, 1997, Berntson et al., 2001 Narella ornata HI Islands 748-1007*** Bayer, 1995 Narella studeri HI Islands Versluys, 1906 Narella n. sp. 1 HI Islands 350-396** Grigg and Bayer, 1976 Narella n. sp. 2 HI Islands 353-417** Grigg and Bayer, 1976

Narella. n. sp. (unbranched) HI Islands Prelim. ID by Cairns Pisces Cruise 2003 Paracalyptrophora n. sp. HI Islands 367-398** Prelim. ID by Cairns Pisces Cruise 2004 Parastenella n. sp. HI Islands 517*** Prelim. ID by Cairns Pisces Cruise 2004 Plumarella n. sp. HI Islands 384-432** Grigg and Bayer, 1976 Thouarella (A.) biserialis HI Islands 439* Nutting, 1908, Grigg and Bayer, 1976 Thouarella (A.) regularis HI Islands 183-722* Wright and Studer, 1889, Nutting, 1908 Thouarella sp. cf. T. (T.) typica HI Islands 350-396** Kinoshita, 1907, Grigg and Bayer, 1976 STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION References Kolliker, 1880, Nutting, 1908 Kolliker, 1998 Nutting, 1908, Chave and Malahoff, Hubrecht, 1885, Nutting, 1908 1998 Chave and Malahoff, fide Williams Prelim. ID by Cairns, Pisces Cruise 2003 Nutting, 1908 Nutting, 1908 1880, Nutting, 1908 Kolliker, Nutting, 1908 Nutting, 1908 Nutting, 1908 Nutting, 1908 1880, Nutting 1908 Kolliker, Nutting, 1908 Nutting, 1908 1870 Kolliker, 1880 Kolliker, Cairns, 1978, 2005 Cairns, 2005 Cairns, 2005 Cairns, 2005 (m Depth Range 426-2286* 196-1650** 225-232* 254-1940** 223-316* 402-530* 1033* 903-1033* 523* 925* 97-421* 1046-2056* 708-1951* 704-2403* 223* 1265-1280* 360-577** 293-377** 322-583** 521-563** PACIFIC ISLANDS PACIFIC Distribution HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands HI Islands Species sp. (short stemmed) sp. Anthoptilum murrayi Calibelemnon symmetricum Echinoptilum macintoshi Funiculina Halipterus willemoesi Kophobelemnon Pennatula flava Pennatula pallida Pennatula pearceyi Pennatula sanguinea Protoptilum wrighti Protoptilum attenuatum Protoptilum studeri Umbellula carpenteri Umbellula gilberti Umbellula jordani abies Virgularia molle Virgularia Distichopora (Haplomerismos) anceps Distichopora asulcata Stylaster griggi Stylaster infundibuliferus Higher Taxon Order Pennatulacea Anthoptilidae Family Family Chunellidae Family Echinoptilidae Family Funiculinidae Family Halipteridae Family Kophobelemnidae Family Pennatulidae Family Protoptilidae Family Umbellulidae Family Virgulariidae Class Hydrozoa Order Stylasterina Family Stylasteridae

193 194 PACIFIC ISLANDS STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION PACIFIC ISLANDS THE IN ECOSYSTEMS STATE CORAL DEEP OF