STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

STATE OF DEEP CORAL ECOSYSTEMS OF THE U.S. PACIFIC COAST: CALIFORNIA TO WASHINGTON

Curt E. Whitmire and M. Elizabeth Clarke

I. INTRODUCTION Beginning in the 1960s and continuing into the current period, there has been an increasing use The U.S. Pacific coast marine region of drop camera systems, submersibles and, most encompasses the continental margin off the recently, ROVs to make in situ observations. states of California, Oregon and Washington. These underwater photographic platforms have Deep corals were first reported here in the 1860s been used to explore numerous seafloor features with descriptions by A.E. Verrill, including two in the region including rocky banks (e.g., Pearcy stony corals, Balanophyllia elegans (1864) and et al 1989; Stein et al. 1992; Pirtle 2005; Tissot et

Paracyathus stearnsii (1869), and one stylasterid, al. 2006), canyons (e.g., Yoklavich et al. 2000), COAST PACIFIC Allopora californica (1866). Because all three escarpments (e.g., Carey et al. 1990; Clague et of the species occur in shallow waters, it is not al. 2001), seamounts (e.g., DeVogelaere 2005) surprising that they were the first in the region and other rocky features (Hyland et al. 2004; to be reported (Cairns 1994; Ostarello, 1973). Brancato et al. 2007). Many of these features Dall (1884) also provided early descriptions of are known to support deep coral communities. hydrocorals off California and Alaska as well as accounts from fishermen of bycatch of Stylaster The disparate data sources that will be referenced sp. off the Farallone Islands, California as early throughout this report have their strengths as 1873. and weaknesses as they inform discussions of deep coral communities and their habitats. In addition to the taxonomic literature, records For example, trawl surveys in the region are of deep corals in the region come from a variety limited to low relief, sedimentary habitats that of other sources including catch records from support relatively few emergent epifauna as regional bottom trawl surveys, bycatch data compared with hard-bottom habitats (Figure collected by fishery observers and observations 3.1). Consequently, bottom trawls rarely sample from underwater vehicles (e.g., submersibles stony corals and stylasterids, but have resulted and remotely operated vehicles (ROVs)). In the in numerous records of pennatulaceans as well early 1970s, the Alaska Fisheries Science Center as fewer records of gorgonians, black corals (AFSC) began conducting triennial bottom trawl and soft corals (Table 3.1). Furthermore, the surveys of demersal fishes in the region. Early level of identification during trawl surveys varied surveys included records of pennatulaceans and according to the priority given for sampling of a few gorgonians. Unfortunately, identifications invertebrates and the level of expertise of the down to any appreciable taxonomic level were biologists onboard. Despite these limitations, initially very limited, typically only to order or trawl surveys encompassed large portions of sometimes family. In 1998, the Northwest the continental margin (including shelf and Fisheries Science Center (NWFSC) began slope depths). Consequently, they contribute conducting annual bottom trawl surveys, but like to discussions of general zoogeography of the early AFSC surveys, identifications of corals some higher coral taxa (e.g., order and family were initially not a priority. levels). On the other hand, in situ photographic surveys in the region primarily target rocky, high relief structures that support diverse benthic Fishery Resource Analysis & Monitoring Division communities, many of which include corals. Northwest Fisheries Science Center Surveys of this type, while limited in extent, National Marine Fisheries Service provide detailed information about the size, 2725 Montlake Blvd. East health and habitat affinities of corals, and their Seattle, WA 98112-2097 relationships between other invertebrates and

109 110 PACIFIC COAST queried. Northwest FisheriesScienceCentersbetween1980 and 2005. A totalof10,526trawlcatchrecordswere trawl surveys,whichwereconductedoff thecoastsofWashington, OregonandCaliforniabythe Alaska and Table 3.1. communities coral protect to enacted measures conservation related and region the in corals to the chapter includes a review of potential impacts addition, In highlighted. are organisms other for that provide vertical structure as potential habitat taxa and region, the in occur to habitat known taxa all general and characteristics. zoogeography The discussions authors attempted to identify and for coral stage geology of the regional set communities. of oceanography deep their descriptions and of region Brief the discussion in includes corals chapter This Family Stylasteridae; Appendix 3.1). Anthoathecatae, Order Hydrozoa, (Class of stylasterid corals species five and families, eleven from species Pennatulacea) (Order 27 pennatulaceans families, of three from Alcyonacea) 36 10 families, eight species of families, true soft corals (Order three from Gorgonacea) (Order gorgonians of species from Antipatharia) corals black (Order of species seven families, seven stony of species from (Class Scleractinia) corals Order Anthozoa, 18 including EEZ the identified within been have orders cnidarian corals six of from species 101 total, bathyal In the meters). to (3900 down intertidal the from zones depth including (EEZ), zone economic exclusive the of boundary seaward the to out shoreline the from and 49°N and 32 between latitudes span coast offPacific corals U.S. deep the of Records specimen collection capabilities. provide ROVs) platforms submersibles, some (e.g., species- however making identifications, of level challenge the is surveys photographic of limitation One fishes. demersal PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF Pennatulaceans Antipatharians Scleractinians Alcyonaceans Stylasterids Gorgonians GeneralstatisticsondeepcoralssampledduringNational MarineFisheriesService(NMFS)bottom Total # Trawls withCorals 2259 1683 150 197 202 26 1 Laboratories. Center forHabitatStudiesatMossLandingMarine information foroff Californiawasprovidedbythe Mapping LabatOregonStateUniversity. Lithology was providedbythe Active Tectonics andSeafloor the continentalmarginoff Washington andOregon were usedforanalysis.Lithologyinformation where thepositionoffishinggearwasknown of Washington, OregonandCalifornia.Onlytrawls 2001-2003 duringsurveysofgroundfishoff thecoast Northwest FisheriesScienceCenter(NWFSC)from gies transectedbybottomtrawlsconductedthe Figure 3.1. % Trawls withCorals 100% 10% 20% 30% 40% 50% 60% 70% 80% 90% 0% <0.1% 16.0% 0.2% 1.4% 1.9% 1.9% Distributionofgeneralseafloorlitholo Rocky R elative Frequency elative R % CoralRecords 100.0% dmentary edim S <0.1% 74.5% 1.2% 6.6% 8.7% 8.9% - STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION and their habitats. Finally, the authors provide have formed over the past tens of millions of recommendations for future research to improve years by hotspot volcanism and by enhanced our understanding of these organisms. melting in association with the migration of the spreading centers over a heterogeneous mantle (Davis and Karsten 1986). One of the largest II. GEOLOGICAL SETTING seamounts – Davidson – has been the site of several ROV surveys that have discovered a The marine region off the coasts of Washington, diverse coral community (see DeVogelaere Oregon and California accounts for about 7% 2005). A number of additional seamounts that (778,628 km2) of the total area of the U.S. EEZ are known to support deep corals lie just to the (NMFS 2007 in prep). The continental margin in west of the EEZ boundary, including Cobb, Brown this region is characterized by a narrow (5-40 km) Bear, Fieberling and Jasper. shelf and steep continental slope, with the shelf break at approximately 200 meters water depth. Other megascale (i.e., km to 10s of km, Greene The outer continental shelf off Oregon and parts et al. 1999) structural features in the region of California are marked by large rocky banks that contribute hard-bottom habitats include the (Figure 3.2), some of which were at or near the Mendocino and Gorda Ridges (Figure 3.2). The surface during the lower sea level stands of the Mendocino Fracture Zone is a 3000-km long COAST PACIFIC glacial epochs. Several of these banks as well transform fault extending from Cape Mendocino, as other bathymetric features such as pinnacles California across the Pacific Plate. A prominent and seamounts create localized upwelling hard-bottom feature associated with this fault conditions that concentrate nutrients, thus driving is the Mendocino Ridge, which shoals to 1100 a high level of biologic productivity. For example, meters water depth and drops 2100 meters to Heceta Bank, which rises over 100 meters above the north-south trending Gorda Ridge. To the the edge of the continental shelf and to within 70 south, the Mendocino ridge slopes 3300 meters meters of the ocean surface, diverts the main flow to the abyssal plain. In contrast to the Mendocino of the California Current, introducing eddies and Ridge, the Gorda Ridge is a seafloor spreading other instabilities that affect areas downstream center where two plates are moving apart, and along the Oregon coast. Smaller rocky allowing molten magma to rise up to form new banks are located off southern California in oceanic crust. It extends 300 km and is bounded what is called the continental borderlands, a by the Mendocino Fracture Zone to the south geologically complex region characterized by and the Blanco Fracture Zone to the north. The deep basins and elevated ridges, some of which Gorda Ridge rises to a maximum height of >1500 breech the surface to form the Channel Islands. meters above the axial valley floor, which ranges Throughout the region, many of these high relief, from 3200 to >3800 meters water depth. Like bathymetric features have been found to support other spreading centers, hydrothermal vents that coral communities (see Love et al. 2007; Tissot support unique communities of chemosynthetic et al. 2006; Tissot et al. in prep; Strom 2006). organisms exist nearby. Morphology of the Mendocino and Gorda Ridges were described by The shelf and slope are cut by numerous Fisk et al. (1993) and Clague and Holmes (1987), submarine canyons, including one of the deepest respectively. and largest on the west coast of North America - Monterey Canyon (Figure 3.2). Deep corals have been discovered at numerous sites here, III. OCEANOGRAPHIC SETTING and also within another large canyon – Astoria – located directly off the mouth of the Columbia Oceanographic circulation in the region is well River. Other major canyons include Juan de Fuca, described and was summarized recently by Quinault, Bodega, Pioneer and Sur Canyons. NMFS (2007 in prep). The U.S. Pacific region is one of the major upwelling areas of the world, Beyond the slope, a number of seamounts rise where nutrient-rich waters support high levels of above the abyssal plain including Thompson, biological productivity. Physical oceanography President Jackson, San Juan, Rodriquez, Taney, varies seasonally and during El Niño and La Guide, Pioneer and Davidson (Figure 3.2). The Nina events or periods of interdecadal climate seamounts off the west coast of North America regime shifts. Major oceanographic currents

111 112 PACIFIC COAST in thischapterarelabeled. Figure 3.2. PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF Bathymetric mapoftheU.S. Pacificcoastregion.Majortopographicfeaturesthat arementioned STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION in the region include the surface- flowing California Current, the Inshore Countercurrent (Davidson Current) and the Southern California Countercurrent, and the subsurface-flowing California Undercurrent. The California Current forms the eastern boundary of a large clockwise circulation pattern in the North Pacific. The California Current is a year- round flow that transports cold, nutrient- rich subarctic water equatorward (Hickey 1998). It extends from the shelf break to about 1000 km offshore with peak speeds at the surface during spring and summer yet significant flows down to 500 meters (Hickey 1998). South of Point Conception, the California

Current splits, with its core continuing COAST PACIFIC farther offshore while a smaller portion turns shoreward both north and south of the Channel Islands (Figure 3.3). Near San Diego, part of the core flow turns northward to form the Southern California Countercurrent, an inshore poleward flow off Southern California. During some years, the Southern California Countercurrent forms a counterclockwise circulation pattern known as the Southern California Eddy Figure 3.3. Map of portion of the Pacific coast of North America (Figure 3.3). During other years, the showing major oceanographic currents. Image source: NMFS Southern California Countercurrent 2007, in prep. rounds Point Conception and combines with the Inshore Countercurrent, a poleward flow al. 2002). A shift in wind direction in the winter inshore of the equatorward California Current north of Santa Cruz, CA creates favorable (Hickey 1979, Figure 3.3). Below these surface downwelling conditions, while upwelling currents lies a narrow, high speed flow known conditions persist year-round south of Santa Cruz as the California Undercurrent, which brings due to modest storm activity (Strub and James warmer, nutrient-poor waters poleward along the 2000). Upwelling plumes also occur at coastal slope from the eastern equatorial Pacific (Pierce headlands, particularly where the California et al. 2000). A major feature of the eastern North Current and Inshore Countercurrent intersect off Pacific and the California Current is a layer where Pt. Conception. oxygen concentrations are low within a depth zone along the upper continental slope, extending Another important hydrographic feature in the to depths greater than 1000 m. This feature lies northern part of the region is the Columbia beneath the California Undercurrent and is called River plume, contributing as much as 90% of the oxygen minimum zone (dissolved oxygen the freshwater input between the Strait of Juan <0.5 ml l-1 (22 mM kg-1) (Deuser 1975). de Fuca and San Francisco Bay (NMFS 2007 in prep). The position of the plume is highly Circulation in the region is driven by the intensity seasonal, generally extending equatorward and and duration of prevailing seasonal winds and offshore in the spring and summer, and poleward storms. Spring and summer northwesterly along the coast in the fall and winter (Thomas winds drive an upwelling system that replenishes and Weatherbee 2006). nutrients to the photic zone, which in turn stimulate biological productivity (Batchelder et According to Briggs (1974), the temperate

113 114 PACIFIC COAST a be cletd t or ie i te Olympic the in sites four at collected been has 1994; Mary Elaine Helix, MMS, pers. comm.) and al. et (Hardin California southern off distributed and haimei Astrangia region, oculata the Madrepora in occur to including known are species elegans) (e.g., corals cup non-branching 3.1). (Appendix solitary,of are coral stony of families records the of Most 18 seven include from coast Pacific species the off corals Stony Scleractinia) a. Stonycorals of specificinformationonstructuralattributes. not be designated as so in this region due to lack considered structure-forming in other regions may are Taxathat region. this in information current to limited are follow that taxa structure-forming of discussions Likewise, region. the in corals of size, colony maximum structure-forming attributes of are specific to exception records the With 3.2). (Table organisms other with associations of number relative the and clumped) form vs. (i.e., solitary dispersion colony spatial to colony maximum of known manner size, is abundance, taxa relative the reefs, not or whether non-branching), vs. branching (i.e., morphology or include structure invertebrates to contributing other Attributes fish. by utilized be can that floor sea the above structure vertical provide to known are they meaning “structure-forming”, as designated are region the in taxa coral Several IV. Province attheStraitofJuandeFuca. northern boundary of the the Oregon placed (or “Oregonian”) (1971) McGowan by zoogeography and Alaska between British Columbia. Another boundary review of Pacific Ocean maritime the Entrance, Dixon the to north Conception Point from extends and region temperate cold the in The Oregon Province is the lower boreal province Mexico. Sur, California Baja Bay, Magdalena to region, extends from Point Conception, CA south warm-temperate the in Province, Diego San The California. and Oregon Washington, coastal off waters include three which of two provinces, includes zoogeographic Pacific northeastern PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF Coenocyathus bowersi Coenocyathus DEEP CORALS STRUCTURE AND HABITAT-FORMING Hwvr a lat ee branching seven least at However, . ohla pertusa Lophelia (Class Anthozoa, Order , , ayitoytu quaylei Labyrinthocyathus edohli oldroydae Dendrophyllia . . , L. pertusa L. cln profunda Oculina Balanophyllia is widely is , ,

ie itiuin, while distributions, wide and sp. 3.1). Antipathidae, Appendix 2002, (Opresko 2003, families, 2005, Schizopathidae and three Cladopathidae from seven only species with speciose very not but coast Pacific the off abundant very are Antipatharians Antipatharia) b. Blackcorals Institution, pers.comm.; Table 3.2). profunda Oculina are corals stony structure-forming Other (Table3.2). importance structural of rating high a given was it California, that fact the and discovery this of Because Pacific. largest the aggregation support of may NMS Coast Olympic the et in site One 2007). (Hyland al. et Brancato 2004; al. Sanctuary Marine National Coast is found in high abundance (including extensive (including abundance high in found is 3.1). (Appendix families 10 of group from species 36 speciose with coast Pacific the off corals most the are Gorgonians Gorgonacea) d. Gorgonians (DeVogelaere etal.2005). one unconfirmed record from Davidson Seamount only with region, the in rare very are corals Gold Zoanthidea) c. Goldcorals of structuralimportance(Table 3.2). reasons, these For 2007). al. et (Love years 140 to aged recently was individuals, invertebrate 2500 over stars, high) dead colony, which brittle was heavily colonized by amphipods, anemones, sponges and crabs. One large (2.1 m crinoids, including invertebrates other with associations epifaunal showed specimens these of Many 2005). Love meters 360 and (Love 90 et al. 2007; Tissot between et al. depths 2006; Yoklavich water and at California southern off banks deepwater several on submersible via observed been has cm, 250 to 10 from height in ranging species described A. abundance. high in occur and cm, >30 heights Bathypathes comm.). pers. Monterey NMS, Burton, Bay (Erica Seamount Davidson Umbellapathes

dendrochristos L. pertusa Antipathes Bathypathes sp. is relatively abundant off southern L. pertusa p hv ol be cletd at collected been only have sp. (Class Anthozoa, Order (Class Anthozoa, Order (Class Anthozoa, Order (Stephen Cairns, Smithsonian Cairns, (Stephen

edohli oldroydae Dendrophyllia Orso 05, newly a 2005), (Opresko r bacig cn reach can branching, are spp. was given a high rating reported in the northeast p ehbt coast exhibit sp. Lillipathes Antipathes Paragorgia arborea Paragorgia Antipathes p and sp. p and sp. and

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Table 3.2. Structure-forming attributes of some deep coral taxa off the U.S. Pacific coast. Relative abundance data was compiled from taxonomic records, in situ photographic surveys, and to a lesser extent bottom trawl surveys. Numbers of species associations were quantified from in situ photographic surveys. Size, morphol- ogy and other colony attributes were informed by taxonomic descriptions. ‘?’ means insufficient data available in the region to comment. Attributes Contributing To Structure

Overall Maximum Associations Colony Rating of Reef- Relative Colony with Other Spatial Structural Taxa Building Abundance Size Morphology Species Dispersion Importance Lophelia pertusa No High Large Branching Many Clumped High Oculina profunda No Low ? Branching ? ? Low Dendrophyllia oldroydae No Medium Medium Branching ? Clumped Medium Antipathes dendrochristos No High Large Branching Many Solitary High

Bathypathes sp. No High Medium Branching ? Solitary Medium COAST PACIFIC Isidella sp. No High Medium Branching ? Solitary Medium Keratoisis sp. No High Medium Branching ? Solitary Medium Paragorgia arborea No High Large Branching Many Solitary High Primnoa pacifica No Low Large Branching Many Solitary High

Table Key Attribute Measure Reef-Building Yes/No Relative Abundance Low/ Medium/ High Size (width or height) Small (< 30cm)/ Medium (30cm-1m)/ Large (>1m) Morphology Branching/ Non-branching Associations None/ Few (1-2)/ Many (>2) Spatial Dispersion Solitary/ Clumped Overall Rating Low/ Medium/ High

“forests” observed along several ridges on 205-234 meters are the southernmost record of Davidson Seamount [DeVogelaere et al. 2005]), the species in the Pacific (Cairns and Barnard can reach heights >1 m and has shown epifaunal 2005). Specimens of Keratoisis and Corallium relationships with numerous other structure- from Davidson Seamount have been aged to forming invertebrates. P. arborea is therefore over 200 and 115 years, respectively (Andrews given a high rating of structural importance et al. 2005). (Table 3.2). Isidella spp. and Keratoisis spp. are found coast wide mostly on the continental slope. e. True soft corals (Class Anthozoa, Order Although both genera can reach heights greater Alcyonacea) than 30 cm, other gorgonians (e.g., P. arborea Only eight species of true soft corals from three and Primnoa pacifica) in the region can reach families occur off the Pacific coast (Appendix heights exceeding 1 meter. Therefore, Isidella 3.1). Anthomastus sp. is abundant and exhibits and Keratoisis were given a medium rating of coast wide distributions, while Gersemia sp. has structural importance (Table 3.2). ROV surveys been caught primarily on the northern Oregon in the Olympic Coast NMS have resulted in slope during trawl surveys. Alcyonium rudyi and numerous observations of gorgonians including Cryptophyton goddarti were described recently large colonies of P. pacifica, numerous colonies (1992 and 2000, respectively) off the Oregon of Plumarella longispina and smaller colonies of coast (Cairns et al. 2002). Other than C. goddarti, Leptogorgia chilensis, Swiftia pacifica,and Swiftia references to Clavularia and Telesto off southern beringi at several sites. Colonies of P. pacifica California (SCAMIT 2001), and Telestula ambigua obtained off La Jolla, CA (north of San Diego) at in deep water off central California (Austin 1985),

115 116 PACIFIC COAST found. typically are stylasterids which at depths shallow and habitats high-relief the target not do trawls bottom because likely most (Table3.1), catches survey trawl in identified been rarely have corals Stylasterid structure-forming. not be are to they considered width, or height in cm rarely 30 region exceed the in specimens most Because while and two The 1983). Cairns 1938; porphyra Stylantheca include Province Oregon the in distributions narrower exhibit that species Other Province. Oregon the throughout found while Province Diego San the Alberto 1983; californicus Cairns 3.1). Appendix comm., 1938; pers. Lindner, the (Fisher in occur to region known are genera three from species five Only depths. water intertidal shelf to the down from habitats rocky high-relief to moderate colonizing observed been have coast Pacific the off corals stylasterid or corals Lace Order Anthoathecatae, SuborderFilifera) g. Stylasteridcorals this report. than by defined as forming structure species be to considered fish some adjacent areas (e.g., Brodeur 2001), they are not of densities higher support to shown been groves have pennatulaceans of Although wide. coast found are which Umbellula 3.1). (Appendix coast Pacific are the off families occur to eleven known from species 27 date, To 3.1). (Table surveys trawl from recorded coral taxon common most the also are They 2001). Brodeur 2005; al. et (Stone offAlaska observed aggregations to similar individuals numerous of submersibles and ROVs either alone or in groves coral abundant taxon in the region and have been most observed from the are Pennatulaceans Pennatulacea) f. Pennatulaceans to bestructure-forming. considered are region the in corals soft true the of none stature, small their of Because the region. in stoloniferans on data other no are there PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF E. pourtalesii Stylatula Stylantheca sp. are the most common taxa, all of all taxa, common most the are sp. s h ol seis nw from known species only the is sp., are flabellate (i.e., fan-shaped) s nrsig Cin 1983). (Cairns encrusting is Anthoptilum grandiflorum (Class Anthozoa, Order and (Class Hydrozoa, Errinopora pourtalesii Errinopora S. petrograpta S. Stylaster . venustus S. Stylaster species (Fisher and is , conducted during each survey varied from year from varied survey each during conducted trawls of number (34.5ºN). the Conception Also, Pt. of south effort survey trawl limited was there 2002, to Prior variable. temporally and spatially however,depth); been effortwater has m survey of the continental shelf and upper slope (10-1600 2005. Cumulatively, both surveys covered much 1980- from records catch on focuses report this therefore limited; very initially was invertebrates of Identification 1998. in surveys ongoing began (NWFSC) Center Science andthe Fisheries Northwest 1971-2001, from coast Pacific the off surveys trawl regional conducted (AFSC) Center Science Fisheries Alaska The 3.1). (Appendix (NMFS) Service Fisheries Marine National the by conducted surveys trawl bottom and records taxonomic from originates region the in corals of Much of the information on general zoogeography V. n Pinia) n syatrd oas (Family corals stylasterid and Primnoidae) and of gorgonians (Corallidae, one Isididae, Paragorgiidae Caryophyllidae), family of black corals and (Antipathidae), four families (Oculinidae corals include representatives from two families of stony California and Oregon Washington, off family). Records and genus (e.g., taxa are higher to records limited many but records, of these of identities some the confirmed have Taxonomists (SIO). Oceanography of Institution (NMNH), Scripps and History Natural of Museum National Smithsonian’s the (MBARI), of Aquarium Institute Bay Research Academy Monterey the California the (CAS), by Sciences the coast off Pacific corals structure-forming and of observations collections of records compiled (2003) Morgan and Etnoyer surveys, trawl to addition In encompassing fifteenspeciesfromsixfamilies. from six families, and 225 records of other corals species thirteen encompassing pennatulaceans of records 316 include trawls SCCWRP (32%) Allen et al. 2002). Catch records from 304 of 957 1998; al. et (Allen 2003 to 1994 from California southern off surveys shelf Project continental different Resource Water (SCCWRP) conducted bottom trawls during three Coastal California Southern the NMFS, to addition 3.1, In (Table 3.4). Figure trawls of coral 5% other only in all occurred while taxa trawls, survey coral of 16% for recorded in were queried Pennatulaceans were occurrences. records catch trawl year.to A7252 of total NWFSC 3274 and AFSC SPECIES AND HABITATS SPATIAL DISTRIBUTIONOFCORAL STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION PACIFIC COAST PACIFIC

Figure 3.4. Maps of frequency of occurrence for two groups of deep coral taxa sampled during NMFS bottom trawl surveys (1980-2005). Frequency defined as number of trawls with corals identified in the catch sample divided by total number of trawls within each 20x20 km cell. Frequency was categorized into three classes: >0-20%, >20-50%, and >50%. Cells where survey trawls occurred but where no corals were identified in the catch sample are labeled as “NO Catch” and are symbolized with an empty box where the underlying bathym- etry shading is visible. Pennatulaceans were singled out because they inhabit different habitat types and were caught much more frequently than other coral taxa. See Table 3.1 for frequency information.

117 118 PACIFIC COAST oe eald nomto o crl aias in habitats coral on information detailed More pertaining to thetwozoogeographic provinces. general sections following the in highlighted are which of the many region, to the in taxa coral of zoogeography contributes information This coast.Pacific U.S. offthe EEZ the corals of much span of records 389 of total A Stylasteridae). PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF ocpin C, n icue te geologically the includes and CA, Conception, Province Diego San Point to north border, Mexican the the from extends of portion U.S. The San DiegoProvince brief descriptionsofthese surveysareprovided. and sources data the possible, When surveys. by provided is region the n situ in photographic

Figure 3.5. Map showing locations of black corals (Order Antipatharia) from NWFSC, AFSC, and SCCWRP trawl surveys, Etnoyer and Morgan (2003) and Tissot et al. (2006). Specific identities from trawl survey catch records are unconfirmed and primarily limited to genus or family level. STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION AFSC, and SCCWRP trawl surveys, Etnoyer AFSC, and SCCWRP PACIFIC COAST PACIFIC Map showing locations of stony corals (Order Scleractinia) from NWFSC, Figure 3.6. and Morgan (2003) Brancato et al. (in review). Specific identities from trawl survey catch records are unconfirmed primarily limited to genus or family level. complex borderlands (Figure 3.2). A number of and Love 2005, Figure 3.5). Lophelia pertusa species found in this region, such as the newly and Desmophyllum dianthus have been observed described black coral, Antipathes dendrochristos on numerous high-relief, hard-bottom features (Opresko 2005) have not been described further below 120 meters near oil platforms surveyed in north. These black corals have been observed via the late 1980s off Pt. Conception, CA (Steinhauer submersible on numerous rocky outcrops in the and Imamura 1990; Hardin et al. 1994). Near province at water depths ranging from 90 to 360 one platform, D. dianthus and L. pertusa were m (Love et al. 2007; Tissot et al. 2006; Yoklavich among the most abundant taxa observed in high- 119 120 PACIFIC COAST (Cairns 1994). 1994). (Cairns Islands including 25 specimens deposited at SIO coral, cup stearnsii The 1994). (Cairns Islands Channel the around especially province arnoldi scleractinian, Another meters. 212 160- from ranging depths water at habitats relief PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF i kon o cu truhu the throughout occur to known is , i as cmo aon te Channel the around common also is , Coenocyathus bowersi Coenocyathus Paracyathus Caryophyllia has been has 1994, Figure 3.6). two records known from the northeast Pacific of (Cairns one only is Island, Anacapa near depth oculata, Madrepora scleractinian, colonial the of record One 1994). and down to 80 meters off Pt. Conception (Cairns of Islands Channel the coasts around California, mainland the off nearshore the from collected collected at 84 meters water meters 84 at collected

Figure 3.7. Map showing locations of gorgonians (Order Gorgonacea) from NWFSC, AFSC, and SCCWRP trawl surveys, Etnoyer and Morgan (2003) and Tissot et al. (2006). Specific identities from trawl survey catch records are unconfirmed and primarily limited to genus or family level. STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION - PACIFIC COAST PACIFIC Map showing locations of true soft corals (Order Alcyonacea) from NWFSC, AFSC, and SCCWRP trawl surveys and Et AFSC, and SCCWRP Alcyonacea) from NWFSC, Map showing locations of true soft corals (Order Figure 3.8. noyer and Morgan (2003). Specific identities from trawl survey catch records are unconfirmed primarily limited to genus or family level.

Gorgonians are not as prevalent as they are in catches from trawl surveys range in water depths the Oregon Province (Figure 3.7); however, this from 77-1400 meters. Pennatulaceans (mostly may be due to sampling bias. Tissot et al. (2006) members of suborder Subselliflorae) have observed 27 specimens from four different been observed from underwater vehicles in the habitat types at 144-163 m. Other records in sedimented flanks of numerous rocky outcrops in the province include Lepidisis sp. at 950 m and the province (9726 specimens from Tissot et al. Keratoisis sp. far offshore San Diego at 3180 and 2006) and are caught more often than other coral 3880 m (Etnoyer and Morgan 2003). Gorgonian taxa in bottom trawls at water depths ranging 121 122 PACIFIC COAST shelf depths, and pourtalesii respectively. Other stylasterids include corals, stylasterid two of extents distributional southern and northern the mark CA) Islands Francisco, San Farallone (off The 1994). (Cairns CAS at deposited specimens 45 including Bay Monterey British and Columbia. state Washington of waters inside including Province Oregon the throughout from 2006). (Aspen habitats hard-bottom moderate-relief on percent and two species of anemones often reached 100 cup corals ( CA, Arguello, Pt. of coverage epifaunal off routes cable submarine proposed of surveys During 36,000 2005). (Pirtle Bank over including habitats of observations 1994), bottom 1983, hard (Cairns nearshore in primarily common are corals cup solitary and Stylasterids Diego at water depths of 37-293 m (Cairns 1994). quaylei province. the in found elsewhere also are stylasterids and Scleractinians Cape Flattery, WA (A.Lindner, pers.obs.). observed in water depths of about 100 meters off coral, stylasterid the of Colonies 1982). Stanley and Cairns 1994; al. et Hardin (see world and region the in elsewhere which is consistent with observations of association with observed. Desmophyllum were colonies living and dead Both 2007). al. et (Brancato depth water meters 250 low-lying mound (<1 meter high) at approximately wide), meters of (tens broad a including 2006, in 2004 (Hyland et al. 2004) and at three other sites surveys, observed on a ROV rock ledge in During 271 meters of water in NMS. – coast Pacific and U.S. the off coral stony structure-forming of Columbia, discoveries recent more the British of one includes and Alaska between boundary maritime the to north CA Pt. Conception, from extends which Province, Oregon The Oregon Province habitats downto90meterswaterdepth. californicus is province the in occur to known coral from 44 to over 1500 meters. The only stylasterid PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF tlse californicus Stylaster is found on Cordell Bank and south to San ohla pertusa Lophelia , which is found off central California at B. elegans wih a be rcre i rocky in recorded been has which , Paracyathus stearnsii Paracyathus p ws lo bevd in observed also was sp. Balanophyllia elegans Balanophyllia L. pertusa aypyla arnoldi Caryophyllia Stylantheca petrograpta and Stylantheca porphyra, Stylantheca Stylaster venustus Stylaster t h Oypc Coast Olympic the at Paracyathus stearnsii (Brancato et al. 2007), and Labyrinthocyathus . pertusa L. is common in common is . venustus S. s known is Errinopora at Cordell at Stylaster Lophelia , which ,

was were two ) ,

was covered with thick, hummocky sediments at sediments hummocky thick, with covered was 200 and 250 meters, where hard rocky substrate between depths at observed were gorgonians of densities high mid-1990s, the in coast Oregon the off dives submersible During 2003). Morgan over 2900 m off San Francisco, CA (Etnoyer and They have also been recorded from water depths 3.6). (Figure Mendocino Cape of north common more also but surveys trawl regional by covered depths all at entire common are and Gorgonians province region. the in taxa antipatharian and 3.5). (Figure slope the with Mendocino largest Cape reported catches from of the northern Oregon north most are prevalent Province Oregon the in corals Black comm.). Puget Sound (Fisher 1938; Alberto Lindner, pers. in and VancouverIsland of part southern the off province the of part northern the in only found is commercial fishing operations. Unfortunately, and surveys operations. trawl fishing bottom commercial during invertebrates fishes other and with collected been have to addition In 2001). Auster 2006; those al. 2005; Pirtle 2005; Parrish 2004; Syms of et and Jones Tissot nature (e.g., the relationships investigated even studies some have and 1991) al. et Hixon 1994; al. other et Hardin 2001; Brodeur 2002; Wing and Krieger and corals structure-forming invertebrates fishes, (e.g., Stone 2006; demersal between associations fine-scale report Pacific north the in Several studies both in the region and elsewhere VI. been observedindensegroves. and habitats sedimentary prefer they Province, Diego San the in As distributed. and but are specimens Most 3.9). (Figure depth and latitude by both region the throughout (Figure 3.8). slope Pennatulaceans the are ubiquitously on distributed only but latitudes all over coral, soft The height. in cm 30 exceed not did sites these organisms encountered), gorgonians observed at all of (21% observed taxa invertebrate abundant second-most the being Despite 2006). (Strom banks rocky submerged of edges southern the Anthoptilum grandiflorum Umbellula Antipathes CORAL COMMUNITIES SPECIES ASSOCIATIONS WITHDEEP Anthomastus sp. are also abundant and widely and abundant also are sp. p ae h ms common most the are sp. in situ in Chrysopathes p, a be caught been has sp., observations, corals observations, , Ptilosarcus Stylatula Stylatula

speciosa

gurneyi has sp.,

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION AFSC, and SCCWRP trawl surveys. AFSC, and SCCWRP PACIFIC COAST PACIFIC Figure 3.9. Map showing locations of pennatulaceans (Order Pennatulacea) from NWFSC, Specific identities from trawl survey catch records are unconfirmed and primarily limited to genus or family level. bottom trawls are of limited precision because consist of partial colonies or skeletal fragments, they often extend over kilometers of seabed, making it difficult to ascertain the size and overall traversing a variety of low-relief habitats. health of the organism. Consequently, the best Furthermore, trawl gears are not designed to source of data on species associations comes target sessile invertebrates. Therefore, catches from direct observations via submersibles or of corals only represent presence data, and other in situ photographic methods (e.g., ROVs). cannot be used to develop standardized indices of Three studies in particular have examined the coral abundance. Lastly, catches of corals often nature of relationships between deep corals and

123 124 PACIFIC COAST (e.g., dianthus two between Caryophyllia – corals existed cup other correlations positive scleractinian, correlation coral, cup positive the of that was taxa two between reported second-strongest Conception. The Pt. spatial off features the bottom hard relief examined high- and (1994) low- both on epifauna of distributions al. et Hardin basketstars, crabs, sponges, brittlestars, anemones,algaeandsalps. crinoids, with corals ( black fishes, to addition In m). (1.0-5.5 small not were them between distances median the existence of functional relationships because the questioned authors the habitats, similar in co-occurred invertebrates large and species fish six these commercially. Although targeted been several subgenus, the and of members rockfish bank cowcod, including subgenus rockfish Sebastomus the of members other and ( rockfish pinkrose m), 1.5 ( ( rockfish cowcod included ( transects along elsewhere of than sponges and corals frequencies large near occurrence higher associations. exhibiting epifaunal species Fish of evidence showed fishes. None of the (3.1%). 27 observations of gorgonians sponges Only by 1.3% of all black coral associations involved followed (6.8%) crinoids mostly were epifauna and them to attached on or lying organisms other had observed corals black 135 the of 15% Approximately sand. and mixed pebbles cobbles, relief, boulders, including areas low rock in highest and were gorgonians corals of black Densities province. the to date in relationships fish-invertebrate of comprehensive study most the includes 2006) (Tissot al. et California southern off banks rocky 11 from observations submersible of analysis Data San DiegoProvince context. et al. in prep) are currently being analyzed in this Tissot 2007; al. et Brancato (e.g., surveys recent video other though 1994), al. et Hardin 2005; other fauna in the region (Tissot et al. 2006; Pirtle PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF Antipathes S. ensifer levis Sebastes empylm dianthus Desmophyllum mhatu californica Amphianthus included galatheid crabs and anemones . rufus S. ; 1.3 m), shortbelly rockfish ( p Ohr soitos with associations Other sp. 14 ) Ms o tee species these of Most m). (1.4 p) ee bevd n association in observed were sp.) ohla pertusa Lophelia ; median distance = 5.5 m), bank m), 5.5 = distance median ; 10 ) sodpn rockfish swordspine m), 1.0 ; aaytu stearnsii Paracyathus . simulator S. Sebastomus n te colonial the and . h authors The ). Moderately . S. jordani 17 m) 1.7 ; have , and D. ; gobies and combfishes were observed more observed often were near gorgonians than expected combfishes by chance in and gobies adult rockfish, juvenile in low- entomelas ( Widow bedrock. and mud occurred of habitats relief Subselliflorae) suborder of while relief varying ( of pennatulaceans habitats rock observed mixed were in gorgonians meters, 250 structure-forming to 55 from depths water at examined dives submersible During between (2005) fishes. demersal and invertebrates megafaunal Pirtle Bank, relationships Cordell At Oregon Province taxa. among relationships functional not any of and indicative currents), to tocommon orientation relief, due (depth, likely habitat their of attributes physical the to affinities most are significant, suggest that these correlations, while statistically xml, h rcn RV uvy f deep-sea of survey ROV recent the example, For may associations. species NMS additional Coast elucidate Olympic the in and Canyon situ in Recent corals. for deep the with analyzed associations be species in to surveys yet have underwater Province Oregon from data much prep), the (in of al. et Tissot and (2005) al. et DeVogelaere(2005), Pirtle of exception the With brittle starsandanemones. crinoids, stars, basket crabs, shrimps, isopods, sp.). Coral epifauna included polychaete worms, ( octopi and cucumbers sea clams, corals, along with sponges, other corals, sea stars, ( ( Rattails 2005). al. et (DeVogelaere current the towards ridges and often oriented to maximize surface area on primarily here observed were Corals meters. 1250 at crest its to seafloor depth water meters 3600 surrounding at the from Cordell rises Davidson to contrast Bank, is a much deeper submarine environment. in Seamount, Davidson contact. physical direct involved none however, adistance m; 1 of within were between fishes and associations gorgonians all of half than Greater ( pennatulaceans near observed and greenspotted rockfish ( dominated habitats, flatfish, poachers, combfishes habitats where they occurred. In sand- and mud- Sebastolobus studies at Davidson Seamount, Monterey Seamount, Davidson at studies Coryphaenoides , oy ( rosy ), p) ee bevd daet to adjacent observed were sp.) Ptilosarcus . rosaceus S. Sebastomus p) n thornyheads and sp.) S. chlorostictus p ad members and sp. ad unknown and ) Ptilosarcus , black-eyed , Graneldone ) were sp.). S. STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Table 3.3. Ratings of potential fishing gear impacts to deep corals off the demersal species both on the U.S. Pacific coast. Each measure of impact is rated as High (H), Moderate shelf and upper slope. Vessel (M), or Low (L). sizes range from 35 to 95 feet Measure of Impact and average 65 feet (NWFSC 2006). Off California and Oregon, an open-access fishery targeting nearshore species is Gear Type comprised of vessels from 10 to 50 feet in length (NWFSC 2007). Region Geographic Gear Impact Charter boats also operate Extent of Use in Extent of Impact Overall Rating of

Severity of Impact out of numerous coastal ports Bottom trawls H H H H and target a variety of pelagic and demersal species. Gear Midwater trawls L L H L types used in the region include Demersal seines L M L L bottom trawls, midwater trawls, Bottom longlines & gillnets M M M M demersal seines, pots, bottom- set gillnets, bottom longlines Pots & Traps M L L L and other hook-and-line gear, COAST PACIFIC Other hook and line L L L L but bottom trawls are the most widely used and potentially coral assemblages at the OCNMS (Brancato et harmful to corals. The degree of impact to corals al. 2007) revealed many species of fishes and from fishing operations depends on the physical large invertebrates (e.g. shrimp, brittle stars, attributes of their habitat (e.g., sediment type, crabs) nestled among the coral structures (see relief), attributes of the gear (e.g., configuration, also. However, the degree to which corals might mode of operation, footprint) and its geographic contribute to the feeding, growth and reproduction extent of use (Table 3.3). These attributes will be of demersal fishes or provide biogenic structure detailed in relation to each gear type used in the for other megafaunal invertebrates is largely region. unknown for most of the Oregon Province. Bottom trawls Bottom trawls are the most widely used fishing VII. STRESSORS ON DEEP gear off the Pacific coast. They are used off CORAL COMMUNITIES Oregon and in federal waters off Washington and California to target numerous species of demersal Compared to other regions in the U.S., the fishes, shrimp, prawns, sea cucumbers and sea Pacific coast from California to Oregon hasa urchins. Bottom trawls also have the greatest narrow continental shelf, which may result in severity ranking of all gear types used in the coral communities here being more susceptible region (Morgan and Chuendpagdee 2003). Gear to coastal activities. Some activities that may components that contact the seafloor include the adversely effect corals include oil and gas doors, bridles, footrope (except in shrimp trawls) development, deployment of gas pipelines and and occasionally the netting (PFMC 2005). Gear communication cables, and marine pollution. configurations depend on target species and However, fishing operations, particularly operating depths, with door separation distances bottom trawling, pose the most immediate and ranging from 34-50 m for shelf trawls and 50- widespread threats to deep coral communities. 200 m for slope trawls (PFMC 2005). Footrope lengths commonly range from 15-34 m for shelf Effects of fishing fisheries. Due to their weight and speed (1.5-2.5 knots) over the seafloor, all trawl components that The temperate, nutrient-rich waters of the contact the seafloor have the potential to snare, California Current support lucrative commercial, undercut or topple emergent structures, including tribal and recreational fisheries. Fishing operations deep corals. The Pacific Council Groundfish in the region are very diverse. A limited-entry Essential Fish Habitat (EFH) Environmental trawl fishery operates from the Canadian border Impact Statement (EIS) identified sensitivity south to Morro Bay, CA targeting numerous of coral habitats to trawl gear as relatively high

125 126 PACIFIC COAST ein ewe uut 01 n uut 2004 August and 2001 August the between in region bycatch coral observed of 92% over for accounted trawls bottom Furthermore, metrics.” many control sites greater than 50% in most measured of and impact losses between differences with organisms large as such boulders; structure, re-arranged bottom as defined in three changes with 0-3 “major of scale a on (2-3) PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF iwtr rws r ue etniey n the in extensively used are trawls Midwater Midwater trawls (Table 3.3). impact of rating high a given were they gears, these to sensitive particularly are habitats extensively throughout the region and deep coral used are trawls bottom Because 3.10). (Figure

Figure 3.10. Map showing locations of deep coral bycatch recorded by fishery observers in the estW Coast Groundfish Observer Program. All observed trips and gear types from August 2001 – August 2004 were queried. Due to limitations of specific identifica- tions, coral bycatch was grouped into two classes: 1) gorgonians and stony corals, and 2) pennatulaceans. Point symbols repre- sent start locations of bottom trawls or longline and pot sets. STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Oregon Province to target Pacific hake (Merluccius and Chuendpagdee (2003), bottom longlines productus). Hake is a schooling, pelagic species and gillnets have a medium to low impact on the and therefore vessels using midwater trawls try physical and biological components of habitat. to avoid contact with the seafloor. As evidenced Furthermore, from August 2001 through August by no records of coral bycatch from this gear type 2004, gillnets and longlines accounted for less between August 2001 and August 2004 and their than 4% of all observed coral bycatch. However, very low rating of habitat impact (Morgan and this low percentage may reflect a sampling bias Chuendpagdee 2003), midwater trawls have low to mobile gears or that corals damaged by these impacts to deep coral communities in the region gears do not reach the surface. Lost longline gear (Table 3.3). has been observed on the seafloor at several sites within the Olympic Coast NMS (Hyland et Demersal seines al. 2004; Brancato et al. 2007). Nevertheless, Demersal seines, also known as Scottish seines, sensitivity of coral habitats to hook and line gear are used in nearshore and shelf areas to catch (e.g., longlines) was classified as low to moderate flatfish (e.g., sand dabs, Petrale sole, English (0.3-1.3) on a scale of 0-3 (PFMC 2005). Due to sole) and chillipepper rockfish (PFMC 2005). this low to moderate sensitivity rating, medium to Demersal seines use a large net attached to low habitat impact rating, and their moderate use long (hundreds of meters) ropes to herd fish in the region, bottom longlines and gillnets were COAST PACIFIC on the seafloor. In contrast to trawl gear, their given a medium rating of overall impact (Table lighter weight and slower movement over the 3.3). seafloor cause little disturbance (PFMC 2005). Corals most likely to be impacted by this gear Other fishing gears are pennatulaceans, because demersal seines Throughout the region, pots (also called traps) are used in areas of unconsolidated sediments are used to catch sablefish, Dungeness and where pennatulaceans inhabit. Due to their other species of crab, spot prawns, spiny limited use in the region; however, demersal lobster and other finfish. Pots can be single seines were given a low rating of impact (Table (e.g., Dungeness crab) or in a series of up to 50 3.3). Furthermore, there are no records of deep attached to a groundline (e.g., sablefish) (PFMC coral bycatch from this gear type. 2005). The effect of pots on the seabed depends on their weight, shape and lateral movement Scallop dredges during retrieval. If the vessel is not directly above Although dredges were once used to target the pot being retrieved, significant contact with weathervane scallops, they are now prohibited in the bottom can occur; however, this movement the region. is typically minimized so as not to put excessive strain on the line and other equipment (PFMC Bottom longlines and gillnets 2005). Severity of impact to coral habitats by Bottom-set gillnets are prohibited offshore pots and traps was ranked as low to moderate Washington and Oregon and in California state (PFMC 2005; Morgan and Chuendpagdee 2003). waters. In federal waters off California, gillnets Factoring in their limited use in the region, pots are used to catch white seabass, bonito, flying were given a low overall rating of impact (Table fish, white croaker, angel shark, California halibut, 3.3). lingcod, mullet and perch (PFMC 2005). Bottom longlines, on the other hand, are used throughout In addition to longlines, other types of hook and the region to target sablefish, rockfish, Pacific line gear (e.g., stick gear, rod and reel, jig gears halibut, cabezon, lingcod or dogfish. Bottom and vertical longlines) are used in the region, longlines are composed of weights, hooks and but their impacts to deep corals are most likely a mainline that contact the seafloor. Gillnets minimal. Sinkers, hooks and lines can snare are anchored by weights and leadlines, which and damage corals, but the area of seafloor weigh about 100 pounds per 100 fathoms of line contacted is small relative to trawls and dredges. (PFMC 2005). These gears can travel significant Consequently, these hook and line gears were distances over the seafloor, particularly during given a low rating of impact (Table 3.3). retrieval when the vessel is not directly over the gear, snaring or undercutting emergent structures (e.g., corals). According to Morgan

127 128 PACIFIC COAST Caryophyllia including taxa, invertebrate four of density the in began at three platforms. They found a decrease drilling after and during before, epifauna bottom et al. (1994) documented the abundance of hard- Hyland rocks. any dislodge and equipment the cool to primarily hole well the down pumped is During drilling, a mixture of water, clay, and barite 2004). NOAA 1997; al. et Raimondi 1981; al. et of introduction (Cimberg materials re-suspended the or suspended through is oiland operations gas offshore from organisms filter-feeding other and corals to threat potential One 2006). (MMS spudded were wells development 23 and producing were leases active 79 of 43 2005, In been drilled at the 23 platforms in federal waters. had wells total 1290 2006, May 1 of As 1998). been erected since 1989 (Nevarez et al. have platforms no although sites, lease existing for comm.). place in are restrictions No pers. MMS, Romero, (John 2012 until affect in is waters federal in state California waters, and a in moratorium on new leases leases gas and oil new on place in currently is moratorium depths 2003, Figures. 3.11-3.13). al. A et water permanent (Love m 363 to 11 at from ranging Channel Barbara Santa the in and Pedro San Arguello, in Pt. off sited 3 currently are waters) waters, state federal 26 in of (23 total platforms A California. southern in platforms offshore from occurred has gas and oil of production 1958, Since Oil andGasExplorationExtraction the world. of parts other in and Ocean Pacific the throughout corals impact adversely to potential the has change climate global future, the In any. if impacts, localized potential these stressors about but they are likely to have only known Unfortunately, is little pollution. and communication cables and pipelines gas of deployment the include adversely corals may impact that activities human be to detrimental to corals shown operations, in the region. been Other has fishing gas development and oil by to caused sedimentation addition In Effects ofotherhumanactivities PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF sp.,afterdrilling began. Arguello, California.Imagesource:MMSPacificOCSRegion. forms andassociatedpipelinesintheSantaMariaBasinoff Pt. Figure 3.11. a einl ald ca osraoy n the in observatory ocean cabled regional a – system cable NEPTUNE the include operations laying cable Future comm.). pers. NMS, Coast in September 2006 (Mary Sue Brancato, Olympic portion of the Olympic Coast NMS was completed that connects the cable U.S. and Japan communication via the northeastern a of reinstallation A depth. water meters 1500 to down buried is and Warrenton, Oregon west-southwest over the shelf Alaska the United Fiber System – West, which include extends from installations cable Recent continental margin off In all three west coast states. 3.11-3.13). the of parts many transect routes cable addition, (Figures. offshore California in facilities southern connect side shore to pipelines platforms production gas Numerous Communication Cables Deployment ofGasPipelinesand Mapshowinglocationsofoilandgasleases,plat - STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION PACIFIC COAST PACIFIC

Figure 3.12. Map showing locations of oil and gas leases, platforms and as- sociated pipelines in the Santa Barbara Channel off southern California. Image source: MMS Pacific OCS Region.

Figure 3.13. Map showing locations of oil and gas leases, platforms and asso- ciated pipelines off San Pedro, Califor- nia. Image source: MMS Pacific OCS Region.

129 130 PACIFIC COAST northern California (Wheatcroft et al. 1997) and 1997) al. et (Wheatcroft California northern been conducted at sites off Oregon (Kulm et al. 1975), have rates accumulation and sediment of deposition Studies region. the in largest the margin include coastal rivers – continental the the to Columbia sediments being of sources Natural Desmophyllum dianthus – corals stony two include correlation that organisms filter-feeding of coverage the and negative a seafloor the of m 2 within flux sediment between The discovered flux. sediment authors suspended to relation in assemblages epifaunal of distribution spatial the Pt. examined off (1994) al. sites et Hardin hard-bottomCA, Conception, At exception. no is coast Pacific U.S. The throughout 1993). (Norse world communities the coral of health the to threat major a still is sedimentation taxa, some by ability this Despite 1995). (Reigl gravity and motion water on 1980), solely rely Shelton alcyonaceans while 1995; sediment (Reigl remove polyps their actively from to ability the have scleractinians that shown have experiments two example, For others. than vulnerable more are taxa some however, 1994); al. et Hardin 1983; 1974; Dodge and Vaisnys 1977; Dodge and Lang negative al. et (Dodge corals report on sedimentation of effects world the throughout Studies Sedimentation (Freiwald etal.2004). repair or placement during used often anchors or cable the heavy the by swaths larger along over or itself pipeline occur can Damage area. the in present if corals to impacts severe cause seabed, they cannot be buried and therefore may Where cables and pipelines be transect hard-bottom could they nearby,sediments. resuspended by smothered or buried located were corals If operations. fishing commercial with interactions avoid to m) 1500 (usually depths deep-water to down buried are pipelines and cables Where feasible, topic. this on effort of lack a to due be areas of known coral habitat. However, this may of region the in communication reports cables or no gas been pipelines transecting have There with theU.S.portiontobeinstalledlater. off Vancouverconstruction, Island, BCareunder route (Figure 3.14). Portions of the cable network the along located nodes instrumented 20 about with cables fiber-optic of network 3000-km a of consist will NEPTUNE Ocean. Pacific northeast PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF . Lophelia pertusa Lophelia and oltns n ep oa cmuiis Hyland communities. coral deep on pollutants of effects the addressed specifically that region the in study one only of aware are authors The Pollution causes. the region on the effects of any of these potential Unfortunately, region. there is little specific information the in in use widespread its and bycatch) of form the in gorgonians and pennatulaceans (mostly corals with interactions documented of its because risk highest Bottom the pose may trawling earthquakes). by turbidity (triggered and currents storms slumps, trawling, bottom include resuspension sediment of causes Other over broadrangesofdepthandlatitude. they did provide average annual deposition rates address impacts to corals or others filter feeders, 2002). Although these studies did not specifically central California (Lewis et al. 2002; Eittreim et al. n cniud viaiiy f e mnrl from minerals key of availability continued involved and depths operating and distance the of Because 2001). (Carney War Cold the during interest the economic provoked Pacific north central in aggregations dense of discoveries later discovered the originally during were They nickel copper. cobalt, and manganese, including minerals initiated. Manganese been nodules proposed are has ores neither containing but coast, been Pacific U.S. the off have operations mining Two 2001). impact (Carney habitat adversely seafloor of areas to larger potential the has mining mineral development, gas and oil to Compared Mineral Mining Stylasterids isnowprohibitedoff California. been of harvest also commercial However, 2005). (NMSP shops shell has in sale for commercially and harvested death, collectors after recreational californicus pigment its Stylaster retains it Because Coral Harvest pollution ondeepcoralsremainuncertain. of effects the world, the throughout and region particulate increased loading. Because of the to lack of information in the due were epifauna toxic to changes biological below any authors that were suggesting The levels, concentrations that CA. Arguello, found Pt. the off on benthos discharges drilling from chemical contaminants of effects the examined (1994) al. et M Challenger HMS s oua among popular is oae although voyage, STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

lower than preindustrial values) and is predicted to decrease carbonate concentrations in the deep ocean (Orr et al. 2005). Corals use carbonate ions to form skeletal components. Two forms of calcium carbonate are common in the oceans: 1) aragonite, used by scleractinians and most stylasterids, and 2) calcite, used by octocorals and about 10% of stylasterids (Cairns and Mcintyre 1992). Aragonite is more soluble than calcite; therefore its depth of saturation (i.e., aragonite saturation horizon (ASH)) is shallower than that of calcite. Furthermore, the ASH in the north Pacific occurs at shallower water depths than the north Atlantic, resulting in dissolution rates twice as high in the upper 1000 m (Feely et al. 2004). Guinotte et al. (2006) hypothesize that the paucity of deep-sea, bioherm-building

scleractinians in the north Pacific is a result of this COAST PACIFIC relatively shallow (50-600 m) ASH. The authors also suggested that stony corals in the north Pacific are already living in marginal aragonite saturation states. Off the U.S. Pacific coast, stony corals are known to occur at water depths down to 578 meters (Cairns 1994). One of the largest accumulations of colonial scleractinian observed thus far in the region (i.e., an extensive low-lying Figure 3.14. Map showing locations of proposed mound of Lophelia pertusa at the Olympic Coast cable routes and node locations for the NEPTUNE NMS) was at a water depth of approximately 250 project. Image credit:: Debbie Kelley, Univ. of Wash- m (Brancato et al. 2007). Guinotte et al. (2006) ington, Center for Environmental Visualization. predicted that the ASH off the entire U.S. Pacific coast would be shallower than 200 m by 2060, terrestrial sources, mining of the nodules was while changes in the calcite saturation horizon never initiated. These dense aggregations occur will be less pronounced (Orr et al. 2005). If outside the EEZ, however, so potential mining those predictions are correct, the distributions operations would not impact U.S. coral resources. of scleractinians in the region could be severely The second proposed offshore mining operation impacted. Stylasterids would most likely be less was for polymetallic sulfides. In 1983, the U.S. affected by changing ASH because they occur at Minerals Management Service issued a draft shallower water depths (<183 m). environmental impact statement for a proposed lease sale of a portion of the seabed on the Gorda Evidence suggests that global climate change Ridge (see Figure 3.2). Despite initial studies may pose other threats to corals. Corals are as to the impact of mining in the vicinity, there sessile filter feeders, most likely feeding on was lack of industry interest. Although it is not suspended organic matter that rains down known whether corals occur on Gorda Ridge, a from the surface or is transported by currents photographic study of the proposed lease site in (Kiriakoulakis et al. 2005). Because many of the 1986 revealed the presence of other suspension organisms that comprise this source of organic feeders (Carney 2001). material (e.g., coccolithophores, foraminiferans, pteropods) use carbonate to form protective Climate Change shells, reduced carbonate concentrations may Although oceans are moderating climate change impact nutrient availability for corals. In addition, by assimilating anthropogenic CO2 emissions, it rising atmospheric CO2 is increasing deep-sea is not without consequence to ocean chemistry. water temperatures (Barnett et al. 2005) and

Uptake of atmospheric CO2 is causing declining altering salinities (Curry et al. 2003), which may pH in ocean surface waters (already 0.1 units in turn cause changes in thermohaline circulation

131 132 PACIFIC COAST n vdo mgr cletd uig numerous information surveys. This ROV and submersible during collected imagery video and photographic and (USGS) University, State Oregon Survey and Geological States United the structural institutions, California, of state the by created maps geologic academic theoil and by industry collected surveys, profiles submersible reflection during seismic and industry oil the by collected samples sediment of extensive database an investigations, geologic collected targeted imagery sonar during large-scale mapping programs and during sidescan and data bathymetric swath including region coast Pacific the for exists information seabed of multitude A Mapping andResearch the region. in programs mapping and These research of synopsis brief a conservation. after below highlighted be habitat will measures coral of effect indirect the had have agencies state and federal recently, a number of measures adopted by these protections of corals have only been implemented specific Although respectively. (MMS), Service Management the Minerals Interior’s and of Department Program Sanctuary Marine NOAA’s National the by regulated outer are shelf the continental on and sanctuaries marine five within national occurring activities Other habitats. subtidal and intertidal to impacts have may that activities other with along states, individual by the regulated are Oregon) in shrimp waters pink (e.g., state within toWashington. solely occur that Fisheries California from Coast Pacific theU.S. off waters in federal resources fishery of management the for responsible are (NMFS) Council) and the National Marine (the Council Fisheries Management Fisheries Service Pacific The VII. or adverseinteractionshavebeenidentified. corals in the region. Thus far, no specific studies on deep on have information might species invasive effects the for search to continues NOAA Invasive Species al. et (Guinotte 2006). conditions environmental in changes such to susceptible particularly be may they environments, nutrient-rich steady-state, in (Joos et al. 1999). Because corals have evolved PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF RESOURCES AND HABITATS MANAGEMENT OFFISHERY feature of these maps is an associated layer of layer associated an is unique maps these A of feature (1999). scheme al. et Greene by characterization developed habitat of a deep-water version modified a on based coded were from tens of meters to a kilometer. Habitat types mud, sand and rock) at horizontal scales ranging (e.g., lithology surficial and canyons) and slope shelf, (e.g., structures physiographic delineating polygons habitat show maps The 2005). al. (Goldfinger et Washington and Oregon the for produced maps University State Oregon at Lab the while Active Tectonics Mapping Seafloor and 2005), Bizzarro and (Greene California for map at Moss Landing Marine Laboratories created the impact statement. The Center for Habitat Studies inform the recent Groundfish EFH environmental to commissioned were projects complimentary Two 3.15). (Figure coast Pacific the off depth) margin (i.e., intertidal out to ca. 3000 meters water continental entire the for habitats benthic of map regional a of creation the in used recently was the Olympic Coast NMS conducted assessments deep coral habitats in the region. In of monitoring Toregular no been has there date, 2004 and 2006, highlighted earlierinthis chapter. were that communities coral deep of discoveries communities. coral Many of these investigations have resulted in deep the document survey to ROV efforts future guide to 2006) (Intelmann continuing its deep sea habitat mapping program also is NMS Coast Olympic 3.16). (Figure NMS Coast the Olympic in features seafloor bottom along the Oregon coast, reefs and Astoria shelf Canyon, outer and the hard on banks rocky Bank, Cordell Canyon, Monterey Seamount, Davidson outcrops off southern California, the Big Sur coast, and banks rocky several include scan) line laser sonar,multibeam and sidescan profiling, seismic (e.g., techniques mapping geophysical remote with ROVs) submersibles, sleds, camera (e.g., have that platforms observational underwater incorporated areas Study benthic of investigations. variety habitat a for maps base provide to coast thePacific off projects mapping seafloor targeted additional conducted have institutions Several federal and state agencies and academic area outtoca.3000meterswaterdepth. km (21,000 9% approximately habitat comprise to estimated these are habitats From rocky maps, press). in al. et (Romsos mapping habitat to utility their for sources data data quality that quantifies data density and ranks 2 o te seafloor the of ) STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION ------

- - Figure 3.15. Map of Pa cific coast of the U.S., showing benthic habitats from the coast line out to ap 3000 proximately meters water depth. Benthic habitat charac terization was modified after Greene et al. incor and (1999) porates informa tion on seafloor lithology (e.g., sedimentary or physi and rocky) ography (e.g., shelf, slope, basin, canyon wall). Maps of and Washington Oregon habitats were created the Tecton Active ics and Seafloor Mapping Lab at Oregon State Map University. hab California of itats was created by the Center for Habitat Studies at Moss Landing Marine Labora tories. PACIFIC COAST PACIFIC

133 134 PACIFIC COAST STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF Figure 3.16. Barbara). Image source:Wakefield etal.2005,Figure 3. Laboratories; MBNMS,MontereyBayNational MarineSanctuary; and UCSB,UniversityofCalifornia, Santa ment ofFishand Wildlife;CBNMS,Cordell BankNationalMarineSanctuary; MLML,MossLanding Marine tory; WSUV, Washington StateUniversity, Vancouver; OSU,OregonStateUniversity; ODFW, OregonDepart Science Center;SWFSC,Southwest FisheriesScienceCenter;PMEL,Pacific Marine EnvironmentalLabora ment ofFishandWildlife;OCNMS, OlympicCoastNationalMarineSanctuary; NWFSC, NorthwestFisheries the participating organizations conducting interdisciplinary studies of fish habitat (WDFW, Washington Depart MapofthePacificcoast of theU.S.,highlightinggrowingnetworkstudyareas andsomeof - - - STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION of deep coral and sponge communities, but it focus largely on pristine or untrawled habitat is unclear how future fiscal environments will and biogenic habitats such as corals that are affect regular monitoring efforts. The Southwest vulnerable to impact from human activities. The Fisheries Science Center (SWFSC) often plan is considered comprehensive because it conducts submersible surveys of hard-bottom addresses impacts from the full range of human habitats off California. Although the primary activities (fishing and non-fishing), and includes focus of these surveys is groundfish habitats, they procedures for adaptation as new information often occur in areas where corals are located and becomes available. have therefore resulted in important discoveries, including that of the newly described black coral, Management measures to minimize adverse Antipathes dendrochristos. Other surveys by the impacts from fishing include marine protected Northwest Fisheries Science Center (NWFSC) areas, reductions in fishing effort, and gear such as those utilizing AUVs are just being restrictions as recommended by the National developed as a routine method for groundfish Research Council (NRC 2002). Over 130,000 mi2 and may provide more routine monitoring of deep (336,700 km2) are now marine protected areas coral habitats coastwide. (MPAs) and fully protected from impacts from bottom trawls, with selected vulnerable habitats

Directed Harvest protected from all fishing gears that contact the COAST PACIFIC Presently, there is no directed harvest of corals bottom (Figure 3.17). The MPAs are distributed in the region. the length of the coast and include both federal and state waters. The MPAs work in concert with Minerals Management Service other spatial management actions taken by the The United States Minerals Management Council such as the Rockfish Conservation Areas Service (MMS) is responsible for the regulation and Cowcod Conservation Areas (Figure 3.18) of development and extraction of offshore that have significantly reduced fishing effort in energy resources, and they regularly conduct habitats important to the adult life-stage of some research in the Pacific outer continental shelf overfished species. Some of these habitats region. Mitigation of potential impacts to the are hard-bottom areas that may include deep nearby marine environment from oil and gas corals. Fishing effort has also been significantly development has been a priority of MMS since oil reduced off the central California coast through a production began of southern California in 1958. collaboration of the fishing industry and the Nature Because of the potential risks from offshore oil Conservancy. A feature of this collaboration is the and gas operations, MMS developed anchoring private purchase of trawl fishing permits by the guidelines to minimize impacts to hard-bottom Nature Conservancy. Finally, gear restrictions communities. These guidelines were crafted as implemented by the Council include coast wide a result of numerous studies conducted before prohibitions on gear types known to have a high and after drilling projects (Mary Elaine Helix, impact on benthic habitats. Prohibited gears MMS, pers. comm.). In addition, a long-term include dredge gear, beam trawl and large monitoring program was conducted in the late footrope bottom trawl. 1980s to evaluate environmental impacts of oil and gas development on marine and coastal Impacts from non-fishing activities were resources (Steinhauer and Imamura 1990). It addressed by the Council through the was these studies that provided some of the first establishment of Habitat Areas of Particular in situ observations of stony corals in the region Concern (HAPC) and publication of recommended including Lophelia pertusa, Desmophyllum conservation measures (PFMC 2005). NMFS is dianthus and Paracyathus stearnsii. now using the HAPC to focus their conservation efforts related to non-fishing activities through Fishery Management Councils the EFH consultation process. The Pacific Fishery Management Council, in cooperation with NMFS, has implemented a The Council and NMFS recognized that the plan comprehensive plan to protect EFH for groundfish to protect groundfish habitat is, in large part, (see PFMC 2005, 2006). The plan was developed based on precautionary management principles in collaboration with NGOs, the fishing industry, due to important gaps in available information. To and the National Marine Sanctuary Program to accommodate the likelihood that new information

135 136 PACIFIC COAST source: PFMC 2006. Essential Fish HabitatEISaspartofthe preferredalternativetominimize adverseimpactsto EFH.Image Groundfish fisherymanagement plan.Conservationareasweredevelopedin thePacificCoastGroundfish Figure 3.17. PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF Map showinglocationsofconservation areasspecifiedin Amendment 19tothePacificcoast STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION PACIFIC COAST PACIFIC

Figure 3.18. Map showing locations of some federal marine managed areas off the U.S. Pacific Coast, includ- ing national marine sanctuaries, the Cowcod Conservation Areas (CCA) and the trawl Rockfish Conservation Areas (RCA). The boundaries of the trawl RCA are adjusted on two-month management cycles in response to bycatch information on overfished species. Shoreward boundaries can be as shallow as the shoreline while seaward boundaries can be as deep as 250 fathoms. An area between the lines approximating the 100- and 150-fathom isobaths (shown on map) has been permanently closed to bottom trawling since inception of the trawl RCA in 2002.

137 138 PACIFIC COAST Fgr 31) Ecp i dsgae marine designated sanctuaries in the areas, conservation or reserves Except 3.18). California (Figure southern and central off located are Islands Channel and Bay, Monterey Farallones, the the of while Gulf Bank, Cordell sanctuaries, four Washington,other northern off located is NMS Coast fivenational Olympic The exist sanctuaries. there marine region Pacific the In National MarineSanctuaries Program. Technology and Research Coral Sea Deep the by identified been areashave corals in sea deep corals where sea deep of protection the for zones designate to authority discretionary have between Councils The corals. sea deep interactions and gear fishing reducing in participants industry fishing assist to technologies develop and research conduct occur, to likely or known are corals sea deep where locations in activity monitor corals, sea deep of locate locations identify, map and will Research Coral Program Technology Sea and Deep The zones. deep coral of designation the Technology allowed and and Program Research Coral Sea Deep a 109-479), Congress directed NOAA to implement and Management Reauthorization Act of Conservation Fishery 2006 Magnuson-Stevens the (P.L.In to process adapt habitatprotectionmeasures. the streamlined a research, established through Council available become will PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF nArl 03 y h sae f aiona (Figure California 3.19). of After implementation in July state 2007, the new the by 2003 April in implemented reserves an state of network complement existing to 3.19) (Figure NMS Islands Channel the within area conservation one and Sanctuary Marine National Program established the a network 2007, of marine reserves May In outside ofdesignatedconservationareas. operations fishing normal during except m), (91 of benthic organisms in waters above 50 fathoms Cordell Bank NMS prohibits the removal or injury the addition, In anchoring. and activities fishing and 3) alteration of the seabed except for normal parts, bait, water or other 2) biodegradable effluents, production, fish except substances or materials of discharge and development exploration, corals, to including harmful but be not limited may to that 1) new activities oil, other gas or mineral gears. however,contact sanctuaries, All prohibit do not regulate fishing including the use of bottom in conservation areas (15 FR 29208). Proposed 29208). FR (15 areas conservation in and recreational take lobster of pelagic fishes will be allowed for be fishing certain will while activities prohibited extractive all reserves, state 232 nmi to areas conservation and reserves of area total to increasing thereby boundary, waters federal state- mile three the to reserves existing it’s of of California is expected to extend the boundaries 110.5nmi total will area conservation and reserves federal 2 and 8.6 nmi 2 and 1.7 nmi 1.7 and 2 , respectively. Like existing 2 , respectively. The state respectively.The , NMS. source: ChannelIslands in July2007.Image will beimplemented 2003 whilefederalMPAs implemented in April areas. StateMPAs were of marineprotected (CINMS) anditsnetwork tional MarineSanctuary Channel IslandsNa showing locationofthe Figure 3.19. Map - STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION rules for marine zoning in the Monterey Bay and The time frame for completion will depend on Cordell Bank NMS are expected in the near available funds. future. Oregon also suffers from lack of seabed State activities information in state waters. To date, less than The state of California in recent years has enacted 5% of Oregon’s ca. 950 nm2 (3263 km2) territorial a series of laws to direct the management of sea is mapped. In March 2006, a consensus state marine resources, including the Marine statement, signed by 20 Oregon marine scientists, Life Management Act (Stats. 1998, ch. 1052), called for support and funds to map the seafloor the Marine Life Protection Act (MLPA, Stats. within Oregon’s territorial sea. Projected costs 1999, ch. 1015), and the Marine Managed are under $6 million. Stated reasons for this Areas Improvement Act (Stats. 2000, ch. 385). mapping plan include management of hazards In particular, a key mandate of the MLPA is to posed by tsunami events, describing nearshore design and manage a network of MPAs to habitats on which nearshore fisheries and protect marine life, habitats, ecosystems, and marine resources depend, and scientific support natural heritage while providing recreational, of two gubernatorial proposals – establishment educational and study opportunities. In addition of a limited network of marine reserves in state to the marine protected areas implemented in waters, and a national marine sanctuary to be COAST PACIFIC 2003, the state, in April 2007 adopted a network sited off Oregon. In response to the scientific of 29 MPAs for their Central Coast Study Region, consensus statement, the Territorial Sea Mapping which encompasses state waters between Pt. Bill (HB 2924), which would appropriate funds for Conception and Pigeon Point (Figure 3.20). seafloor mapping, was introduced in February These include new and expansions of existing 2007. Passage of HB 2924 along with the MPAs that cover approximately 204 mi2 (528 establishment of reserves and a national marine km2) or 18% of state waters in the central coast sanctuary are still pending. region, including 97 mi2 (251 km2) of no-take reserves (Office of Administrative Law Notice File In September 2006, the three west coast governors Number Z06-1031-05). While corals (specifically signed a joint agreement to collaborate on critical hydrocorals) are listed as “benefit species” for ocean and coastal protection and management only three of the MPAs, the protection of both issues. Short-term priorities for this collaboration shallow and deep hard bottom habitats is an include increased funding for mitigation of objective for this network of MPAs. Planning nonpoint source pollution, opposition to oil and and scoping of additional MPAs began in early gas leasing, exploration and development, and a 2007 for the next study region, the North Central regional research plan to support ocean observing Coast, which encompasses state waters from programs and seafloor and habitat mapping. Pigeon Point north to about Pt. Arena.

To date, only about a third of the seafloor in VIII. REGIONAL PRIORITIES TO California state waters has been mapped at any UNDERSTAND AND CONSERVE DEEP appreciable scale (Rikk Kvitek, CSUMB, pers. CORAL COMMUNITIES comm.). The Seafloor Mapping Lab of California State University Monterey Bay (CSUMB) and Given the limitations of existing information off the its partners are currently engaged in the North U.S. Pacific coast, it is clear that more targeted Central Coast Mapping Project, which will produce data collections and mapping efforts are needed. multibeam base maps for state waters seaward Because many collections are made from long of the 20-meter isobath from Año Nuevo north to trawls that can traverse several habitats, it is Pt. Arena. Another project mapping the coastal impossible to determine specifically the habitat area in the Santa Barbara Channel involves the from which these species were collected. Seafloor Mapping Lab, USGS and the California Therefore, to date, it is difficult to map corals at Ocean Protection Council. Over the next 5-6 a regional scale showing the appropriate habitat years, the California Coastal Conservancy and associations. Furthermore, coral specimens are Ocean Protection Council hope to complete continuously collected during in situ photographic comprehensive mapping for the remaining 66% of surveys, regional trawl surveys and by fisheries state waters (Rikk Kvitek, CSUMB, pers. comm.). observers. Rapid identification of these samples is

139 140 PACIFIC COAST Game. and marinerecreationalmanagement areas(SMRMA).Imagesource:California DepartmentofFishand of California,includingstate marineconservationareas(SMCA),parks(SMP), marinereserves(SMR) Figure 3.20. PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF Mapshowinglocationsof marine protectedareasintheCentralCoaststudyregion forthestate STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION needed to mitigate further impacts to deep corals identification of corals and potentially other and their habitats. Finally, the establishment of difficult to identify structure-forming invertebrates a network of conservation areas in the region from the region’s ongoing trawl surveys and provides an unprecedented opportunity to observer program and will provide validations monitor coast wide recovery of benthic habitats of identifications during targeted deep coral (including those that support coral communities) surveys. from fishing impacts. Deep corals are vulnerable to a variety of Mapping activities including oil and gas exploration and Because little is known about the nature of fishing; however, little is known about the potential relationships between corals, other invertebrates recovery rates of these species. There are a and demersal fishes off the Pacific coast, the variety of ways to monitor recovery. For instance, highest mapping priority is to quantify those age and growth information can be obtained relationships. To date, few studies (see Tissot through a combination of biochemical studies, et al. 2006; Pirtle 2005; Hardin et al. 1994) 14C and other radiometric dating, stable oxygen in the region have examined the nature of isotope data from the corals themselves and relationships between corals, other structure- other biochemical analyses. In addition to basic forming invertebrates and fishes, though analysis age information, it should be possible to associate COAST PACIFIC of recent surveys is ongoing (see Brancato et al. growth stages of corals to environmental change. 2007). In order to evaluate the importance of Working with biochemists at collaborating corals to their benthic communities, future in situ academic institutions, the physiological potential surveys will need to incorporate a more holistic for recovery of deep corals via geochemical and investigation of species relationships and habitat biochemical analyses of age and growth as well characteristics. as growth potential can be investigated.

Surveys are needed that specifically monitor Other gaps in information result from the fact the abundance and distribution of corals in that few targeted surveys have been conducted representative habitat types so that accurate in the region. Nearly thirty species of stylasterid and comprehensive maps of their distribution corals have been observed in waters off Alaska and abundance can be made. Targeted surveys including the Aleutian Islands coral gardens and on representative habitats using underwater parts of the Bering Sea and Gulf of Alaska (Stone platforms (e.g., submersibles, ROVs, AUVs) in and Shotwell, Chapter 2). In contrast, only five conjunction with optical survey systems (e.g., species of stylasterids have been observed video, laser line scan) are needed. Survey areas in the Oregon Province with an additional two can be selected using maps of surficial geologic species found off British Columbia (Appendix habitats that were developed for the Pacific coast 3.1, Jamieson et al. 2006). This may be a result Groundfish EFH EIS (see Figure 3.15) or higher of sampling bias; nonetheless, more focus needs resolution maps of specific areas. In addition, to be given to the shallower depths where these areas of repeated coral bycatch identified during taxa are commonly found. Also, reef-building mapping of existing observer and trawl survey scleractinians (e.g., Lophelia pertusa) while data, should be intensively mapped to establish common in the San Diego Province do not form the extent of these “hot spots.” expansive reefs like those in the north Atlantic. While ocean chemistry may be a factor (see Research Guinotte et al. 2006; Orr et al. 2005), it’s clear, Coral information can be collected ancillary to especially with the recent discovery of a Lophelia other survey and monitoring activities, however mound in the Olympic Coast NMS that more it is difficult for non-specialists on these surveys attention needs to be given to these taxa. to provide specific identifications. Collaborations with systematics experts at museums through Although more research on deep coral a pilot project in the NWFSC’s Genetics and communities is needed, data mining opportunities Evolution Program to develop molecular methods do exist. Since the late 1980s, underwater vehicles for the rapid identification of Pacific coast and camera sleds have been used to survey corals are already in place. The development the benthic environment off the Pacific coast. A of this capability will provide for species-level product of these surveys is hours upon hours of

141 142 PACIFIC COAST part, corals in the region do not build reefs with reefs build not do region the in corals part, most the For observers. fishery by and surveys been also trawl bottom of catch the in have wide coast recorded gorgonians and corals black coral, dendrochristos tree Christmas discovered recently the Yoklavichincluding 2005), Love and 2006; al. et Tissot 2007; al. et (Love California 2007) and on numerous rocky banks off southern 2005), in the Olympic Coast NMS (Brancato et al. 2005), in Monterey Canyon, at Cordell Bank (Pirtle coral discovered at Davidson Seamount (DeVogelaere deep of amount habitat. Significant coral communities have been considerable a hosts Washington to California from coast Pacific The IX. • • • • • Summary ofresearchpriorities: for EFH as corals demersal fishes. deep evaluating for basis informational the providing as well as California, deep corals and of associated fishes abundance and habitats and off distribution the georeferenced on database a in resulted has collaboration This University. State Washington at ecologists fishery biologists at the SWFSC and invertebrate is a longterm and productive collaboration among invertebrates structure-forming other and deep corals of surveys visual past from data existing ecology rescuing of example their great A 2006). Strom into (see insights and corals deep of observations additional provide may consuming, time- while videos, these of Review data. video PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF Data mining of video records from numerous from records video of mining Data rocky shallow in effort survey more Focus and corals of rates recovery Investigate specimens coral of identification Timely of nature the describe and Quantify species associations. and habitat identify significant any to and rocky corals deep region of the throughout surveys features ROV and submersible to coastalactivities. habitats where corals may be more susceptible stressors, of variety particularly fishing. a from habitats their fisheries observers. by and surveys trawl region during collected other with fishes. corals deep structure-forming invertebrates and demersal of associations CONCLUSION (Opresko 2005). Pennatulaceans, Antipathes RV) n mr rcnl, autonomous these recently, however, (AUVs); more vehicles underwater and (ROVs) vehicles collected operated remotely submersibles, been using a has in scope, but geographic information, limited detailed the More these on within species. methods collection the information of limitations mapping provide will continued and ongoing are surveys Some these of surveys. trawl was bottom taxa during collected coral higher-level of zoogeography regional the on information recent the of Much investigate andquantifytheserelationships. taxa. More targeted studies are needed to further of these represent obligate relationships between been reported, there is have no fishes direct evidence and that invertebrates any other with corals of associations Although NMS. Coast Olympic and Province Diego San the in stony reef-building two corals - only of observations huh sal i aes hr de corals these deep surveys, trawl where Unlike found. areas are in usually fishes, demersal though on focus often most surveys (e.g., trawl marks in sediments, derelict gear) in gear) derelict sediments, in marks trawl (e.g., seafloor the with interactions fishery of evidence Additional observers. fishery by reported been has widespread corals to impacts fishing of evidence Direct namely fishes. demersal target to trawls fishing, bottom of use commercial is and region the in significant habitats coral to most threat immediate the However, coral habitats. deep impact and shape both that events anactive of (e.g., subduction) prolonged and episodic (e.g., turbidity currents) both is part with the margin, of coast continental coast Pacific eastern the the U.S., Unlike provinces. latitudinal extent encompassing two large distinct zoogeographic a and upwelling coastal shelf, continental narrow a include distributions coral Unique attributes of the region that may influence ongoing basis. Observers currently collect this information on an begleaned from information collected by fisheries can observers. impacts fishing monitoring as well Additional as corals of distribution the sites. on information these of monitoring routine on a repeated basis, and therefore do not provide conducted seldom are but habitats coral deep of fishes. mapping localized provide surveys photographic demersal and invertebrates other corals, deep between relationships information the on provide can surveys photographic Lophelia pertusa and Madrepora oculata L.

pertusa n situ In in the in n situ in

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION the vicinity of corals also is provided from in situ X. REFERENCES photographic surveys, such as those conducted recently in the Olympic Coast NMS (Hyland et al. Allen MJ, Groce AK, Diener D, Brown J, Steinert 2004; Brancato et al. 2007). With recent fishery SA, Deets G, Noblet JA, Moore SL, Diehl management measures (e.g., area closures, gear D, Jarvis ET, Raco-Rands V, Thomas C, restrictions), however, the risk posed by bottom Ralph Y, Gartman R, Cadien D, Weisberg trawling has been significantly reduced. Although SB, Mikel T (2002) Southern California conservation areas encompass many known coral Bight 1998 Regional Monitoring Program: habitats coastwide, recent discoveries of diverse V. demersal fishes and megabenthic coral communities outside of conservation areas invertebrates. Southern California Coastal (e.g., in the Olympic Coast NMS) warrant further Water Research Project, Westminster, consideration of protection. California

Allen MJ, Moore SL, Schiff KC, Weisberg SB, Diener D, Stull JK, Groce A, Mubarak J, Tang CL, Gartman R (1998) Southern California Bight 1994 Pilot Project:

V. Demersal fishes and megabenthic COAST PACIFIC invertebrates. Southern California Coastal Water Research Project, Westminster, California

Andrews AH, Cailliet GM, Kerr LA, Coale KH, Lundstrom C, DeVogelaere A (2005) Investigations of age and growth for three deep-sea corals from the Davidson Seamount off central California. Pages 1021-1038 in Freiwald A, Roberts JM (eds.), Cold-water corals and ecosystems. Springer-Verlag Berlin Heidelberg

Aspen Environmental Group (Aspen) (2006) Tranquillon ridge oil and gas development project: Environmental impact report, State clearinghouse no. 2006021055, Prepared for County of Santa Barbara Planning and Development

Auster PJ (2005) Are deep-water corals important habitat for fishes? Pages 747-760 in Freiwald A, Roberts JM (eds.), Cold-water corals and ecosystems. Springer-Verlag Berlin Heidelberg

Barnett TP, Pierce DW, AchutaRao K, Gleckler PJ, Santer BD, Gregory JM, Washington WM (2005) Penetration of human-induced warming into the world’s oceans. Science 309:284–287

143 144 PACIFIC COAST Cairns SD, Stanley GD (1982) Ahermatypic coral Cairns SD, Bayer FM (2005) A review of the genus the of revision generic A (1983) SD Cairns temperate the of Scleractinia (1994) SD Cairns Brodeur RD (2001) Habitat-specific distribution of Briggs JC (1974) Marine Zoogeography. McGraw- Intelmann J, Hyland CE, Bowlby MS, Brancato C (2005) Goldfinger SA, Heppell MA, Bellman Strub PM, Kosro JA, Barth HP, Batchelder PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF aia cnevto rglto via regulation patterns temporal and spatial of analysis conservation habitat Evaluation of a U.S. west coast groundfish 15:36-47 California Oceanography program. System Current Pacific Northeast The (2002) GLOBEC SR Ramp Mackas BM, Hickey DG, DL, Ainley FB, Powell Schwing LW, TM, Botsford MD, Ohman Tynan WT,CT, Peterson RD, Brodeur PT, Coral ReefSymposium: 1:611-613 International Fourth counterparts. the of fossil Proceedings and Living banks: 77:225-256 Science Marine of Bulletin species. new Gorgonacea: two of description the with Primnoidae), (Octocorallia: Primnoa of MarineScience33:427–508 Part Hydrozoa), 1, Description of the genera. Bulletin (Coelenterata: Stylasterina to ZoologyNo.557 Contributions Smithsonian Pacific. North Shelf Research21:207-224 Continental Sea. Bering Canyon, Pribilof ( perch ocean Pacific Hill BookCompany. New York, 475pp McArthur IICruise AR06-06/07, 46pp Ship NOAA Report: Cruise Washington. Olympic Coast National in Marine assemblages Sanctuary, sponge and coral deep of Observations (2007) K Brenkman SS, 2900 Fisheries and Aquatic Sciences 62:2886- of effort.Canadian Journal fishing trawl of Sebastes alutus Sebastes ) in ) Cimberg RL, Gerrodette T, Muzik K (1981) Habitat applicability Management (2001) RS Carney PA Rona DL, Stein GL, Taghon AG, Jr. Carey al A 18) n oe yrcrlia from Hydrocorallinae some On (1884) WA Dall Curry R, Dickson B, Yashayaev I (2003) A change Geology, (1987). ML Holmes DA, Clague S, Goffredi A, Davis R, Zierenberg DA, Clague, No. 601, U.S. Office of Marine Pollution Marine of Office U.S. 601, No. Alaska coral. Final Report, Research Unit requirements and expected distribution of Orleans, New Louisiana, USA,174pp Service, Management Minerals Interior the of 2001- Dept. MMS U.S. 095. Study OCS Report. studies. Final Mexico of Gulf of reevaluation and ecology deep-sea contemporary of zone. economic Springer, New exclusive York A States’ Ridge: the United in Gorda center spreading seafloor (ed) GR In: McMurray 225-240 Pages Valley. Axial Gorda Ridge the in fishes and megaepifauna benthic of ecology Distributional (1990) 115 2:111- Washington of Society Biological Alaska and California. Proceedings of the 426:826–829 Ocean over the past four decades. Nature Atlantic the of balance freshwater the in pp563-580 USA, Texas, Houston, Earth Series, Science Resources, Mineral for and Council Energy Baja Circum-Pacific to California. Sea adjacent basins—Beaufort and ocean America of North margin continental western the of potential resource and Geology A, (eds.) JG Grantz Vedder DW, Scholl In the Ridge. Gorda of potential mineral and petrology, Ridge. Gorda the RIDGE Events,11:5–12 to expedition 2000 MBARI’s (2001) K Damm Von S, Ross V,Orphan E, Olsen N, Maher J, McClain Assessment, Alaska Office,54pp STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Davis EE, Karsten JL (1986) On the cause Etnoyer P, Morgan L (2003) Occurrences of of the asymmetric distribution of habitat-forming deep sea corals in the seamounts about the Juan de Fuca Northeast Pacific Ocean. A report to Ridge: Ridge crest migration over a NOAA’s Office of Habitat Conservation. heterogeneous asthenosphere. Earth and Marine Conservation Biology Institute, Planetary Science Letters 79:385-396 Redmond, Washington, USA, 33pp

Department of Fisheries and Oceans Canada Feely RA, Sabine CL, Lee K, Berelson W, Kleypas (DFO) (2000) Hexactinellid sponge reefs J, Fabry VJ, Millero FJ (2004) Impact of

on the British Columbia continental shelf: anthropogenic CO2 on the CaCO3 system geological and biological structure. DFO in the oceans. Science 305:362–366 Pacific Region Habitat Status Report 2000/02 Fisher WK (1938) Hydrocorals of the North Pacific Ocean. Proceedings of the United States Deuser WG (1975) Reducing environments. National Museum 84:493-554 In Riley JP, Skirrow G (eds) Chemical Oceanography, 2nd ed., vol 3. Academic Fisk MR, Duncan RA, Fox CG, Witter JB

Press, pp1-37 (1993) Emergence and petrology of COAST PACIFIC the Mendocino Ridge. Marine and DeVogelaere AP, Burton EJ, Trejo T, King CE, Geophysical Research 15:283-296 Clague DA, Tamburri MN, Cailliet GM, Kochevar RE, Douros WJ (2005) Deep- Freiwald A, Fosså JH, Grehan A, Koslow T, sea corals and resource protection at Roberts JM (2004) Cold-water coral reefs. the Davidson Seamount, California, UNEP-WCMC, Cambridge, UK, 85pp U.S.A. Pages 1189-1198 in Freiwald A, Roberts JM (eds.), Cold-water corals Fosså JH, Mortensen PB, Furevik DM (2002) and ecosystems. Springer-Verlag Berlin The deep-water coral Lophelia pertusa in Heidelberg Norwegian waters: distribution and fishery impacts. Hydrobiologia 471:1-12 Dodge R, Aller R, Thomsom J (1974) Coral growth related to resuspension of bottom Greene HG, Bizzarro JJ (2005) Essential fish sediments. Nature 247:574-577 habitat characterization and mapping of the California continental margin. In Dodge R, Lang J (1983) Environmental correlates Risk assessment for the Pacific coast of hermatypic coral ( Montastrea annularis) groundfish FMP. Prepared for the Pacific growth on the East Flower Gardens Bank, States Marine Fisheries Commission, north west Gulf of Mexico. Limnology and Portland, Oregon, USA, Appendix 3 Oceanography 28:228-240 Greene HG, Yoklavich MM, Starr RM, O’Connell Dodge R, Vaisnys J (1977) Coral populations VM, Wakefield WW, Sullivan DE, McRea and growth patterns: responses to Jr JE, Cailliet GM (1999) A classification sedimentation and turbidity associated scheme for deep seafloor habitats. with dredging. Journal of Marine Research Marine benthic habitats and their living 35:715-730 resources: monitoring, management and applications to Pacific Island countries. Eittreim SL, Xu JP, Noble M, Edwards BD (2002) Oceanologica Acta 22:663-678 Towards a sediment budget for the Santa Cruz shelf. Marine Geology 181:235-248

145 146 PACIFIC COAST Hyland J, Cooksey C, Bowlby E, Brancato MS, Brancato E, Bowlby C, Cooksey J, Hyland Fish (1991) WG Pearcy BN, Tissot MA, Hixon system current California The (1979) BM Hickey of oceanography Coastal (1998) BM Hickey Hardin DD, Toal J, Parr T, Wilde P, Dorsey K (1994) Guinotte JM, Orr J, Cairns S, Freiwald A, Morgan R, Milstein R, Robison C, Romsos C, Goldfinger PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF itiuin f epsa scleractinian deep-sea of distribution the alter chemistry seawater in changes human-induced Will (2006) R George L, USA, Appendix 2 Oregon, Portland, Fisheries Commission, Marine States Pacific the for FMP. Prepared Washington, groundfish coast Pacific and Oregon 1.0. Version for maps lithology seafloor Interim (2005) B Myers SC, USA,13pp Charleston, Research, Biomolecular and Health Environmental Coastal NOAA/NOS for Center 15. NCCOS NOS Tech. Memo. NOAA 2. II Leg McARTHUR AR-04-04: Cruise Ship Cruise NOAA for Report (OCNMS). Sanctuary National Marine Coast Olympic the at impacts fishing/harvest to susceptibility assemblages their and of coral/sponge survey pilot deepwater A (2005) S. Intelmann California, USA,410pp Minerals Management Service, Camarillo, Interior the of Dept. U.S. 91-0052, MMS Study OCS Report. the Final Banks], Daisy of banks and Coquille, [Heceta, Northwest Pacific rocky of assemblages in Progress facts. Oceanography 8:191-279 and hypotheses – New Inc, York, Sons, pp345-393 & Wiley John 11. Vol. Sea, The (eds.) KH Brink AR, Robinson In: Island. Vancouver of to California tip Baja the from America North western Marine Environmental Research37:165-193 factors. physical of importance the California: Basin, Maria Santa the in epifauna hard-bottom in variation Spatial and Ecology in Environment 4:141-146 Frontiers corals? In ik seset o the for assessment Risk aisn S Plern , esn (2006) S Jessen N, Pellegrin GS, Jamieson Effort Mapping Habitat (2006) SS Intelmann D, Coats M, Steinhauer D, Hardin J, Hyland iikuai K Hre E Wlf A Freiwald GA, Wolff E, Harper K, Kiriakoulakis coral deep-water in Fish (2004) MJ Kaiser O, Marchal TF, Stocker GK, Plattner F, Joos and national of review A (2002) KA Johnson rts Clmi. eerh Document Research Columbia. British cold coastal of areas explored in of corals water zoogeography and Taxonomy MD. 29pp National Marine Sanctuary Program, Silver Spring, and Administration, Oceanic Atmospheric National Commerce, Department of U.S. NMSP-06-11. Series Needs. Marine Sanctuaries Conservation Future and Status Marine Current – National Sanctuary Coast Olympic the at Marine California. Environmental Research37:195-229 Arguello, slope Point and shelf off continental outer on development the oil offshore Environmental of (1994) impact J Jeff R, Green Springer-Verlag BerlinHeidelberg (eds.), Cold-water corals and ecosystems. Pages 715-729 in Freiwald A, Roberts JM nutrition. their for implications and results initial North-East Atlantic: the from corals deep-water two of isotopes nitrogen and Lipids (2005) JM Roberts A, Grehan A, Science 304:1595 Editor. the to Letters habitats. atmospheric CO future on feedbacks cycle carbon marine Schmittner A (1999) Global warming and 57, 72pp NOAA NMFS-F/SPO- habitats. Memorandum benthic Technical effects on the fishing of on literature international 45pp 2006/062, Fisheries and Oceans Canada, 2 . Science128:464-467 STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Koenig CC, Shepard AN, Reed JK, Coleman Morgan LE, Chuendpagdee R (2003) Shifing FC, Brooke SD, Brusher J, Scanlon KM Gears: addressing the collateral impacts (2005) Habitat and fish populations in the of fishing methods in U.S. Waters. Pew deep-sea Oculina coral ecosystem of the Science Series, Washington DC, 42pp western Atlantic. In: Barnes PW, Thomas JP (eds.) Benthic habitats and the effects National Marine Fisheries Service (NMFS) of fishing. American Fisheries Society, (2007, in prep) Our living oceans. The Bethesda, Maryland, USA, pp795-806 status of habitats of U.S. living marine resources, 2006. U.S. Dep. Commer., Krieger KJ, Wing BL (2002) Megafaunal NOAA Technical Memorandum NMFS-F/ associations with deepwater corals SPO-75 (Primnoa sp.) in the Gulf of Alaska. Hydrobiologica 471:83-90 National Marine Sanctuary Program (NMSP) (2005) Biological and historical/cultural Kulm LD, Roush RC, Harlett JC, Neudeck resources of the study area for the Channel RH, Chambers DM, Runge EJ (1975) Islands National Marine Sanctuary Oregon continental shelf sedimentation: management plan update. In U.S. Dept.

Interrelationships of facies distribution of Commerce. National Oceanic and COAST PACIFIC and sedimentary processes. Geology Atmospheric Administration. National 83:145-175 Marine Sanctuary Program. Channel Islands National Marine Sanctuary Draft Lewis RC, Coale KH, Edwards BD, Nowicki- Management Plan / Draft Environmental Douglas J, Burton EJ (2002) Accumulation Impact Statement. Silver Spring, MD. rate and mixing of shelf sediments in the Monterey Bay National Marine Sanctuary. National Research Council (NRC) (2002) Effects Marine Geology 181:157-169 of trawling and dredging on seafloor habitat. National Academy Press, Love MS, Yoklavich MM, Black BA, Andrews AH Washington D.C. (2007) Age of black coral (Antipathes dendrochristos Opresko, 2005) colonies, Nevarez L, Molotch H, Shapiro P, Bergstrom R with notes on associated invertebrate (1998) Petroleum extraction in Santa species. Bulletin of Marine Science. Barbara County, California: an industrial 80:391-400 history. U.S. Dept. of Interior Minerals Management Service, OCS Study Love MS, Schroeder DM, Nishimoto MM MMS98-0048 (2003) The ecological role of oil and gas production platforms and natural Norse, EA (1993) Global marine biological outcrops on fishes in southern and central diversity. A strategy for building California: a synthesis of information. conservation into decision making. Island U.S. Dept. of Interior U.S. Geological Press, Washington DC, 383pp Survey, Biological Resources Division, Seattle, Washington, USA, OCS Study Northwest Fisheries Science Center (NWFSC) MMS 2003-032 (2007) Data report and summary analyses of open access fixed-gear fisheries in Minerals Management Service (MMS) (2006) waters less than 50 fathoms. Northwest Current facts and figures in the Pacific Fisheries Science Center, Seattle, OCS region as of May 1, 2006. U.S. Washington, USA Minerals Management Service, Online: http://www.mms.gov/omm/pacific/ Northwest Fisheries Science Center (NWFSC) offshore/currentfacts.htm (2006) West coast groundfish observer program data report and summary analyses of limited-entry trawl permits. Northwest Fisheries Science Center, Seattle, Washington, USA

147 148 PACIFIC COAST Pacific Fishery Management Council (PFMC) Council Management Fishery Pacific (PFMC) Council Management Fishery Pacific (PFMC) Council Management Fishery Pacific the of history Natural (1973) GL Ostarello Doney L, Bopp O, Aumont VJ, Fabry JC, Orr the of Revision (2002) DM Opresko the of Revision (2003) DM Opresko of species new A (2005) DM Opresko PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF at I Shzptia. Zoologische Anthozoa). Schizopathidae. II. (Cnidaria: Part Antipatharia Zoologische Anthozoa). Cladopathidae. Mededelingen (Leiden)77:495-536. III. (Cnidaria: Part Antipatharia bight. Zootaxa852:1-10 Antipatharia) from the southern California Anthozoa: (Cnidaria: coral antipatharian Pacific Fishery Management Council, Portland, Oregon, USA,756pp Management Fishery Pacific species. migratory highly for coast west US for statement and impact environmental plan management Fishery (2003) Council, Portland,Oregon,USA impact Management Fishery environmental Pacific statement. Final impacts: adverse of minimization and designation habitat fish essential plan: fishery management groundfish coast Pacific (2005) Oregon, USA Portland, Council, Management Fishery and Pacific impacts. adverse of designation minimization habitat plan: fish Essential management fishery groundfish (2006) Amendment 19 to the Pacific coast 564 145:548– Bulletin Biological The 1866). hydrocoral calcifying organisms.Nature437:81–86. on impact its and century twenty-first the over acidification ocean Anthropogenic (2005) A Yool Y, Yamanaka MF, Sarmiento Weirig IJ, Totterdell CL, RD, Slater R, Schlitzer Sabine JL, KB, GK, Rodgers Plattner RG, Najjar A, Mouchet P, Monfray R, Matear E, Lindsay Maier-Reimer RM, K, Key F, Joos A, Ishida N, Gruber A, Gnanadesikan RA, Feely SC, Mededelingen (Leiden)76:411-442. looa californic Allopora (Verrill, a ite L 20) aia-ae assessment Habitat-based (2005) JL Pirtle Pierce SD, Smith RL, Kosro PM, Barth JA, Wilson EK, Pikitch MA, Hixon DL, Stein WG, Pearcy the of landscape Foraging (2004) FA Parrish hlo GB (1980) GAB Shelton Taxonomic (1998) (eds.) JA Blake PV, Scott Milstein R, Robison C, Goldfinger CG, Romsos on deposition sand of Effects (1995) B Riegl (1997) PR Krause AM, Barnett PT, Raimondi nv, acue, ahntn USA, 64pp Washington, Vancouver, megafaunal Univ., State Washington Thesis, MS California. structure-forming invertebrates and fishes on Cordell Bank, of Sea ResearchII47:811–829 Deep- Pacific. North mid-latitude the of poleward the boundary eastern the along undercurrent of Continuity (2000) CD Bank. FisheryBulletin87:955-965 observations of deep-reef fishes of Heceta Submersible (1989) RM Starr WH, Barss Univ. Hawaii, Manoa,USA,146pp diss., Ph.D seal. monk Hawaiian ilgcl soito o te United the of Kingdom 60: pp517-528 Association Biological Marine the of Journal coral. deep-water a in behaviour and conduction electrical History, SantaBarbara,CA,USA Natural of Museum Barbara Santa the of Publications Special 3. Volume Channel. Barbara Santa western and Santa Basin Maria the of fauna benthic the of atlas of Canada,SpecialPaper47 Association Geological Characterization. and HG (eds) Greene Mapping Oregon the Seafloor BJ, for Habitat Todd In of USA. Washington, forthe margins map seafloor continental habitat regional geologic a surficial of Development RL, Chaytor JD, Wakefield WW (in press) corals. Marine Biology121:517-526. alcyonacean and scleractinian adults. and 16:1218-1228 larvae ToxicologyChemistry Environmental and marine on invertebrate muds drilling of effects The ohla pertusa Lophelia (L.): STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Smith CR, Hamilton SC (1983) Epibenthic Syms C, Jones GP (2001) Soft corals exert no megafauna of a bathyal basin off southern direct effects on coral reef assemblages. California: patterns of abundance, Oecologia 127:560-571 biomass, and dispersion. Deep-Sea Research 30:907-928 Thomas AC, Weatherbee RA (2006) Satellite- measured temporal variability of the Southern California Association of Marine Columbia River plume. Remote Sensing Invertebrate Taxonomists (SCAMIT) of Environment 100:167-178 (2001) A taxonomic listing of soft bottom macro- and megainvertebrates from Tissot BN, Yoklavich MM, Love MS, York K, Amend infaunal and epibenthic monitoring M (2006) Benthic invertebrates that programs in the Southern California form habitat on deep banks off southern Bight. Ed. 4. San Pedro, California, USA, California, with special reference to deep 192pp sea coral. Fishery Bulletin 104:167-181

Stein DL, Tissot BN, Hixon MA, Barss WH (1992) Tissot BN, Wakefield WW, Puniwai NPF, Pirtle J, Fish-habitat association on a deep reef at York K, Clemons JER (in prep) Abundance

the edge of the Oregon continental shelf. and distribution of structure-forming COAST PACIFIC Fishery Bulletin 90:540-551 megafaunal invertebrates, including deep corals, on Heceta Bank, Oregon, 2000- Steinhauer M, Imamura E (1990) California OCS 2002 phase II monitoring program: Year three annual report: Volume I. U.S. Dept. of Yoklavich MM, Love MS (2005) Christmas tree Interior Minerals Management Service, corals: A new species discovered off Pacific OCS Region, Los Angeles Southern California. Journal of Marine Education 21:27-30 Stone RP (2006) Coral habitat in the Aleutian Islands of Alaska: depth distribution, fine- Yoklavich MM, Greene HG, Cailliet GM, Sullivan scale species associations, and fisheries DE, Lea RN, Love MS (2000) Habitat interactions. Coral Reefs 25:229-238 associations of deep-water rockfishes in a submarine canyon: an example of a Stone RP, Masuda MM, Malecha PW (2005) natural refuge. Fishery Bulletin 98:625- Effects of bottom trawling on soft-sediment 641 epibenthic communities in the Gulf of Alaska. In: Barnes PW, James JP (eds.) Wakefield WW, Whitmire CE, Clemons JER, Benthic habitats and the effects of fishing. Tissot BN (2005) Fish habitat studies: American Fisheries Society Symposium combining high-resolution geological and 41, Bethesda, Maryland, USA, pp461- biological data, In: Barnes PW, Thomas 475 JP (eds) Benthic habitats and the effects of fishing: American Fisheries Society Strom NA (2006) Structure-forming benthic Symposium 41:119-138 invertebrates: habitat distributions on the continental margin of Oregon and Wheatcroft RA, Sommerfield CK, Drake DE, Washington. MS Thesis, Oregon State Borgeld JC, Nittrouer CA (1997) Rapid and Univ., Corvallis, Oregon, USA, 96pp widespread dispersal of flood sediment on the northern California margin. Geology Strub PT, James C (2000) Altimeter-derived 25:163-166 variability of surface velocities in the California Current System: 2. Seasonal Williams GC (1995) Living genera of sea circulation and eddy statistics. Deep-Sea pens (Coelenterata: Octocorallia: Research II 47:831-870 Pennatulacea): illustrated key and synopses. Zoological Journal of the Linnean Society 113:93-140

149 150 PACIFIC COAST STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF Appendix 3.1. Taxonomic list of known species of deep corals off the U.S. Pacific Coast, California to Washington. List includes both confirmed taxonomic records and some unconfirmed records of genera identified from surveys using bottom trawls and underwater vehicles.These unconfirmed records are denoted by the genus name followed by the word “species” (abbreviated as “sp”). Occurrence in two zoogeographic provinces (San Diego and Oregon) is noted. Known depth distributions are noted and originate from taxonomic records, published papers, and National Marine Fisheries Service (NMFS) bottom trawl surveys. Other abbreviations include the Olympic Coast (OCNMS) and Monterey Bay (MBNMS) National Marine Sanctuaries.

Higher Taxon Coral Taxa San Diego Oregon Depth (m) Source Phylum Cnidaria Class Anthozoa Subclass Hexacorallia Order Scleractinia Family Fungiacyathidae Fungiacyathus marenzelleri Vaughan, 1906 x x Cairns (1994) Family Micrabaciidae Leptopenus discus Moseley, 1881 x Cairns (1994) Family Rhizangiidae Astrangia haimei Verrill, 1866 x x 1-53 Cairns (1994) Family Oculinidae Oculina profunda Cairns, 1991 x 119 -578 Cairns (1994) Cairns (1994), Etnoyer and Morgan Madrepora oculata Linnaeus, 1758 x 84 (2003) Family Caryophylliidae Caryophyllia arnoldi Vaughan, 1900 x x 183 - 505 Cairns (1994) Labyrinthocyathus quaylei Durham, 1947 x x 37 - 293 Cairns (1994) Crispatotrochus foxi Durham and Barnard, 1952 x 82 Cairns (1994) Paracyathus stearnsii Verrill, 1869 x x Cairns (1994) Paracyathus montereyensis Durham, 1947 x 75 - 146 Cairns (1994) Coenocyathus bowersi Vaughan, 1906 x x 9 - 302 Cairns (1994), NMFS Nomlandia californica Durham and Barnard, 1952 x 82 Cairns (1994) Desmophyllum dianthus (Esper, 1794) x x Cairns (1994) Cairns (1994), Etnoyer and Morgan Lophelia pertusa (Linnaeus, 1758) x x (2003), OCNMS, unpublished data Family Flabellidae Javania californica Cairns, 1994 x 62 - 170 Cairns (1994) Polymyces montereyensis (Durham, 1947) x x 69 - 212 Cairns (1994), NMFS Family Dendrophylliidae Balanophyllia elegans Verrill, 1864 x x Cairns (1994), Pirtle (2005) Dendrophyllia oldroydae Oldroyd, 1924 x 99 - 366 Cairns (1994) STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION Source Tissot et al. (2006) Tissot NMFS, Etnoyer and Morgan (2003), MBNMS, unpublished data Opresko (2003) Opresko (2003), NMFS Opresko (2003), MBNMS, unpublished data NMFS, Etnoyer and Morgan (2003), MBNMS, unpublished data NMFS, MBNMS, unpublished data MBNMS, unpublished data MBNMS, unpublished data NMFS Austin (1985) MBNMS, unpublished data Austin (1985) MBNMS, unpublished data, OCNMS, unpublished data Montagne and Cadien (2001) Montagne and Cadien (2001) Montagne and Cadien (2001) Cairns et al. (2002), NMFS Cairns et al. (2002), NMFS, OCNMS Austin (1985) MBNMS, unpublished data NMFS, Etnoyer and Morgan (2003), MBNMS, unpublished data PACIFIC COAST PACIFIC 90 - 360 86 - 710 82 - 1162 Depth (m) 808 - 1165 296 - 1400 136 - 1243 531 - 1243 129 - 1200 1127 - 1159 1127 1357 - 2447 x x x x x x x x x x x x x x x x x x Oregon x x x x x x x x x x x x San Diego Opresko, 2003 Opresko, 2005 Bayer, 1958 Bayer, Opresko, 2003 Opresko, 2003 Broch, 1935 Verrill, 1868 Verrill, Verrill, 1868 Verrill, Coral Taxa Nutting, 1909 (Kukenthal, 1913) Verrill, 1868 Verrill, Verrill, 1868 Verrill, sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. Antipathes dendrochristos Antipathes Chrysopathes formosa Chrysopathes speciosa pseudotristicha Trissopathes Bathypathes Lillipathes Umbellapathes Acanthogorgia Calcigorgia spiculifera Anthothela pacifica Chrysogorgia Radiceps Corallium Adelogorgia phyllosclera Eugorgia rubens Heterogorgia tortuosa Leptogorgia caryi Leptogorgia chilensis Stenogorgia kofoidi Acanella Isidella Higher Taxon Order Antipatharia Order Antipathidae Family Family Cladopathidae Family Schizopathidae Subclass Octocorallia Order Gorgonacea Acanthogorgidae Family Anthothelidae Family Family Chrysogorgiidae Family Coralliidae Family Gorgoniidae Family Isididae

151 152 PACIFIC COAST STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF

Higher Taxon Coral Taxa San Diego Oregon Depth (m) Source Keratoisis flabellum Nutting, 1908 x Etnoyer and Morgan (2003) Keratoisis philippinensis Wright and Studer, 1910 x Etnoyer and Morgan (2003) NMFS, Etnoyer and Morgan (2003), Keratoisis sp. x x 516 - 1707 MBNMS, unpublished data Etnoyer and Morgan (2003), MBNMS, Lepidisis sp. x unpublished data Family Muriceidae Muricea californica Aurivillius, 1931 x Grigg (1970) Muricea fruticosa Verrill, 1869 x Grigg (1970) Ljubenkov (1986), Montagne and Thesea sp. A x Cadien (2001) Ljubenkov (1986), Montagne and Thesea sp. B x Cadien (2001) Etnoyer and Morgan (2003), MBNMS, unpublished data, OCNMS, unpublished Family Paragorgiidae Paragorgia arborea (Linnaeus, 1758) x 185 - 1743 data Paragorgia sp. x x Etnoyer and Morgan (2003) Family Plexauridae Swiftia spauldingi (Nutting, 1909) x x Austin (1985) Swiftia beringi Nutting, 1912 x OCNMS, unpublished data Swiftia kofoidi (Nutting, 1909) x x Austin (1985) Swiftia pacifica (Nutting, 1912) x 78 - 1186 NMFS, OCNMS, unpublished data Swiftia simplex (Nutting, 1909) x x 62 - 1075 Austin (1985), NMFS Swiftia torreyi (Nutting, 1909) x x Austin (1985) Family Primnoidae Amphilaphis sp. x 114 - 114 NMFS Callogorgia kinoshitae Kukenthal, 1913 x x 127 - 464 NMFS, Etnoyer and Morgan (2003) Calyptrophora sp. x Carey et al. (1990) Narella sp. cf. bowersi Nutting, 1908 x Carey et al. (1990) Parastenella doederleini Wright and Studer, 1889 x x Etnoyer and Morgan (2003) Parastenella sp. x Etnoyer and Morgan (2003) Plumarella longispina Kinoshita, 1908 x Etnoyer and Morgan (2003) Primnoa pacifica Kinoshita, 1907 x x OCNMS, unpublished data STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION Source Etnoyer and Morgan (2003) Cairns et al. (2002) NMFS Cairns et al. (2002) Austin (1985) Hochberg (1978), Montagne and Cadien (2001) Austin (1985) Montagne and Cadien (2001) Austin (1985) Montagne and Cadien (2001) Berntson et al. (2001) Hochberg and Ljubenkov (1998) Austin (1985) Austin (1985) Hochberg and Ljubenkov (1998), NMFS NMFS Hochberg and Ljubenkov (1998) Hochberg and Ljubenkov (1998) Austin (1985) Montagne and Cadien (2001) Austin (1985) Austin (1985) Hochberg and Ljubenkov (1998) Hochberg and Ljubenkov (1998) PACIFIC COAST PACIFIC intertidal 293 - 972 Depth (m) 103 - 1543 516 - 1083 x x x x x x x x x x x x x x x x x x Oregon x x x x x x x x x x x San Diego Nutting, 1912 Pasternak, 1960 (Verrill, 1879) (Verrill, Studer, 1894 Studer, Studer, 1894 Studer, Williams, 2000 Verrill, 1882 Verrill, Coral Taxa Nutting, 1912 (Ehrenberg, 1834) Moroff, 1902 Moroff, Kolliker, 1880 Kolliker, Nutting, 1909 Nutting, 1909 Kukenthal, 1913 Kolliker, 1874 Kolliker, (Pallas, 1766) Pfeffer, 1886 Pfeffer, Verseveldt and van Ofwegen, 1992 Verseveldt sp. sp. sp. A A sp. sp. H sp. Thouarella Alcyonium rudyi Anthomastus ritteri Cryptophyton goddarti Clavularia Clavularia ambigua Telestula Telesto Gersemia rubiformis Renilla koellikeri Renilla reniformis Kophobelemnon affine Kophobelemnon biflorum Kophobelemnon hispidum Anthoptilum grandiflorum Anthoptilum murrayi Funiculina parkeri Distichoptilum gracile Helicoptilum rigidum Stachyptilum superbum Scleroptilum Umbellula lindahli Umbellula magniflora Halipteris californica Higher Taxon Order Alcyonacea Order Alcyoniidae Family Family Clavulariidae Family Nephtheidae Order Pennatulacea Suborder Sessiliflorae Family Renillidae Family Kophobelemnidae Anthoptilidae Family Family Funiculinidae Family Protoptilidae Family Stachyptilidae Family Scleroptilidae Family Umbellulidae Family Halipteridae

153 154 PACIFIC COAST STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION THE IN ECOSYSTEMS STATE CORAL DEEP OF

Higher Taxon Coral Taxa San Diego Oregon Depth (m) Source Halipteris sp. x Suborder Subselliflorae Family Virgularidae Acanthoptilum album Nutting, 1909 x x Hochberg and Ljubenkov (1998) Acanthoptilum gracile (Gabb, 1863) x Hochberg and Ljubenkov (1998) Stylatula elongata (Gabb, 1862) x x Hochberg and Ljubenkov (1998) Stylatula gracilis Verrill, 1864 x x 64 - 1243 Hochberg and Ljubenkov (1998), NMFS Ljubenkov (1991), Montagne and Stylatula sp. A x Cadien (2001) Virgularia agassizii Studer, 1894 x Montagne and Cadien (2001) Virgularia californica Pfeffer, 1886 x Montagne and Cadien (2001) Virgularia cystiferum (Nutting, 1909) x x Austin (1985) Virgularia sp. cf glacialis Kolliker, 1870 x Austin (1985) Family Pennatulidae Pennatula californica Kukenthal, 1913 x x 465 - 991 Hochberg and Ljubenkov (1998), NMFS Ptilosarcus gurneyi (Gray, 1860) x x 11 - 922 Hochberg and Ljubenkov (1998), NMFS Ptilosarcus undulatus Verrill, 1865 x Cairns et al. (2002) Class Hydrozoa Order Anthoathecatae Suborder Filifera Fisher (1938), Alberto Lindner, pers. Family Stylasteridae Errinopora pourtalesii (Dall, 1884) x 49-183 comm. Fisher (1938), Alberto Lindner, pers. Stylantheca petrograpta (Fisher, 1938) x intertidal comm. Fisher (1938), Alberto Lindner, pers. Stylantheca porphyra Fisher, 1931 x intertidal comm., Etnoyer and Morgan (2003) Fisher (1938), Alberto Lindner, pers. Stylaster californicus (Verrill, 1866) x x 35-90 comm., Etnoyer and Morgan (2003) Fisher (1938), Alberto Lindner, pers. Stylaster venustus (Verrill, 1868) x 49-84 comm., Etnoyer and Morgan (2003)